Vol. 31, No.3 & 4 Fall/Winter 1998 THE GREAT LAKES ENTOMOLOGIST

PUBLISHED BY

THE MICHIGAN ENTOMOLOGICAL SOCIETY THE GREAT lAKES ENTOMOLOGIST

Published by the Michigan Entomological Sociely

Volume 31 No.3 & 4

ISSN 0090-0222

TABLE OF CONTENTS

A new species of Eularsopolipus [Aoori: PodapolipidaeJ from Harpo/us pennsylvanicus (Coleoptera: Carabidae) from East Lansing, Michigan Robert W, Husband ..... , . , ...... , , . , ...... , .... , . , . , 141

Evidence of long range transport of a dark morph swallowtail butterfly (Papilia g/aucus) on a storm front into northern Michigan (Lepidoptera: Papilionidae) J. Mark Scriber, Mark D. Deering and Aram D, Stump, .. , . , ..... , ... , ...... 151

Effects of enamel point on the behavior and survival of the periodical cicada, Magidcada sepfendecim (Homoptera) and the lesser migratory grasshopper, Me/aoop/us saoguiojpes (Orthoptero). John R. Cooley, G.S. Hammond and D,C. Marshall .... , . , .... , .. , . , ... , .. , .... , 161

The parasitoid complex of first generation Ostrinja nubi/a/is (Lepidoptera: Pyralidae) larvae in northwest Ohio Daniel M. Pavuk and Laura l. Hughes .. , ...... , , . , , .... , . , , . , , .... 169

Temperature effects on development in Aphe/inus a/bipodus [Hymenoptera: Aphelinidae) from two geogrophic regions Jang-Hoan Lee and Norman C. Elliott ...... 173

Phenology and infestation patterns of the cottonwood twig borer (Lepidoptera: Tortricidae) in Iowa Joel D. McMillin, Michael J. Anderson, Elizabeth E. Butin and Elwood R. Hart ...... 181

The sequential relationship of body oscillations in the paper wasp, Polis/es Fuscatus (Hymenoptera: Vespidae) Bobbi J. Harding & J. Gamboa ...... 191

Additional Siphonaptera records from small mammals in the central Upper Peninsula of Michigan William C. Scharf and Patrick E. Lederle ...... 195

Psi/olrela indedsa and Agarodes dislinclus [Trichoptera: Odonloceridae, Sericostomalidae): New state records and notes on the habitat of these species in Michigan Ethan Bright and Doug Bidlack ...... 199

New larval variants and distributional records for Plauditus ceslus (Ephemeroptera: Baetidae) C. R. lugo-Ortiz and W. P. McCafferly...... _...... 201

Distribution of He/aerina tilia (Odonata: Calopterygidae) in the eastern Great lakes Region Paul D. Pratt and Paul M. Calling ...... 203

Gleaning on Coreidae [Heteropteraj byTachop/eryx /horeyi (Odonata: PetaluridaeJ Robert D. Waltz ...... 209

Enallagma anno, a damseilly new to Ihe Great Lakes region (Odonata: Coenagrionidae) Mark F. O'Brien ond Poul D. Pratt ...... 211

COVER PHOTO Stream Mayfly [Ephemeroplera: Heptageniidae), Minnesota. Photo by David C. Houghton. THE MICHIGAN ENTOMOLOGICAL SOCIETY

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Copyright © 1998. The Michigan Entomological Society 1998 THE GREAT LAKES ENTOMOLOGIST 141

A NEW SPECIES OF EUTARSOPOLIPUS (ACARI: PODAPOLIPIDAE) FROM HARPALUS PENNSYLVANICUS (COLEOPTERA: CARABIDAE) FROM EAST LANSING, MICHIGAN

Robert W. Husband]

ABSTRACT Eutarsopolipus /ischeri n.sp. (Acari: Podapolipidae) is described from the carabid beetle, Harpalus pennsylvanicus (DeGeer) and compared with the other species in the pterostichi group of Eutarsopolipus. Keys to groups of Eutarsopoiipus and to species in thepterostichi group are provided.

Mites in the family Podapolipidae (Acari: Heterostigmata) are all para­ sites of insects. The genus Eutarsopolipus is restricted to Carabidae (Coleoptera) and occurs worldwide. The genus was erected for E. lagenae­ fdrmis Berlese 1913. More than 30 species have been discovered and most of them are described by Regenfuss (1968, 1974). Previous species recorded from Michigan carabid beetles include E. regenfussi Husband and Swihart 1984, E. porten Husband 1993 and E. crassisetus Regenfuss 1968. A closely related species in the pterostichi group is E. inermis Regenfuss 1974, col­ lected in Georgia, U. S. A. The purpose of this paper is to describe Eutarsopolipus fischeri, new species, collected from Harpalus pennsylvanicus (DeGeer), compare E. /is­ cheri with species in the pterostichi group of Eutarsopolipus as defined by Regenfuss 1968 and provide a key to distinguish the pterostichi group from 7 other groups of Eutarsopolipus. Species in the pterostichi group are: Eutar­ sopolipus pterostichi Regenfuss 1968 from Germany and Ukraine, E. vernalis Regenfuss 1968 from Germany, E. inermis Regenfuss 1974 from Georgia, U.S.A. and E. diunculosus Eidelberg 1994 from Ukraine. Measurements were taken with the aid of a Zeiss microscope with a stage micrometer and drawing tube. All measurements are in micrometers (}lm). Terminology is based on Lindquist (1986).

Eutarsopolipus fischeri Husband, new species Female (Figs. 1,2)- Gnathosoma length 45-48 width 42-46. Palp length 10-17; cheliceral stylets 33-34, dorsal gnathosomal setae 17-19, ventral setae 5-6. Stigmata not evident. Idiosoma- Length 332-450, width 240-350. Prodorsal plate setae vI 5-6 v 6-7, sc 32-42, setae situated near the posterior margin of prodor sal plate. Distance between setae VI 33-38; v2 slightly lateral to a line connecting vI

IBiology Department, Adrian College, Adrian, MI 49221. 142 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4

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Figure 1. Eutarsopolipus fischeri n. sp., adult female, dorsal aspect. 1998 THE GREAT LAKES ENTOMOLOGIST 143

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2

Figure 2. Eutarsopolipus fischeri n. sp., adult female, ventral aspect. 144 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4

Table 1. Leg setation for femur, genu, tibia, tarsus for E. fischeri and other species in the pterostichi group. Solenidia are included. Leg! Leg II Leg III F G Ti Ta F G Ti Ta F G Ti Ta E. pterostichi 3 2 6 8 0 0 4 7 0 0 4 6 E. vernalis 3 2 6 8 0 0 4 7 0 0 4 6 E. inermis 2 0 6 8 0 0 4 6 0 0 4 5 E. diunculosus 3 0 5 9 0 0 4 6 0 0 4 4 E. fischeri 2 0 7 8 0 0 4 6 0 0 4 6

SC • and 2 Plate C length 50-58, width 210, setae c1 3-5, ~ 5-6. Plate D length 47-65, width 178-188, setae d 3-5. Plate EF length 4·,-57, width 130-150, setae (5. Venter with apodemes 1,2 well developed, meeting sternal apodeme me­ dially; sternal apodeme not extending beyond junction with apodemes 2. Coxal setae 1a 2, 2a 3; setae 1a situated equidistant to apodemes 1, 2 and sternal apodeme. Distance between setae 1a 22-34, 3a and 3b 18-23. Coxal setae 3a 4-5, 3b 7. Legs- Leg setation as in Table 1. Ambulacrum I with a terminal stout claw, ambulacra II, III with 2 strong claws. Tarsus I subunguinal seta spine­ like, 2 terminal spines on each of tarsi II, III. Tarsus I solenidion Cl) 3; tarsus II solenidion Cl) not evident. Tibia I solenidion cI> 8-10, 112 to 213 width of base of tarsus I. Tibial I, II, III setae d 22-30,7-10 and 6-8 respectively. Male (Figs. 3,4)- Gnathosoma length 26-29, width 27. Palps 13; che­ liceral stylets 19-22; dorsal gnathosomal setae 2-4, ventral setae 2. Idiosoma- Length 161-162, width 104-112. Prodorsal setae VI 2, v2 2, setae sc2 48. Distance between setae v1 25-26, distance between setae ~ 42-43; setae v2 lateral to a line connectmg setae VI and SC 2• Plates C and V fused, setae cl> v2 2, setae d, and (obscured. Genital capsule length 25-38, width 27-30. Venter- Apodemes 1,2 and sternal apodeme conspicuous; coxae III sepa­ rated medially. Setae la, 2a, and 3a vestigial, 3b obscured. Legs- Leg setation as in Table 1. Ambulacrum with 1 straight claw, am­ bulacra II, III with minute claws. Single tarsal I ventroterminal spine, 2 ter­ minal spines on each of tarsi II, III. Tarsus I solenidion Cl) 3, tarsus II solen­ dion Cl) 2-3; tibia I solendion cI> 7-8, about 3/4 width of base of tarsus I. Larva (Fig. 5)- Gnathosoma length 31, width 31. Palps 10, cheliceral stylets 25, dorsal setae 17, ventral setae 3. Idiosoma- Length 200, width 100. Prodorsal plate bluntly rectangular, setae VI 5, v2 3, sC 2 74. Distance between setae v2 greater than the distance ; VI SC . between setae vl setae v2 lateral to a line connecting setae and setae 2 Plates C,D fused anteromedially, setae c 3, c 4, setae d 4: distance between . setae d 34. Setae c2, anterior to setae ci Plate EF oval, setae {4. Plate H oval, setae hI 43, h2 16, distance between setae hI 7. Venter- A:podemes 1,2 and sternal apodeme conspicuous but weakly scerotized. Setae 1a m, 2a m, 3a 3 and 3b 4. Distance between setae 3a and 3b 14. Legs- Leg setation as in Table 1. Ambulacrum I with 2 slender claws. Ambulacra II, III with minute claws. Single tarsus I spine short and stout 5, 1998 THE GREAT LAKES ENTOMOLOGIST 145

'£\vo terminal tarsi II, III spines 5-7. Tarsus I solenidia ro 3 tarsus II solenid­ ion ro 2. Tibia I solenidion Ii> 6, seta k 3. Setae tc' 8, tc"ll. Eggs- Length 192-217, width 100-119. Type data- Holotype female: from East Lansing, Michigan, U.S.A. from under elytra of Harpalus pennsylvanicus (DeGeer) (Carabidae) collected by Roland L. Fischer, 26 July 1947 (RWH 22798-1). Allotype (RWH 22798-2) with same data as holotype and both deposited in the Museum of Entomol­ ogy, Michigan State University, East Lansing, MI, U.S.A. Paratypes: 2 fe­ males without larval exoskeletons, 4 females with partial exoskeletons oflar­ vae, 4 males and 2 larvae on slides and females, males and eggs with the same data as the holotype stored in 70% ethanol. One female and one male depsoited in the collection to the Museum of Zoology, University of Michigan. Balance of paratypes in the collection of the author. Diagnosis- Eutarsopolipus fischeri and E. inennis lack ventral femur I setae in all stages. Larval female and male E. fischeri have minute but well sclerotized ambulacral II, II claws. Four related species in Europe and the United States lack claws in male and larval stages. Female E. fischeri have ambulacral I claws while female E. inermis lack ambulacral I claws. Length of cheliceral stylets in female E. fischeri are 34 versus 48 in E inermis. Che­ liceral stylets of male E. fischeri are 19 versus 27 in E. inennis. Larval E. fis­ cheri have cheliceral stylets, 25, versus 49 in E. inennis. Dorsal gnathosomal setae are 17 in E. fischeri versus 30 in E. inennis. Setae hz are 16 in E. fis­ cheri versus 51 in E. inermis. Regenfuss (1968) proposed 7 groups of Eutarsopolipus based on 11 female characters, 2 larval female characters and 1 male character. The Ochoai group was added by Husband (1995). The following key uses characters of only adult females for simplicity. Some species do not have all of the charac­ ters of the group as noted by Regenfuss (1974). However, all of the species will fit in groups listed in the keys below.

Key To Groups ofEutarsopoljpus Based onAdult Females L Without leg II, III claws ...... 2 With leg II, III claws ...... 4 2. Stigmata and trachea evident, plates, C, D present ...... 3 Stigmata and trachea not evident, plates C, D not present .... .8tammeri 3. Strong claw on leg I, with genu III seta, femur I l' long (15 11m or longer) . Acanthomus No strong claw on leg I, without genu III seta, femur I l' short (1-5 11m ) .. Biunguis 4. Without genu III seta ...... 5 With genu III seta ...... ochoai 5. Femur I seta l' short, less than 10 \lm ...... 6 Femur I seta I'10ng, 11-21 J.l.ID •••••••.•••.••.•••••..••••.•• •Myzus 6. Stigmata and trachea prominent ...... 7 Stigmata and trachea not prominent ...... Pterostichi 7. Setae h present, cheliceral stylets shorter than 75 11m ...... Desani Setae h not present, stylets longer than 140 11m ...... Lagenae/ormis

Key To Species in the Pterostichi Group Based onAdult Females L Femur I v" seta short but evident ...... 3 Femur I v" seta not present ...... 2 146 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No, 3 &4

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Figure 3. Eutarsopolipus fischeri n. sp., male, dorsal aspect. 1998 THE GREAT LAKES ENTOMOLOGIST 147

4

Figure 4. Eutarsopolipus fischeri n. sp., male, ventral aspect. 148 THE GREAT LAKES ENTOMOLOGIST Vol. 31/ No. 3 & 4

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5

Figure 5. Eutarsopolipus fischeri n. sp., larval female, dorsal aspect. 1998 THE GREAT LAKES ENTOMOlOGIST 149

2. No claw on leg I...... Eutarsopolipus inermis Regenfuss Prominent claw on I ...... Eutarsopolipus fischeri n. sp. 3. Leg I with I claw ...... 4 Leg I with two claws...... Eutarsopolipus diunculosus Eidelberg 4. Setae v2 on or medial to a line connecting setae ul and sc2 •••••••••••••• ...... Eutarsopohpus pterostichi Regenfuss Setae v2 disticntly lateral to a line commecting setae VI and sc,2' ...... Eutarsopolipus uernallS Regenfuss

DISCUSSION Synapomorphies shared by species in the pterostichi group are no stig­ mata, no setae on genua II and III, males and larvae with either no ambu­ lacral II and III claws or much reduced claws, and femur I seta v" short or absent. Other characters shared include genital plates which are about as long as wide, females with leg II, III claws (except E. inermis), strong claw on leg I (except E. inermis), and setae VI and u2 evident (Regenfuss, 1968, 1974). American species and E. diunculosus laCK setae on genu I. Eidelberg (1994) noted ambulacrum I with 2 claws in female E. diunculosus instead of 1 claw as in all other adult female Eutarsopolipus. As more species are discovered, it will be interesting to note whether the lack of ventral femur I setae and reduced or absent claws will continue to be characteristic of species in the pterostichi group of American versus European Eutar­ sopolipus.

ACKNOWLEDGMENTS The assistance of Fred Stehr, Director of the Museum of Entomology, Michigan State University, in acquiring specimens and advice on host beetles and procedure is appreciated.

LITERATURE CITED Berlese, A. 1913. Acari NuovL Redia 9:27-87. Eidelberg, M. 1994. Mites of the family Podapolipidae (Heterostigmata: Tarsonemina) of Ukraine and adjacent areas with description of a new species. Verstin. Zool. I: 37--43. (in Russian) Husband, R. W. 1993. A new Eutarsopolipus sp. (Acari: Podapolipidae) from Harpalus herbivagus Say (Coleoptera: Carabidae) from Michigan. Great Lakes Entomol. 26(2): 1-13. Husband, R. W. 1995. A new species of Eutarsopolipus (Acari: Podapolipidae) from Costa Rican Pasimachus spp. (Coleoptera: Carabidae). Entomol. Mitt. zool. Mus. Hamburg Bd. 11(151): 157-165. Husband, Robert W. and Cheryl D. Swiliart. 1986. A new species of mite (Acari: Po­ dapolipidae) from a Michigan carabid beetle, Chlaenius pennsylvanicus. Great Lakes Entomol. 19: 107-113. Lindquist, E. E. 1986. the world genera Tarsonemidae (Acari: Heterostigmata): a mor­ phological, phylogenetic, and systematic revision with reclassification of family group taxa in Heterostigmata. Mem. Ent. Soc. Canada 136:1-517. 150 THE GREAT lAKES ENTOMOLOGIST Vol. 31, No.3 & 4

Regeniuss, H. 1968. Untersuchungen zur Morphologie, Systematik und Okolgie der PodapoJipidae (Acarina, Tarsonemini). Z. wiss. Z001. 177(3/4): 183-282. Leipzig. Regenfuss, H. 1974. Neue ektoparasitische Axten der familie Podapolipidae (Acari: Tarsoneminil von Carabiden. Mitt. Hamburg. Zoo!. Mus. lnst. 71: 147-163. Ham­ burg. 1998 THE GREAT LAKES ENTOMOLOGIST 151

EVIDENCE OF LONG RANGE TRANSPORT OF A DARK MORPH SWAl­ LOWTAil BUTIERFLY (PAPILIO GLAUCUSj ON A STORM FRONT INTO NORTHERN MICHIGAN (lEPIDOPTERA: PAPILIONIDAE)

J. Mark Scriber 1, Mark D. Deering 1, and Aram D. Stump1

ABSTRACT

A dark morph female Papilio glaucu8 was captured in the middle of a P. canadensis population in the center of Dickinson County, in Michigan's Upper Peninsula. We are unaware of any dark female ever being captured at this latitude, and none within 400 km has ever before been reported. Mor­ phological wing traits all score this dark female as P. glaucus, with all 31 of the other (yellow) females captured from the same group of nectar sources scoring as P. canadensis. Two diagnostic electrophoretic allozymes (PGD and LDH) show all 24 males to have 100% P. canadensis alleles, and 98% canadensis alleles were seen for the HK locus. The dark female had the P. glaucus X-linked allele for PGD, the P. canadensis X-linked allele for LDH, and was heterozygous for diagnostic HK alleles which are autosomaL All other yellow females that were examined were pure canadensis type for the sex-linked PGD and LDH alleles. One yellow female was heterozygous for the autosomal HK diagnostic alleles. A long distance "blow-in" on a strong weather front from the southwest is suspected as the most likely explanation for the appearance of this dark morph female of mixed (introgressed) genetic background so far from its nearest known source. The blow-in was not likely to have been in a previous year since this dark morph female would require both (1) a dark morph mother with the Y-linked dark gene and (2) a father that was part P. glaucus (i.e. without the color suppressor genes and with a PGD-100 allele), None of the males from this area had PGD or LDH alleles of the canadensis type this year or ever previously, making it unlikely that the blow-in occurred in a previous year.

On 25 June 1997, the day after a strong storm with tornado watches blew across Wisconsin and Michigan from the southwest, we captured a dark morph tiger swallowtail (Papilio glaucus) female in Dickinson County, of the Upper Peninsula of Michigan. This female, captured nectaring on clover at 11:25 hr, was the largest of 32 females collected at that population site. All other 31 females were yellow and of the P. canadensis type, based on key morphometric wing band distinguishing characters (Table 1; and Luebke et al. 1988; Scriber 1990). Dark female morphs are believed to occur only in P. glaucus and not in P. canadensis (Scriber et al. 1996), and have never before been reported anywhere for the Upper Peninsula of Michigan. This observa-

IDepartment of Entomology, Michigan State University, East Lansing, MI 48824 152 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4

Table 1. Wing characteristics of 31 females captured on 25 June 1997 in Dickinson County, Michigan with the dark morph female of P. glaucus (see Scriber 1982; Luebke et al. 1988). Proportional width % black Female Forewing Submarginal band anal cell Mean colorl langth mm2 yellowFW2 HW3 class age4 1 Yellow 47 band 65 1.0 2 Yellow 47 band 65 4.0 3 Yellow 49 band 60 1.5 4 Yellow 48 band 60 2.0 5 Yellow 49 band 70 1.0 6 Yellow 47 band 70 1.0 7 Yellow 49 band 85 2.5 8 Yellow 47 band 75 1.5 9 Yellow 48 band 55 1.5 10 Yellow 48 band 75 1.5 11 Yellow 48 band 65 2.0 12 Yellow 51 band 55 1.5 13 Yellow 46 band 55 2.5 14 Yellow 47 band 50 1.5 15 Yellow 52 band 55 1.0 16 Yellow 48 band 80 1.0 17 Yellow 48 band 75 2.0 18 Yellow 46 band 55 2.0 19 Yellow 45 band 80 2.0 20 Yellow 51 band 55 1.5 21 Yellow 48 band 70 1.0 22 Yellow 48 band 70 2.0 23 Yellow 49 band 85 2.0 24 Yellow 49 band 55 2.0 25 Yellow 46 band 50 2.5 26 Yellow 47 band 70 1.0 27 Yellow 48 band 60 2.0 28 Yellow 48 band 65 2.0 29 Yellow 47 band 80 2.0 30 Yellow 48 band 70 2.5 31 Yellow 47 band 65 2.5 32 dark 52 spots 30 2.0 1 See Scriber et al1996 (for discussion and genetics of dark morph distribution). 2See Luebke et al1988 (size can vary according to foodplant reared upon, but has a ge­ netic basis). aSee Scriber, 1982 (P. canadensis usually greatly exceed 50% in band width; P. glaucus are usually less than 40%). 4See Lederhouse 1978 (1=perfect, fresh, 4=very worn). tion is 320-400 km further north than any dark morph female of which we are aware (Fig. 1, Scriber 1996a). From 1978 to 1997, we have previously col­ lected more than 1,100 females of P. canadensis in northern Michigan and Wisconsin, and more than 1,200 females of P. glaucus in southern parts of these states and adjacent Illinois and Ohio (Scriber 1994, and unpublished), While dark morph females were reported from Toronto, Canada (Phil 1998 THE GREAT lAKES ENTOMOLOGIST 153

Figure 1. Distribution records more than 4,400 dark morph female Papilio glaucus from numerous Museum records and personal collections (modified from Scriber 1996a). The percentage of dark morph females at the northern tier of counties is only 5% Wisconsin, 6% Michigan, and 3% in Massachusetts (Scriber et al 1996). The closest previous county record of any dark morph in Wisconsin is 150-250 miles from our Dickinson County capture site (inci­ dated with a star symbol in the upper peninsula of Michigan).

Schappert, pers. comm.) and Midland, Michigan in the early 1990s (after sev­ eral of the warmest years on record for the growing season degree-day ther­ mal accumulations), the northward movement of P. glaucus genes across the Great Lakes hybrid zone has apparently been minimal and only in the range of 50-100 miles (Scriber and Gage 1995). Nonetheless, the possibility of a relic glaucus-type gene pool persisting in this small pocket of several hun­ dred miles out of the geographic range of the species and 400 km from the northern limits of the species' distribution could not be ruled out entirely. 154 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No.3 &4

.it• •• ""1""----- •• (T)(l) 00

PROPO;1 nON 01" O~ ALlELES: C"'NAOENS!$ TY?E tlH ~dh ;>gd Hk

Figure 2. Electorphoretic allozyme frequencies (males only) for LDH, PGD, and HK across the hybrid (transition) zone between P. canadensis in the north and P. glaucus in the south (modified from Scriber 1996). The shaded region on the map corresponds to the average seasonal total degree dayaccu­ mulations (1400-1500 degrees C, base 10) that delineate the northernmost limits to bivoltine populations of P. glaucus, as well as the previous northern­ most limits to dark morph females (see Fig. 1). The Dickinson County dark morph location is indicated with a star (1997 allozyme frequencies of males are presented in Table 2).

Our analyses here consisted of electrophoretic and morphometric compar­ isons and consultation of weather records to examine the likelihood of relic vs. vagrant P. glaucus phenotype and genes.

METHODS Morphological characters used as potentially diagnostic traits between P. glaucus and P. canadensis include the wing lengths, submarginal yellow band on the ventral forewings, and the anal cell black band width (Scriber 1982, Luebke et al. 1988, Scriber 1990). Biochemical allozyme electrophore­ sis differences include hexose kinase (HK), lactate dehydrogenase (LDH), and phosphogluconate dehydrogenase (PGD). Standard techniques were used for enzyme analysis (Hagen and Scriber 1991). Weather data was obtained from the National Weather Service midwest regional data center. 1998 THE GREAT LAKES ENTOMOlOGIST 155

Table 2. Wing characteristics and allozyme alleles of 24 male P. canadensis captured in the immediate vicinity of the females (Table 1). Proportional Alleles** (Allozymes) Forewing Submarginal width % black mm2 yellowFW2 band anal cell HW LDH PGD 1 46 band 50 110 80 90 2 46 band 70 110 80 90 3 44 band 70 110 80 90 4 44 band 70 110 80 90 5 45 band 80 110 80 90 6 46 band 60 110 80 90 7 46 *band 60 110 80 90 8 46 band 50 110 80 90 9 42 band 60 110 80 90 10 45 band 70 1101100 80 90 11 44 band 60 110 80 90 12 44 band 80 110 80 **901110 13 45 band 60 110 80 90 14 43 band 70 110 80 90 15 44 band 50 110 80 90 16 43 band 60 110 80 90 17 42 band 80 110 80 90 18 42 band 40 110 80 90 19 44 *band 45 110 80 90 20 44 band 60 110 80 90 21 44 *band 60 110 80 90 22 42 band 70 110 80 **90/110 23 42 band 85 110 80 **90/110 24 47 band 70 110 80 90 *FW submarginal showing some "intermediate" dimpling (see Luebke et al. 1988). **Both alleles PGD 90 and PGD 110 are P. canadensis (see Hagen and Scriber 1989 and Fig. 2).

RESULTS The 31 yellow females (Table 1), and all of the 24 male swallowtail but­ terflies we collected in that same area were clearly of the P. canadensis type with regard to key morphological wing characters (Luebke et al. 1988). Hybrid individuals are not always easily identified by a few key wing characters, and we brought the specimens from this population back to Michigan State University to examine for biochemical (allozyme) traits which we have determined to be excellent species diagnostic tools (Fig. 2, Hagen et al. 1991; Scriber 1996a). The results of these analyses showed the 24 males from this population to have 100% of the alleles (PGD-90 and 110 and LDH-80) to be P. canadensis type and 97.9% of the HK alleles to be P. canadensis type (HK-110; Table 2). One of the males was heterozygous (HK 110/100). With females, only 18 were able to give excellent allozyme bands for all 3 diagnostic enzymes. The others appear to have lost some stain intensity, per­ haps due to the fact that they weakened or died in their oviposition arenas and even with immediate freezing, some enzyme activity was apparently lost 156 THE GREAT lAKES ENTOMOLOGIST Vol. 31, No.3 & 4

for electrophoretic detection. Everyone of the 19 females with LDH stains was the P. canadensis type (even the dark morph)! It is interesting that only the dark morph female exhibited the PGD-100 (P. glaucus-type) allele of the 20 that exhibited PGD stain. However nearly all the females had canadensis alleles, except this dark female and one yellow one that was heterozygous 110/100 for the HK locus of 22 females showing the HK stain. This suggests a complicated genetic scenario for this dark morph female. If the LDH and PGD alleles are sex-linked and fixed (diagnostic) for the two swallowtail species (Hagen and Scriber 1989, 1991; Hagen et al. 1991), then the father of the dark morph female apparently passed an X-chromosome with both P. canadensis and P. glaucus alleles (suggesting a previous crossover event). This type of male crossover event was once before suggested in a male from a South Manitou Island population from the northern end of Lake Michigan (Scriber 1994). The father of our Dickinson County dark female could also have been P. glaucus-like in the HK-100 autosomal allele he contributed.

DISCUSSION In summary, we have found for this Dickinson County population some evidence of a low level of genetic introgression from P. glaucus in 1 male het­ erozygous for the autosomal HK alleles, and evidence of genetic introgression at the HK locus of 2 heterozygous females including the dark morph, which had one additional glaucus allele (PGD). These results are unlike all previ­ ous data from the Upper Peninsula in which diagnostic alleles thus far ana­ lyzed were clearly P. canadensis, without heterozygotes (Scriber 1996a, Scriber et al. 1996, and Fig. 2). If a pocket of genetically glaucus-type individuals existed in this Dickin­ son County, Michigan population, we would have expected some morphomet­ ric traits to clearly be evident in both males and females, but they were not. Furthermore, unless the nationwide pattern of dark/yellow morph female in­ heritance is inaccurately understood, we should see a dark morph female only if 1) her mother was also dark (since it is a Y-linked trait) and 2) her fa­ ther was homozygous or heterozygous for the "enabler" gene, or for lacking the "suppressor" gene on the X-chromosome (Scriber et al. 1996). In other words, this dark female required both a dark morph mother and a father that was partly P. glaucus, both of which are highly unlikely based on the total lack of dark morphs ever seen in the Upper Peninsula and the large number of morphologicallbiochemical analyses of the males that were P. canadensis-like in this population. The uncommon and unexpected appearance of dark female daughters from yellow P. glaucus mothers can be explained fairly easily, and in most cases it was likely that the yellow mother was carrying the dark allele on her Y-chromosome (Clarke and Sheppard 1962) with a "suppressor" (or lack of "enabler") on her X-chromosome (Scriber et al. 1996). These cases historically are almost always from parents close to the Great Lakes hybrid zone where genetic introgression is most likely (250-300 miles from our dark morph Dickinson Co. female). In contrast, while the appearance of any dark morph daughters from a P. canadensis mother has never been reported from nature, it has occurred in one interspecific hybrid brood from our lab studies (Scriber and Evans 1986). In this case a yellow P. canadensis female from northern Wisconsin produced 21 dark and 12 yellow (213 dark) daughters (male mate was an aberrant colored P. glaucus from Ohio). In this exceptional brood (#1132; Scriber and Evans 1986) it could not be resolved as to whether the rare dark daughters were the result of a cross-over event from the Y-chromo­ 1998 THE GREAT LAKES ENTOMOlOGIST 157

some or due to a nondisjunction of the sex chromosome (Clarke et al. 1976, and C.A. Clarke, pers. comm.). While we now have evidence that crossovers on the X-chromosome of male P. canadensislglaucus hybrids have occurred in nature (Scriber 1994), and that introgression from canadensis may explain "spring form" glaucus that look like canadensis (Scriber 1990), the cause of the occurrence of dark females from a canadensis mother has never been re­ solved. Therefore, this female dark morph found in Dickinson County could not automatically have been assumed to be a "pure" P. glaucus individual. In fact, we have observed that this dark female has the P. canadensis sex-linked allele (LDH-80) as well as being heterozygous for the HK diagnostic locus. The pattern of sex-linked diapause physiology which we have docu­ mented for P. glaucus and P. canadensis (Rockey et al. 1987) clearly indicates that Dickinson County, Michigan has insufficient degree-days of thermal unit accumulation to ever support two generations of Papilio (even on the most nutritious host plant species; Scriber and Lederhouse 1992). This area also is near a large "cold pocket" which further shortens the growing season and which has apparently selected for pupation at small weights (Ayres and Scriber 1994) resulting in smaller adult females (as observed in forewing lengths that in this vicinity are generally less than 50 mm Scriber 1994, 1996b and Table 1). The total average growing season degree-day accumula­ tions of thermal units (To> 10°C) for this site in central Dickinson County is only 1000 (Eichenlaub et al. 1990). In one year out of 10 it is only 830, which is barely enough for even a single generation of P. canadensis to be completed (Scriber and Lederhouse 1992). Unfortunately, we couldn't assess the oblig­ ate diapause trait nor sex-linked oviposition preferences because our dark morph female did not lay any eggs. Long range transport of insects is believed to occur for other Lepidoptera, and the black cutworm (Agrotis ipsilon) reaches the northern Great Lakes and Canada on winds from the Southwest in mid-June (Showers 1977). This is about the same time of the year as this dark female Papilio was discov­ ered. It has been suspected during the 1980s that 1st instar ballooning gypsy moth larvae (Lymantria dispar) were blown into Wisconsin westward across Lake Michigan (Wisconsin Department of Agriculture, pers. comm.), but con­ vincing evidence is still lacking. Two day-flying Callosamia promethea moths (Saturniidae) were marked and recaptured in 3 days at 14 km and 36 km (both upwind, Toliver and Jeffords 1981). Hurricanes have apparently brought several tropical species into Texas (Neck 1978). However, to our knowledge, nothing as large as this female tiger swallowtail has been docu­ mented blowing in for distances 400-480 km (Gatehouse 1997), as we think is likely here. The arrival of monarch butterflies in Great Britain has been attributed to hurricanes assisted by the jetstream

ACKNOWLEDGMENTS We thank Drs. Guy Bush and Jim Smith for use of their laboratories for our biochemical analyses and Fred V. Nurnberger for obtaining the National Weather Service midwest regional data. We are also grateful for partial re­ search support for Mark Deering and Aram Stump from the Michigan Agri­ cultural Experiment Station (MAES projects #1644 and #1722) and NSF pro­ ject DEB9510044. We also thank Ann Swengel and Brian Scholtens for very thoughtful and helpful reviews of this manuscript.

LITERATURE CITED Ayres, M. P. and J. M. Scriber. 1994. Local adaptations to regional climates in Papilio canadensis (Lepidoptera: Papilionidae). EcoL Monogr. 64:465-482. Clarke, C. A and P. M. Sheppard. 1962. The genetics of the mimetic butterfly Papilio glaucus. Ecology 43:159-161. Clarke, C. A, P. M. Sheppard, and U. Mittwoch. 1976. Heterochromatin polymorphism and colour pattern in the tiger swallowtail butterfly, Papilio glaucus L. Nature 263:585-587. ~ __~ __a. __ ~ ______~ ______-'...

1998 THE GREAT LAKES ENTOMOLOGIST 159

Coombs, S. and V. Tucker. 1995. Comments on monarchs in England. Monarch Newsletter. Dec. 1995, p. 4. Eichenlaub, V. L., J. R. Harman, F. V. Nurnberger, and H. J. Stolle. 1990. The climate atlas of Michigan. Univ. of Notre Dame Press, Notre Dame, IN. 165 pp. Gage, S. 1996. Ten years of data reveal gypsy moth patterns. GatherlScatter National Laboratory for Computational Science and Engineering 12:20-21. Gatehouse, A G. 1997. Behavior and ecological genetics of wind-borne migration by in­ sects. Ann. Rev. Entomol. 42:475-502. Hagen, R. C. and J. M. Scriber. 1989. Sex linked diapause, color, and allozyme loci in Papillo glaucus: Linkage analysis and significance in a hybrid zone. Heredity 80:179-185. Hagen, R. C. and J. M. Scriber. 1991. Systematics of the Papilio glaucus and P. troilus groups (Lepidoptera: Papilionidae): inferences from allozymes. Ann. Entomol. Soc. Amer. 84:380-395. Hagen, R. H., R. C. Lederhouse, J. Bossart, and J. M. Scriber. 1991. Papillo canadensis and P. glaucus (Papilionidae) are distinct species. Jour. Lepid. Soc. 45:245-258. Lederhouse, R. C. 1978. Territorial behavior and reproductive ecology of the black swallowtail butterfly, Papilio polyxenes asterias Stoll. Ph.D. Dissertation, Cornell University, Ithaca, NY. Diss. abst. Ent. order no. 7902342. Lederhouse, R. C., M. P. Ayres, and J. M. Scriber. 1995. Physiological and behavioral adaptations to variable thermal environments in North American swallowtail but­ terflies. Pp. 71-82. In: (J. M. Scriber, Y. Tsubaki, and R. C. Lederhouse, eds.) Swal­ lowtail butterflies: Their ecology and evolutionary biology. Scientific Pub. Gainesville, FL. Luebke, H. J., J. M. Scriber, and B. S. Yandell. 1988. Use of multivariate discriminant analysis of male wing morphometries to delineate the Wisconsin hybrid zone for Pa­ pilio glaucus glaucus and P. g. canadensis. American MidI. Nat. 119:366-379. Neck, R. W. 1978. Climatic regimes resulting in unusual occurrences of Rhopalocera in Central Texas in 1968. Jour. Lepid. Soc. 32:111-115. Rockey, S. J., J. H. Hainze, and J. M. Scriber. 1987. Evidence of a sex-linked diapause response in Papilio glaucus subspecies and their hybrids. Physiological Entomol. 12:181-184. Scriber, J. M. 1982. Foodplants and speciation in the Papilio glaucus group. pp. 307-314. In: (J.H. Visser and AK. Minks, eds.) Proc. 5th International Symp. on In­ sect Plant Relationships, PUDOC, Wageningen, Netherlands. __. 1990. Interaction of introgression from Papilio glaucus canadensis and dia­ pause in producing 'spring form' Eastern tiger swallowtail butterflies, P. glaucus. Great Lakes Entomoi. 23:127-138. __. 1994. Climatic legacies and sex chromosomes: latitudinal patterns of voltinism, diapause, size and host-plant selection in 2 species of swallowtail butterflies at their hybrid zone. pp. 133-171. In: (H.V. Danks, ed.) Insect life-cycle polymorphism: the­ ory, evolution and ecological consequences for seasonality and diapause control. Kluwer Academic Publ. Dordrecht, Netherlands. 1996a. Tiger tales: Natural history of native North American swallowtails. Amer. Entomol. 42:19-32. __. 1996b. A new cold pocket hypothesis to explain local host preference shifts in Papilio canadensis. 9th Intern. Symp. Insects and Host Plants. Entomologia expt. & appl. 80:315-319. Scriber, J. M., M. D. Deering, L. N. Francke, W. Wehling, and R.C. Lederhouse. 1998. Notes on the swallowtail population dynamics of southcentral Florida (Highlands County). Holarctic Lepidoptera (in press). Scriber, J. M. and M. H. Evans. 1986. An exceptional case of paternal transmission of the dark form female trait in the tiger swallowtail butterfly, Papilio glaucus (Lepi­ doptera: Papilionidae). J. Research Lepid. 25: 110-120. Scriber, J. M. and S. H. Gage. 1995. Pollution and global change: Plant ecotones, but­ 160 THE GREAT LAKES ENTOMOLOGIST Vol. 3 T, No.3 & 4

terfiy hybrid zones, and biodiversity. pp. 31S-3M.In: (J.M. Scriber, Y. Tsubaki. and RC. Lederhouse, eds.) Swallowtail butterflies: Their ecology and evolutionary biol­ ogy. Scientific Pub., Gainesville, FL. Scriber, J. M., R H. Hagen, and R C. Lederhouse. 1996. Genetics of mimicry in the tiger swallowtail butterflies, Papilio glaucus and P. canadensis (Lepidoptera: Papil­ ionidae). Evolution 50:222-236. Scriber, J. M. and R C. Lederhouse. 1992. The thermal environment as a resource dic­ tating geographic patterns of feeding specialization of insect herbivores. pp. 429-466. In: (M.R Hunter, T. Ohgushi, and P.W, Price, eds.) Effects of Resource Dis­ tribution on Animal-Plant Interactions. Academic Press, Showers, W. B. 1997. Migratory ecology of the black cutworm. Ann. Review Entomol. 42:393-425. Toliver, M. E. and M. R. Jeffords. 1981. Long distance dispersal by Callosamia promethea (Saturniidae). Jour. Lepid. Soc. 35:76. 1998 THE GREAT LAKES ENTOMOLOGIST 161

EFFECTS OF ENAMEL PAINT ON THE BEHAVIOR AND SURVIVAL OF THE PERIODICAL CICADA, MAG/CICADA SEPTENDECIM (HOMOPTERA) AND THE LESSER MIGRATORY GRASSHOPPER, MELANOPWS SANGUINIPES IORTHOPTERA}.

J. R. Cooley, G. S. Hammond, and D. C. MorsholP

ABSTRACT We present information compiled from several studies on the effects of methods for marking individual arthropods on their longevity and behavior. Results from our own research on effects of enamel paint marking on two in­ sect species, the periodical cicada, Magicicada septendecim, and the lesser migratory grasshopper, Melanoplus sanguinipes, are also presented. Neither species showed any adverse survivorship or behavioral effects from marking.

The ease of use of different marking methods and their appropriateness for particular study designs are well reviewed (Southwood 1978, Walker and Winewriter 1981, Kearns and Inouye 1993). Above and beyond logistical con­ cerns, the possible effects on the stu dects further constrains the choice of marking method. Table 1 summ 'terature reporting effects of indi­ vidual marking (not mass-marking) methods on study organisms. We located sources using published reviews and electronic databases (Zoological Record, Wilson Indexes to Journal Articles, and Biological Abstracts) and included only papers containing data specifically addressing possible mortality or morbidity due to marking. Of several hundred papers examined, we found only 19 fitting these criteria. This table is neither exhaustive nor prescrip­ tive, but rather supplements earlier reviews. Few general trends are appar­ ent, except that marks applied topically to sclerotized parts seemed to have few deleterious effects, while dusts, powders, or, in the case of soft-bodied in­ sects, some solvents, must be used cautiously. The lack of general trends means that controlled testing of a marking method is the only way to ensure its appropriateness for any given study. Individual marking has proven effective in our research on periodical ci­ cadas [Magicicada septendecim (L.)] and lesser migratory grasshoppers [Melanoplus sanguinipes (F.)], allowing us to track the behaviors of individ­ ual insects or classes of insects as they emerge, mature, mate, and die. One marking technique we have used is to apply colored enamel paint dots to the dorsal thoracic surface (Testors® lead-free gloss enamel; Testors Corp., Rock­ ford IL, 61104), or to use paint pens (Uni®·Paint Fine Line PX-21 paint markers; Eberhard Faber Inc., Lewisburg TN 37091). Because published reo

lInsect Division, The University of Michigan Museum of Zoology, Ann Arbor, MI, 48109-1079 'l'able 1. Reported effects of methods for marking individual insects, grouped by general method and organized chronologically within 0­ each grouping. I'V Reference Marker Taxon Effect* This study Enamel paint (xylol, propanol based) Magicicada septendecim (Homoptera) Melanoplus sanguinipes (Orthoptera) None Hunter 1960 Enamel paint (xylol, propanol based) Laelaspis georgiae (mite) None et al. 1990 Enamel paint (xylol, propanol based) Leptin.otarsa decemlin.eata (Coleoptera) None Hager and Kurczewski 1986 Enamel paint (xylol, propanol based) Ammophila harti (Hymenoptera) None§ -i Greenslade 1964 Enamel paint (petroleum-solvent based) Nebria breuicollis (Coleoptera) Nonet I m Paulson and Akre 1991 Enamel paint (petroleum-solvent based) Formica neoclara (Hymenoptera) None G) ;:<:l Dobson et al. 1958 Oil-based paint Lepthohylemia coarctata (Diptera) None m ~ Dobson et al. 1961 Lacquer-based paint (nitrocellulose) Lepthohylemia coarctata (Diptera) Toxic :; A Davey 1956 Oil-based paint (Rubine 'funer or Schistocerca gregaria (Orthoptera) m Monolite Yellow in wood rosin and kerosene) None C/l m Gallepp and Hasler 1975 Typewriter correction fluid Brachycentrus spp. (Trichoptera) Behavioral Z -i changes 0 Fales et al. 1964 Fingernail polish (acetone based) Musca autumnalis (Diptera) Toxic ~ 0r- Garcia and Paleari 1990 Fingernail polish (acetone based) Charidotis punctatostriata (Coleoptera) None 0 G) Dreyer and Fingernail polish (acetone based) Clauigralla tomentosicollis (Hemiptera) Vi -i Baumgartner 1997 None Brussard 1971 "Sharpie" marker (xylol solvent) Erebia epipsodea (Lepidoptera) None Rieske and Raffa 1990 Acrylic paint Hylobius pales (Coleoptera) ~ None w ~ Caprio et at. 1990 Paper labels affixed with cynoacrylate glue Leptin.otarsa decemlineata (Coleoptera) None - Z Caprio et at. 1990 Acetone wash, followed by Leptin.otarsa decemlineata (Coleoptera) ~ paper labels aft'"txed with cynoacrylate glue None w Qo Hart and Resh 1980 Plastic tags aftixed with wire or elastic band Dicosmoecus giluipes (Trichoptera) None ..,.. Sempala 1981 Fluorescent powder Aedes africanus (Diptera) -0 combined with Duco paint Toxic -0 CD Gangwere et al. 1964 Colored thread or paper attached with glue Acheta domesticus (Orthoptera) None:\: Stuart 1986 Polyester fiber tied around body Leptothorax spp., Harpagoxenus spp. (Hymenoptera) None Gangwere et al. 1964 Melted crayon, colored ink, Aniline dyes, or Acheta domesticus (Orthoptera) Titanium oxide spray None:\: Gangwere et al. 1964 Metallic dust Acheta domesticus (Orthoptera) None:\: Gangwere et al. 1964 Fluorescent powder, pigment in oil spray, Acheta domesticus (Orthoptera) Toxic in :2 Oil and Cellulose paints, dry pigment large doses m Gangwere et al. 1964 Wing amputation or clipping Periplaneta americana (Blattaria) None for G) ;>C moderate m treatment ~ Mead-Briggs 1964 Tarsal clipping SpilopsyUus cuniculi (Siphonaptera) None S; A Mascanzoni and Wallin 1986 Radar tags Pterostichus melanarius, P. niger, Harpalus m rufipes, Carabus granulatus (Coleoptera) None C/l m Z * "None:" no effects reported; "Toxic:" increased mortality reported; "Injury:" unspecified damage or mortality noted. --i § Pers. Comm. o t Dispersal attributable to handling effects noted. ~ o :\: Method proved to be impermanent and of limited use. 5 G) ~

0­ W 164 THE GREAT LAKES ENTOMOlOGIST Vol. 31, No.3 & 4

ports contain no clear information on the toxicity of the solvents in these paints (xylene, xylol and n- propoxy propanol) we conducted experiments to determine whether this marking method has adverse effects on our study an­ imals.

MATERIALS AND METHODS Periodical cicadas. We compared the survivorship and mating activity of marked and unmarked adult Magicicada septendecim in a recently logged clearing along Route 639 in the Horsepen Lake State Wildlife Management Area, Buckingham County, VA, in mid-May 1996. Two cages were con­ structed by enclosing living vegetation with a 1 x 2 m piece of black fiber­ glass window screen; each cage was stocked with 10 unmarked and 10 marked mature, unmated female cicadas. We marked each cicada by using a flat toothpick to apply one yellow and two blue dots of Testors® enamel paint. We used no anesthetic during marking, and we were careful to prevent paint from contacting the wing articulations or the articulation between the head and thorax. The total surface area covered by paint was between 6 and 10 mm2. Unmarked cicadas were handled in similar fashion. To allow matings, on May 20, 1996, we captured 20 sexually mature adult males from the sur­ rounding chorus and placed 10 in each female cage. Cages were censused on May 23, 24, 29, 30, 31, June 1, 2, and 4. At each census, we counted and removed dead cicadas within each cage, recorded whether or not they were marked or showed evidence of a mating sign (or "seminal plug;" White 1973), and preserved them in 70% ethanoL We re­ moved, examined, and preserved all remaining cicadas at the final census. Lesser migratory grasshoppers. We collected male and female nymphs of Melanoplus sanguinipes (F.), the lesser migratory grasshopper, from the grounds of the Matthaei Botanical Gardens at the University of Michigan, Ann Arbor, Michigan. We raised the grasshoppers to maturity in 50 x 40 x 40 em fiberglass screen cages, on a diet of romaine lettuce, wheat bran and wheat seedlings ad libitum. Cages were kept at room temperature (20°C ± 2°), but provided with a 90 watt incandescent light bulb on a 13:11h light: dark cycle to allow for thermoregulation. Twenty one-week-old adults of each sex were chosen for marking and handled individually; half (10 of each sex) were marked with two dots of paint on the pronotum. We used four different colors of Uni®-PaintFine Line paint markers. All 40 individuals were maintained for 6 days in a 50 x 40 x 40 cm cage under the same conditions described above. We examined the cage twice every day, once between 09:00 and 12:00 and once between 13:00 and 19:00, and recorded the paint/no-paint status of all copulating individuals, as well as any deaths.

RESULTS Periodical cicadas. We compared the survivorship of the marked and unmarked cicadas using Cox regression analysis, implemented with the PHREG function in SAS (SAS Institute Inc., 1996). This technique allowed us to include individuals surviving at the end of our observations. The small difference in survivorship curves for marked and unmarked individuals was not statistically significant (Wald Chi-Square = -2.32, 1 d.f., p = 0.13), and the trend was in the direction of marked cicadas seeming to survive longer than unmarked cicadas. Mortality rates in the two cages differed (Wald Chi­ 1998 THE GREAT LAKES ENTOMOlOGIST 165

10

c'" 6

3 I CD 5-+­ ,Q E i 4

0 ~.

Square = 14.02, 1 d.f., p = 0.0002), probably due to the death of substantial amounts of vegetation in one cage, although the increased mortality in this cage was distributed equally among marked and unmarked cicadas. A chi-square goodness-of-fit analysis of seminal plug presence/absence data reveals no significant pattern. Marked and unmarked females were equally likely to exhibit this indicator of mating status. Data are summa­ rized in Figure 1 and Table 2. Lesser migratory grasshoppers. We observed 31 copulations over 6 days. Of these, 8 were between unmarked females and unmarked males, 6 were between marked females and unmarked males, 10 were between un­ marked females and marked males, and 7 were between marked females and marked males. There was no difference in likelihood of marked or unmarked males or females being in copula (Chi-Square Goodness-of-Fit Test, X2=1.31, p»0.10). During the 6 days after marking there was no mortality or appar­ ent illness in any of the marked or unmarked individuals.

DISCUSSION We have uncovered no evidence that careful marking with enamel paints affects cicadas' or grasshoppers' mortality during the short « 12 day) study period. Furthermore, marking does not affect the behavior or attractiveness of female cicadas in any way that alters likelihood of mating. For the grasshoppers, paint marking appeared to have no impact on the sexual be­ havior of either sex. 166 THE GREAr LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4

Table 2. Results of "sperm plug" analysis, demonstrating that marking had no effect on tendency of female Magicicada septendecim to mate. Thtal "sperm plug" Females present X2 p --...... ------..".----..".--"------...:..:...----...:..:...-­ Females in replicate A with "sperm plug" Marked 10 6 Unmarked 10 7 Thtal 20 13 0.038 p»O.l (n.s.)

Females in replicate B with "sperm plug" Marked 10 5 Unmarked 10 6 Thtal 20 11 0.045 p»O.l (n.s.)

Females in combined replicates with "sperm plug" Marked 20 11 Unmarked 20 13 Thtal 40 24 0.083 p»O.l (n.s.)

From our experiences in the field, we can draw attention to several pit­ falls best avoided when marking insects with enamel paints. First, paint ap­ plied to body parts that undergo significant flexion (such as the wings or dor­ sum of the thorax in flying insects) tends to wear or flake off, usually beginning a week to ten days after marking, but sometimes noticeably after only a day or two. Secondly, the pigments in enamel paints fade under daily exposure to sunlight; thus some colors may become indistinguishable within a week of marking. The difficulty in distinguishing similar pigments such as blue and violet can be exacerbated by the varying light conditions in the field; the contrast between two given pigments differs depending on atmos­ pheric conditions, orientation of the marked insect, or time of day. Redun­ dancy in the marking scheme alleviates this problem to some extent. Finally, enamel paint spots fade and flake oft'when marklpd specimens are preserved in alcohol (we have experience with ethanol and denatured alcohol; other commonly available alcohols probably give similar results).

ACKNOWLEDGMENTS Funding was provided by the Insect Division of the University of Michi­ gan Museum of Zoology.

LITERATURE CITED Brussard, P. F. 1971. Field techniques for investigations of population structure in a "ubiquitous" butterfly. Jour. Lepidop. Soc. 25: 22-29. Caprio, M. A, Miller, D., and Grafius, E. 1990. Marking adult Colorado potato beetles, Leptinotarsa decemlineata (Coleoptera: Chrysomelidae), using paper labels. Great Lakes Entomol. 23: 13-18. F"".----~------I1'-....-

1998 THE GREAT LAKES ENTOMOLOGIST 167

Davey, J. T. 1956. A method of marking isolated adult locusts in large numbers as an aid to the study oftheir seasonal migrations. Bull. EntomoI. Res. 46: 797-802. Dobson, R. M., Stephenson, J. W., and Lofty, J. R. 1958. A qnantitiative study of a pop­ ulation of wheat bulb fly, Leptohylemia coarctata (FalI.), in the field. Bull. EntomoL Res. 49: 95-111. Dobson, R. M., and Morris, M. G. 1961. Observations on emergence and life-span of wheat bulb fly, Leptohylemia coarctata (Fall.), under field cage conditions. Bull. En­ tomol. Res. 51: 803-821. Dreyer, H., and Baumgartner, J. 1997. Adult movement and dynamics of Clavigralla tomentosicollis (Heteroptera: Coreidae) populations in cowpea fields of Benin, West Mrica. Jour. Econ. Entomol. 90: 421-426. Fales, J. H., Bodenstein, O. F., Mills, G. D., Jr., and Wessel, L. H. 1964. Preliminary studies on face fly dispersion. Ann. Entomol. Soc. Am. 57: 135-137. Gangwere, S. K, Chavin, W., and Evans, F. C. 1964. Methods of marking insects, with especial reference to Orthoptera (Sens. Lat.). Ann. EntomoI. Soc. Am. 57: 662-669. Gallepp, G. W. and Hasler, A D. 1975. Behavior of larval caddisflies (Brachycentrus spp.) as influenced by marking. Am. MidI. Nat. 93: 247-254. Garcia, M. A, and Paleari, L. M. 1990. Marking cassinidae (Coleoptera: Chrysomeli­ dae) larvae in the field for population dynamics studies. Entomol. News 101: 216-218. Greenslade, P. J. M. 1964. The distribution, dispersal, and size of a population of Ne­ bria brevocillis (F.) with comparative studies on three other Carabidae. J. Anim. Ecol. 33: 311-333. Hager, B. J., and F. E. Kurczewski. 1986. Nesting behavior ofAmmophila harti (Fer­ nald) (Hymenoptera: Sphecidae). Am. MidI. Nat. 116: 7- 24. Hart, D. D. and Resh, V. H. 1980. Movement patterns and foraging ecology of a stream caddisfly larva. Can. J. ZooL 58: 1174-1185. Heller, K G. and vonHelverson, O. 1990. Survival of a phaneropterid bush-cricket studies by a new marking technique (Orthoptera: Phaneropteridae). Entomol. Gener. 15: 203-208. Hendricks, D. E., and Graham, H. M. 1970. Oil-soluble dye in larval diet for tagging moths, eggs, and spermatophores of tobacco budworms. J. Econ. EntomoL 63: 1019-1020. Hunter, P. E. 1960. Plastic paint as a marker for mites. Ann. Entomol. Soc. Am. 53: 698. Kearns, C. A, and Inouye, D. W. 1993. Techniques for pollination biologists. University Press of Colorado, Niwot CO. Mascanzoni, D., and Wallin, H. 1986. The harmonic radar: a new method oftracing in­ sects in the field. EcoL EntomoL 11: 387-390. Mead-Briggs, A R. 1964. Some experiments concerning the interchange of rabbit fleas, Spilopsyllu8 cuniculi (Dale), between living rabbit hosts. J. Anim. Ecol. 33: 13-26. Paulson, G. S., and Akre, R. D. 1991. Role of predaceous ants in pear psyUa (Ho­ moptera: Psyllidae) management: estimating colony size and foraging range of Formica neoclara (Hymenoptera: Formicidae) through a mark-recapture technique. Jour. Econ. Entomol. 84: 1436-1440. Rieske, L. K and Raffa, K F. 1990. Dispersal patterns and mark-and-recapture esti­ mates of two pine root weevil species, Hylobius pales and Pachylobius picivorus (Coleoptera: Curculionidae), in Christmas tree plantations. Environ. EntomoI. 19: 1829-1836. SAS Institute Inc. 1996. SASISTAT® Software: Changes and Enhancements through Release 6.11. Cary, NC: SAS Institute Inc. Sempala, S. D. K 1981. The ecology of Aedes (Stegomyia) africanus (Theobald) in a tropical forest in Uganda: mark-release-recapture studies on a female adult popula­ tion. Insect Sci. Application 1: 211-224. 168 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4

Southwood, T. R. E. 1978. Ecological methods, with particular reference to the study of insect populations. Chapman and Hall, London. Stuart, R. J. 1986. Use of polyester fibers to mark small leptothoracine ants (Hy­ menoptera: Formicidael. Jour. Kans. Entomol. Soc. 59: 566-568. Walker, T. J., and Winewriter, S. A 1981. Marking techniques for recognizing individ­ ual insects. Florida Entomol. 64: 18-29. White, J. 1973. Viable hybrid young from crossmated periodical cicadas. Ecology 54: 573-580. 1998 THE GREAT LAKES ENTOMOLOGIST 169

THE PARASITOID COMPLEX OF FIRST GENERATION OSTR/N/A NUB/LAUS (LEPIDOPTERA: PYRALIDAEI LARVAE IN NORTHWEST OHIO

Daniel M. Pavuk l and Laura l. Hughes]

ABSTRACT A survey of first generation European corn borer (Ostrinia nubilalis) lar­ vae was conducted during 1997 in six cornfields located in northwestern Ohio in order to determine the larval parasitoid complex utilizing this host. Collected larvae were held under constant conditions until the larvae com­ pleted development or parasitoids emerged. The following species were recorded: Eriborus terebrans (Hymenoptera: Ichneumonidae), Macrocentrus grandii (Hymenoptera: Braconidae), Sympiesis viridula (Hymenoptera: Eu­ lophidae), and Lixophaga sp. (Diptera: Tachinidae), Levels of parasitism in the different fields ranged from 14.3 to 83.3%. Future research will include surveys of additional fields and sampling of O. nubilalis over the entire sea­ son in the northwest Ohio region.

The European corn borer, Ostrinia nubilalis (Hubner) (Lepidoptera: Pyralidae), is a widespread and significant pest of both field and sweet corn in the United States (Showers et aL 1989). Because O. nubilalis is not native to North America, it became the focus of a classical biological control pro­ gram after it was introduced and discovered. Between 1920 and 1938, 24 parasitoid species were released in the United States from Europe and the Orient (Baker et al. 1949). In the North Central Region, the introduced par­ asitoids that were commonly recovered in European corn borer surveys dur­ ing subsequent years were Lydella thompsoni (Herting) (Diptera: Ta­ chinidae), Macrocentrus grandii (Goidanich) (Hymenoptera: Braconidae), and Eriborus terebrans (Gravenhorst) (Hymenoptera: Ichneumonidae), In Ohio, the most commonly recovered parasitoids in recent years have been E. terebrans and M. grandii. Collections of second generation larvae have been made by researchers in various parts of the state (e.g., Pavuk & Stinner 1992, Mason et al. 1994). The last comprehensive, statewide survey was per­ formed by Mason et al. (1994); this survey focused on overwintering, second generation European corn borer larvae present in eight states, including Ohio. The parasitoids recovered were species mentioned above (Mason et aI. 1994). In addition, another tachinid, Lixophaga sp., was found in western Ohio and western North Carolina, but was extremely uncommon (Mason et al. 1994). We were interested in determining, in a preliminary survey within the

lDepartment of Biological Sciences, Bowling Green State University, Bowling Green, OH 43403-0212. 170 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No.3 & 4 northwestern Ohio region, which parasitoids were attacking O. nubilalis first generation larvae. During late July and early August 1997 (28 July-6 Au­ gust), we collected first generation larvae from six cornfields in Hancock, Henry, and Wood Counties. These are contiguous counties in the northwest­ ern region of Ohio. All of the fields sampled were similar in terms of agro­ nomic practices, with the exception of two fields (Shaffer Fields 1 and 2) that were planted using a narrow row spacing (51 cm, or 20 in). The narrow row cornfields did not appear to have substantially smaller or larger European corn borer larval populations than cornfields planted using typical row spac­ ing (76 cm, or 30 in) (Pavuk and Hughes, personal observation). Within each field, 25 plants were dissected at each of four locations; two locations were on two edges of the field, and two locations were situated in the approximate center of the field, 25 meters apart. All of the larvae were 4th or 5th instars at the time of collection. The larvae were maintained in the laboratory under constant conditions (16 h: 8h, L:D; 25° C, and apyroximately 70% RH) until either parasitoids emerged or the larvae completed development. We ex­ cluded larvae that died from disease or unknown causes when calculating percent parasitism. The most common parasitoid was Eriborus terebrans; Macrocentrus grandii was the second most common species reared from the larvae (Table 1). In addition, one larva was parasitized by the eulophid Sympiesis viridula, and two specimens of Lixophaga sp. were reared from larvae. Populations of O. nubilalis were extremely small, especially in com­ parison to the previous growing season (1996) (D. M. Pavuk, personal obser­ vation). The second generation was also very small, and few larvae were col­ lected from the same cornfields where the first generation larvae were found. No parasitoids were reared from the second generation larvae collected (n < 10 larvae for each field sampled). We observed a wide range in percent parasitism of larvae collected from the six fields; total percent parasitism ranged from 14.3% to 83.3% (Table 1). The highest level of parasitism occurred in the largest field surveyed (Ben­ nett Field 1). However, this field had an extremely small O. nubilalis larval population (only 6 larvae were collected), and the five larvae that were para­ sitized all came from an edge near a large wooded area. This may indicate that parasitoids were moving easily from the neighboring woodlot to the field edge; the woodlot may have been providing su.itable refuges for the para­ sitoids. The use of adjacent woodlots by parasitoids of European corn borer, especially Eriborus terebrans, was suggested by Landis & Haas (1992) and Dyer & Landis (1997). Another factor that may eventually have effects on parasitism rates of O. nubilalis larvae is the increasing use of Bt-corn (transgenic corn in which the Bacillus thuringiensis toxin-producing gene has been incorporated). Hilbeck et aL (1998) found that Chrysoperla carnea larvae reared on Bacillus thuringiensis-fed O. nubilalis larvae had a higher mortality rate than C. camea immatures that were fed B. thuringiensis-free prey. Parasitoids may also possibly suffer sub-lethal effects and increased mortality while develop­ ing in or on O. nubilalis larvae that are feeding on Bt-corn plants. Also, re­ duced popUlations of O. nubilalis due to the planting of Bt-corn may make it more difficult for parasitoids to locate larvae, which could adversely affect the persistence of parasitoid populations. We estimated that the amount of Bt-corn planted.in the tri-county region we surveyed to be less than 5% in 1997; however, this amount is likely to increase in the next several years. It will be interesting to track the changes in parasitism of O. nubilalis that may occur because of increasing use of Bt-corn. We intend to continue our studies of the parasitoids of Ostrinia nubilalis in northwest Ohio. More intensive collecting of O. nubilalis larvae may re­ Table 1. Parasitoids Reared from First Generation Ostrinia nubilalis Larvae, 1997. -0 -0 No. of Larvae co Location & Field* No. of Larvae Parasitoid Parasitized % Parasitism Wood Co.; Shaffer - Field 1 (-20 ha;51 cm row spacing; 24 Eriborus terebrans 7 29.2% COITl fields on Nand Wedges) Macrocentrus grandii 3 12.5% Total: 41.7%

Wood Co.; Shaffer - Field 2 (-10 ha; 51 cm row spacing; 11 Eriborus terebrans 4 36.4% ::r:-l trees on Sand Wedges) Macrocentrus grandii 2 18.2% m Total: 54.6% G) Al Hancock Co.; Welly • Field 1 m (-5 ha; 76 cm row spacing; 17 Eriborus terebrans 5 29.4% ~ woods on E and Wedges) Macrocentrus grandii 2 };: 11.8% A Sympiesis viridula m 1 5.9%

Henry Co.; Bennett- Field 2 (-10 ha; 76 cm row spacing; 14 Eriborus terebrans 2 Total: 14.3% hedgerows on E and Sedges)

*Some type of reduced tillage used in all fields, e.g., chisel plowed in the fall and field cultivated in the spring. 'I 172 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No.3 &4 veal trends in parasitism and will enable us to more thoroughly elucidate the parasitoid complex of this important corn herbivore.

ACKNOWLEDGMENTS We thank Rich Bennett, Lyle and Phil Shaffer, and Brian Welly for allow­ ing us to collect Ostrinia nubilalis larvae from their cornfields. We also thank Curtis Young of Ohio State University and Alan Sundermeier, Henry County Cooperative Extension, for their assistance with this project.

LITERATURE CITED Baker, W.A, W.G. Bradley, and C.A Clark. 1949. Biological control of the European corn borer in the United States. USDA Technical Bulletin 983. Dyer, L.E. and D.A. Landis. 1997. Influence of noncrop habitats on the distribution of Eriborus terebrans (Hymenoptera: Ichneumonidael in cornfields. Environ. Entomol. 26: 924-932. Hilbeck, A, M. Baumgartner, P.D. Fried, and F. Bigler. 1998. Effects of transgenic Bacillus thuringiensis corn-fed prey on mortality and development time of immature Chrysoperla carnea (Neuroptera: Chrysopidael. Environ. Entomol. 27: 480-487. Landis, D.A and M.J. Haas. 1992. Influence of landscape structure on abWldance and within-field distribution of European corn borer (Lepidoptera: Pyralidael larval parasitoids in Michigan. Environ. Entomol. 21:409-416. Mason, C.E., RF. Romig, L.E. Wendel, and L.A. Wood. 1994. Distribution and abWl­ dance of larval parasitoids of European corn borer (Lepidoptera: Pyralidael in the East Central United States. Environ. Entomol. 23: 521-531. Pavuk, D.M. and B.R Stinner. 1992. Influence of weed commWlities in corn plantings on parasitism of Ostrinia nubilalis (Lepidoptera: Pyralidael by Eriborus terebrans (Hymenoptera: Ichneumonidael. BioI. Control 2: 312-316. Showers, W.B., J.F. Witkowski, C.E. Mason, D.D. Calvin, RA Higgins, and G.P. Dively. 1989. European corn borer development and management. North Central Regional Extension Publication No. 327. 1998 THE GREAT LAKES ENTOMOlOGIST 173

TEMPERATURE EFFECTS ON DEVELOPMENT IN APHELINUS ALBIPODUS (HYMENOPTERA: APHELINIDAE) FROM TWO GEOGRAPHIC REGIONS

Jang-Hoon lee l ,2 and Norman C. Elliottl

ABSTRACT Aphelinus albipodus Hayat & Fatima was imported to the United States for classical biological control of the Russian wheat aphid, Diuraphis noxia (Mordvilko). Temperature effects on development of A. albipodus from two geographic regions (hereafter referred to as strains) were measured using the Russian wheat aphid as host. Temperature thresholds for egg to mummy, mummy to adult, and egg to adult development were 8.9,10.9, and 9.7°C for A. albipodus collected near Pingluo, China, and were and 8.5, 10.3, and 9.6°C for A. albipodus collected near Urumqi, China. The time required to develop from egg to adult did not differ among strains. However, when total imma­ ture development was partitioned into egg to mummy and mummy to adult, the time required for development through these two periods differed among strains. The Urumqi strain developed faster than the Pingluo strain from egg to mummy, while the Pingluo strain developed faster from mummy to adult. Degree-day requirements for egg to mummy development were 135 and 104 for the Pingluo and Urumqi strains, respectively. Corresponding require­ ments for mummy to adult development were 70 and 101 degree-days. The ability to vary immature development rate in response to climate or other factors could have adaptive significance because it would permit the para­ sitoid to exploit environments over a broad geographic range.

The Russian wheat aphid, Diuraphis noxia (Mordvilko), is a recently in­ troduced pest in North America (Stoetzel 1987). The aphid spread rapidly throughout the western cereal growing regions of United States and Canada (Jones et at 1989) where it is now a serious pest of wheat and barley. Apheli­ nus albipodus Hayat & Fatima is one of several exotic parasitoid species im­ ported into the United States from several regions including Europe and Asia for classical biological control of the Russian wheat aphid (Tanigoshi et at 1995). Recoveries of A. albipodus were recently reported from Colorado (EI· liott et aL 1995a) and Washington State (Tanigoshi et al. 1995), indicating that the ecological requisites are present for the parasitoid to establish repro­ ducing popUlations in these areas. The parasitoid is sometimes abundant in the field. Developmental response to temperature is one of several biological char­ acteristics that can influence the ability of a parasitoid to survive and be­

lUSDA, ARS, Plant Science Research Laboratory, 1301 N. Western St, Stillwater, OK 74075. 2Current address: Department of Applied Biology, Dongguk University, Seoul, Korea 100-715. ==~======~~======--~======----~~

174 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 &4

come an effective biological control agent in a new region. The development threshold and degree-day requirements are two parameters for describing developmental response of insects to temperature. The developmental threshold is defined as the temperature below which no measurable develop­ ment occurs, and the development time in degree-days is defined as the amount of heat above the developmental threshold required to complete de­ velopment through a particular life stage (Campbell et al. 1974). The timing of appearance of insect populations in the field is largely determined by their developmental responses to temperature, which can be summarized in terms of the developmental threshold and degree-day requirements for develop­ ment (Frazer & Gilbert 1976, Gilbert et al. 1976). The appearance of natural enemy populations early in the growing sea­ son, when aphid populations are low is thought to be particularly important in aphid biological control. A high ratio of natural enemies to aphids at that time may allow the natural enemies to suppress the aphid population early in the growing season before the period of rapid aphid population growth (van Emden 1988). Knowledge of the developmental threshold and degree­ day requirements for development can yield insight as to whether a para­ sitoid to be used as a prospective biological control agent will exhibit early appearance and activity in the field. Morris (1971) and Morris and Fulton (1970) were among the first to ob­ serve that insect populations in different geographic areas sometimes exhibit different developmental responses to temperature in laboratory studies. They found a difference in the degree-day requirements for development, but not the developmental threshold, of fall webworm populations, Hyphantria cunea (Drury) (Lepidoptera), from New Brunswick and Nova Scotia. Geo­ graphic populations of parasitoids are sometimes differently adapted to cli­ mate, due to natural selection in their region of origin (Messenger 1971). As a result, inherent differences sometimes exist among strains in their response to temperature or other climatic variables that influence their ability to es­ tablish in a region where they are released. Furthermore, if they do estab­ lish, these differences can influence the ability of a strain to control the tar­ get pest in the region (Messenger 1971, Messenger and van den Bosch 1971). This study was conducted on A. albipodus obtained from two geographic regions (hereafter referred to as strains). Our objectives were to determine if differences existed among the A. albipodus strains in the effects of tempera­ ture on immature development that might foretell differences in climatic adaptation among the strains. The information obtained may be useful for assessing whether strain-specific differences in biology exist that might be exploited in developing release strategies for A. albipodus, and in assessing the likelihood that the parasitoid will be effective as a biological control agent of the Russian wheat aphid.

MATERIALS AND METHODS The A. albipodus laboratory colonies used in this study were obtained from the USDA, ARS European Biocontrol Laboratory, Montpellier, France. The parasitoid strains were collected from Russian wheat aphid hosts in wheat fields near Pingluo (38° 5'N, 106° 30'E) and Urumqi (43° 45'N, 87° 45'W) China in the spring of 1992. From 100 to 200 individuals of each strain were received as founder colonies in the summer of 1992 and were main­ tained on Russian wheat aphids for 5-6 generations prior to initiating the study. Laboratory colonies were maintained in a plant growth chamber at 16L:8D, 22±0.5°C, and 55±5% relative humidity. 1998 THE GREAT LAKES ENTOMOlOGIST 175

To establish an experiment, 50 1st-3rd instar Russian wheat aphid nymphs were placed on each of 5-7 barley (Hordeum vulgare L.) seedlings growing in 10-cm diameter plastic pots (one seedling per pot) using an artist's brush. Each potted plant was then covered with a vented clear plastic cylindrical cage (10-cm diameter - 3D-em height). After allowing the aphids to settle for 4 h, 10 active female parasitoids were randomly taken from a stock colony using an aspirator and introduced into the cage. After 4 h expo­ sure, the parasitoids were removed from the cage and the caged plant was placed in a growth chamber maintained at a constant temperature of 14, 18, 22, or 26°C (all ±0.5°C), 16L:8D, and 55±5% relative humidity. The plants were checked daily, and mummies that formed were removed from plants, placed in a plastic petri dish (5-cm diameter - l.4-em height), and returned to the plant growth chamber. Petri dishes were shaded from direct light in order to minimize heat build-up. Mummies were examined daily for adult emergence. The number of days required for development from egg to mummy formation and the number of days for development from mummy formation to adult emergence from the mummy were recorded for each mummy that formed. Developmental threshold temperatures and develop­ ment times in degree-days for the period from egg to mummy, mummy to adult, and egg to adult were estimated by least squares regression [PROC REG (SAS Institute 1988)]. Average development rate (l/development time in days) for each temperature was calculated based on all individuals that developed to a particular stage at that temperature. The developmental threshold and degree-day requirements for development from egg to adult emergence were estimated from a linear regression of development rate (Y) against temperature (X):

Y =a + bX

The developmental threshold is the intercept of the X-axis (-alb) and de­ velopment time in degree-days is the reciprocal of the regression coefficient (lib) (Campbell et al. 1974). Standard errors of the estimates of the develop­ mental threshold and degree-day requirement were calculated using formu­ lae presented in (Campbell et al. 1974). Tests of the statistical hypotheses that developmental thresholds and degree-day requirements did not differ among strams were made using t-te8t8 (Campbell et al. 1974). Developmen­ tal times were compared among strains using ANOVA and the LSD test [FROC GLM (SAS Institute 1988)].

RESULTS The number of days required for development from egg to mummy, mummy to adult, and egg to adult for the two A. albipodus strains are listed in Table 1. Development times for the three developmental periods decreased as temperature increased for both strains. The two strains showed similar developmental responses to temperature for the egg to adult period. For ex­ ample, the Pingluo strain required 41.8 d (SE 0.27) to complete develop­ ment at 14°C, whereas the Urumqi strain required 42.8 d (SE =0.28), At 26°C both strains required 12.2 d to develop from egg to adult. When egg to adult development time was partitioned into times from egg to mummy and mummy to adult, differences in developmental times were evident among strains. The Urumqi strain developed faster from egg to mummy at all tem­ peratures than did the Pingluo strain, with the difference averaging 9.5,3.7, 1.6, and 2.2 d at 14, 18, 22, and 26 Q C, respectively. Conversely, the Pingluo 176 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4

Table 1. Average number of days for three developmental periods for Aphelinus al­ bipodus at four temperatures. Standard errors are enclosed in parentheses. Temperature (00) Place of OriginlLife 14 18 22 26 Pingluo, China sample size 46 53 64 51 egg-mummy formation 27.0* (0.23) 15.0* (0.11) 10.0* (0.10) 8.00* (0.07) mummy formation-adult 14.8* (0.27) 11.1* (0.18) 7.6* (0.18) 4.2* (0.08) egg-adult 41.S (0.27) 26.1 (0.18) 17.6 (0.19) 12.2 (O.OS) Urumqi, Ohina sample size 51 55 56 49 egg-mummy formation 17.5* (0.27) 11.3* (0.15) S.4* (0.15) 5.8* (O.OS) mummy formation-adult 25.3* (0.23) 14.1 * (0.23) S.S* (0.13) 6.4* (O.OS) egg-adult 42.S (0.2S) 25.4 (0.24) 17.2 (0.17) 12.2 (0.09) Mean developmental time for a particular stage differs significantly (P < 0.01) among strains if followed by an asterisk.

strain developed faster from mummy to adult than the Urumqi strain at all temperatures. In this case, 10.4, 3.0, 1.2, and 2.2 fewer days were required by the Pingluo strain to complete mummy to adult development at 14, 18, 22, and 26°C, respectively. The Pingluo strain had a lower developmental threshold than the Urumqi strain for all three developmental periods, although the difference was not statistically significant for any period (Table 2). Both strains re­ quired 205 degree-days above their respective developmental thresholds to complete development from egg to adult (Table 2). However, a greater num­ ber of degree-days were required by the Pingluo strain to complete egg to mummy development (135 degree-day, SE =0.6) than the Urumqi strain (104 degree-days, SE == 2.5). The Pingluo strain required fewer degree-days to complete mummy to adult development (70 degree-days, SE = 2.2) than the Urumqi strain (101 degree-days, SE = 1.7).

DISCUSSION Degree-day requirements for egg to mummy and mummy to adult devel­ opment differed significantly among strains in this study. More time was spent developing from egg to mummy and less from mummy to adult by the Pingluo strain when compared with the more northern Urumqi strain. Bernal and Gonzalez (1995) suggested that the living host and the mummy may afford different levels of buffering to developing parasitoids against am­ bient temperature as an explanation of why developmental thresholds, de­ gree-day requirements, or both parameters might vary for pre-mummy and post-mummy immature development of a parasitoid. We can speculate that developmental rate is inversely related to the de­ gree of protection from high temperatures afforded during the particular de­ velopmental period. For example, the Pingluo strain originated from a slightly warmer climate than the Urumqi strain and spends a smaller pro­ portion of its total immature development period inside the mummy than the Urumqi strain. This could be because the developing parasitoid is not often 1998 THE GREAT LAKES ENTOMOLOGIST 177

Table 2. Developmental thresholds (SE) and degree-day requirements (SE) for two strains of Aphelinu8 albipodu8. Also listed are regression equations relating develop­ mental rate to temperature were calculated from mean developmental rate at the four temperatures (n 4). Thermal Degree- Regression Place of Origin/Life Stage threshold days equation r2 Pingluo, China egg-mummy formation 8.9 (0.05) 135* (0.6) Y =-O.066+D.0074X 0.99 mummy formation-adult 10.9 (Q.32) 70* (2.2) Y =-0.156+D.0143X 0.83 egg-adult 9.7 (Q.15) 205 (2.8) Y =-0.048+0.0049X 0.96 Urumqi, China egg-mummy formation 8.5 (0.29) 104* (2.5) Y =-0.082+0.0096X 0.89 mummy formation-adult 10.3 (0.17) 101* (1.7) Y =-0.101+0.0099X 0.94 egg-adult 9.6 (0.13) 205 (2.4) Y =-0.047+0.0049X 0.97 Developmental thresholds and degree-days requirements differ significantly (P<0.05) among strains iffollowed by an asterisk. exposed to extreme temperatures while inside the mummy in the cooler cli­ mate, whereas extreme temperatures are more frequently encountered at Pingluo. Thus, the Pingluo strain may have adapted to minimize the time spent in this exposed state. Historical records indicate that the climate at Pingluo is slightly warmer than at Urumqi. For example, average maximum daily temperature during March-May is 17.7°C at Pingluo versus 14.1°C at Urumqi; while average maximum temperatures during June-August are about 1°C higher (29.3°C) at Pingluo than at Urumqi (Hoare 1998). Factors other than climate, for ex­ ample reducing exposure time for parasitism by conspecific or other para­ sitoid species or by hyperparasitoids, could also be responsible for the vari­ able development times of different life stages of these strains. The ecological significance of the variation in developmental threshold and degree-day requirements during portions of the immature development period is unknown to us. However, we can speculate that it is an adaptation by the parasitoid to minimize the time spent in life stages with potentially high mortality subject to constraints on the total time required to complete immature development. In this case the parasitoid would minimize mortality over a fixed generation time, in the respective climates, and as a result in­ crease its fitness. The ability to vary immature development rate in response to climate in new environments could have adaptive significance because it would permit the parasitoid to exploit environments over a broad geographic range. Estimates of developmental thresholds reported for total immature de­ velopment ofA. albipodus were 9.7 and 9.6°C for two geographic regions in China (this study) and 8.2°C for a strain from Tacheng, China (Bernal and Gonzalez 1996). These estimates are substantially greater than those for species ofAphidiidae that parasitize the Russian wheat aphid. For example, estimates of the- developmental threshold for Aphidius matricariae Haliday were 1.4°C for a Czechoslovakian strain (Miller and Gerth 1994) and 4.5 C for a strain from Iraq (Bernal and Gonzalez 1993). For D. rapae, Bernal and Gonzalez (1993) estimated the developmental threshold at 2.1°C for a Pak­ istanian strain, and Elliott et al. (1994) estimated it at 3.5°C for a Syrian strain. Elliott et al. (1995a) observed a developmental threshold of 2.8°C for 178 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4 an Argentinean strain of Aphidius colemani Viereck. Although the develop­ mental threshold differs among aphidiid species and among geographic strains of the same species, all reported values for aphidiids are 1.9Q C or more lower than the lowest estimate reported for A. albipodus. The developmental threshold for A. albipodus is several C higher than that of the Russian wheat aphid (Michels and Behle 1988, Kieckhefer and El­ liott 1989). This indicates that development of A. albipodus populations would lag substantially behind those of the Russian wheat aphid. However, the existence of geographic variation in developmental parameters of A. al­ bipodus suggests that the degree of difference in developmental response to temperature between the host and parasitoid varies regionally. For example, in geographical areas where the temperature transition from cold to warm occurs over a short time interval in the early spring, the time lag between Russian wheat aphid and A. albipodus development would be reduced. Therefore, the comparison of developmental thresholds and degree-day re­ quirements for immature development do not necessarily provide an accu­ rate picture of the degree of synchrony between the parasi toid and host.

ACKNOWLEDGMENTS We thank Wade French, Tim Johnson, and Perry Shelby for technical as­ sistance with the project. We also thank Keith Hopper and Dominique Coutinot (USDA, ARS European Parasite Laboratory) and Larry Ertle, Ken Swan, and Joe Tropp (USDA, ARS Beneficial Insects Research Laboratory, for overseas collection, quarantine, and laboratory rearing of the parasitoids used in this study. This article reports the results of research only. Mention of a proprietary product does not constitute an endorsement or recommenda­ tion for its use by the USDA.

LITERATURE CITED Bernal, J., and D. Gonzalez. 1993. Temperature requirements of four parasites of the Russian wheat aphid Diuraphis noxia. Entomol. Exp. Appl. 69: 173-182. Bernal, J., and D. Gonzalez. 1995. Thermal requirements of Diaeretiella rapae (M'In­ tosh) on Russian wheat aphid (Diuraphis noxia Mordwilko, Hom., Aphididae) hosts. J. Appl. Entomol. 119: 273-277. Bernal, J., and D. Gonzalez. 1996. Thermal requirements of Aphelinus albipodus (Hayat & Fatima) (Hym., Aphelinidae) on Diuraphis noxia (Mordwilko) (Hom., Aphi­ didae) hosts. J. Appl. Entomol. 120: 631-638. Campbell, A., B. D. Frazer, N. Gilbert, A. P. Gutierrez, and M. Mackauer. 1974. Tem­ perature requirements of some aphids and their parasites. J. Appl. Ecol. 11: 431-438. Elliott, N.C., J. D. Burd, J. S. Armstrong, C. B. Walker, D. K. Reed, and F. B. Peairs. 1995a. Release and recovery of imported parasites of the Russian wheat aphid in eastern Colorado. Southwest. Entomol. 20: 125-129. Elliott, N.C., J. D. Burd, S. D. Kinder, and J. H. Lee. 1995b. Temperature effects on de­ velopment of three cereal aphid parasitoids (Hymenoptera: Aphidiidae). Great Lakes Entomol. 28: 199-204. Elliott, N.C., D. K. Reed, B. W. French, and S. D. Kindler. 1994. Aphid host effects on the biology of Diaeretiella rapae. Southwest. Entomol. 19: 279-283. Frazer, B.D., and N. Gilbert. 1976. Coccinellids and aphids. J. Entomol. Soc. British Columbia 73: 33-56. Gilbert, N., A. P. Gutierrez, B. D. Frazer, and R. E Jones. 1976. Ecological relation­ ships. Addison-Wesley, San Francisco. pp. 93-157. 1998 THE GREAT LAKES ENTOMOLOGIST 179

Hoare, R 1998. WorldClimate. World Climate. http://www.worldclimate.com. Jones, J.W., J. R Byers, R A. Butts, and J. L. Harris. 1989. A new pest in Canada: Russian wheat aphid, Diuraphis noxia (Mordwilko) (Homoptera: Aphididae). Can. Entomol. 121: 623-624. Kieckhefer, R.W., and N. C. Elliott. 1989. Effect of fluctuating temperatures on devel­ opment of immature Russian wheat aphid (Homoptera: Aphididae) and demographic statistics. J. Econ. Entomol. 82: 119-122. Messenger, P.S. 1971. Climatic limitations to biological controls. Pages 97-114 in Proc. Tall Timbers Conference on Ecological Animal Control by Habitat Management. Tal­ lahassee, Florida. Messenger, P.S., and R van den Bosch. 1971. The adaptability of introduced biological control agents. Pages 68-92, in: C. B. Huffaker (ed.) Biological Control. Plenum Press, New York. Michels, G.J., and R. W. Behle, 1988. Reproduction and development of Diuraphis noxia (Homoptera: Aphididae) at constant temperatures, J. Econ. Entomol. 81: 1097­ 1101. Miller, J.C., and W. J. Gerth. 1994, Temperature-dependent development of Aphidius matricariae (Hymenoptera: Aphidiidae), as a parasitoid of the Russian wheat aphid. Environ. Entomol. 23: 1304-1307. Morris, RF. 1971. Observed and simulated changes in genetic quality in natural popu­ lations ofHyphantria cunea. Can. Entomol. 103: 893-906. Morris, RF., and W. C. Fulton. 1970. Models for the development and survival of Hy­ phantria cunea in relation to temperature and humidity. Mem. Entomol. Soc. Can. 70: 1-60. SAS Institute. 1988. SAS/STAT users guide. SAS Institute, Cary, N.C. Stoetzel, M.B. 1987. Information on and identification of Diuraphis noxia (Homoptera: Aphididae) and other aphid species colonizing leaves of wheat and barley in the United States. J. Econ. Entomol. 80: 696-704. Tanigoshi, L.K., K. S. Pike, R H. Miller, T.D. Miller, and D. Allison. 1995. Search for, and release of parasitoids for the biological control of Russian wheat aphid in Wash­ ington State (USA). Agric., Ecosyst. and Environ. 52: 25-30. van Emden, J.F. 1988. The potential for managing indigenous natural enemies of aphids on field crops. Phil. Trans. Royal Soc. London B 318: 183-201. 1998 THE GREAT LAKES ENTOMOLOGIST 181

PHENOLOGY AND INFESTATION PATIERNS OF THE COTIONWOOD TWIG BORER (LEPIDOPTERA: TORTRICIDAE) IN IOWA

Joel D. McMillin1,3, MichaelJ. Anderson2, Elizabeth E. Butin2, and Elwood R. Hartl, 2

ABSTRACT Cottonwood twig borer, Gypsonoma haimbachiana (Lepidoptera: Totrici­ dae), phenology and infestation patterns on Populus spp. were examined over a 2-year period in Iowa. Weekly sampling of infested shoots during the host growing season verified the existence of five instars. Head capsule size in­ creased nonlinearly from the first to the fifth instar and corresponded to a concomitant geometric increase in the volume of larval feeding galleries. The sampling indicated that the cottonwood twig borer had two generations per year in Iowa. Corresponding with the two generations, two peaks of larval abundance were observed; one in the second week of June and the other in the first week of August. Greater volume of feeding galleries occurred in the early season generation compared with the late season generation. Sampling of infested shoots revealed that more than 80% of infested terminals con­ tained only one active attack (freshly bored hole in tree terminal with frass present); more than 88% of feeding galleries contained only one larva; and more than 80% of the larvae were found in the first active attack nearest the terminal apex. These data were compared with results published on the phe­ nology and attack patterns of the cottonwood twig borer in the southern United States.

The cottonwood twig borer, Gypsonoma haimbachiana (Kearfott) (Lepi­ doptera: Tortricidae), infests Populus species and hybrids throughout the range of its hosts (Morris et al. 1975). Feeding by larvae causes a shortening of internodal distance and increased forking, breakage, and mortality of ter­ minal and lateral shoots (Solomon 1995). This feeding damage can threaten the economic viability of short-rotation Populus plantations (Morris 1967, Payne et al. 1972). Although the phenology and infestation patterns of the cottonwood twig borer have been reported for short-rotation cottonwood plan­ tations in the southern United States (Morris 1967, Stewart and Payne 1975), similar studies have not been conducted in other regions of its range. Because insect phenology and morphology can vary by geographic location (references cited in Goettel and Philogene 1979), more information is needed to understand the type and timing of damage caused by the cottonwood twig borer before economic injury levels can be determined or management pro­ grams can be developed.

1 Department of Forestry, Iowa State University, Ames, lA, 50011. 2 Department ofEntomoiogy, Iowa State University, Ames, IA, 50011. 3 Current address: USDA Forest Service, Pactola Ranger District, Rapid City, SD 57702 182 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4

The life cycle and phenology of the cottonwood twig borer has been docu­ mented in the southern United States (Morris 1967). Females oviposit small clusters of eggs on the upper surface of leaves. The first instars mine major leaf veins until the first molt and then tunnel into tender shoots at the base of developing leaves. Instars 2 tbrough 5 feed on actively growing tissues in shoots for approximately 21 days. Pupation occurs in bark crevices or in the litter beneath trees. Larvae overwinter in hibernacula on the bark or leaf scars of cottonwood trees (Stewart and Payne 1976). Four to five generations occur per year, with a summer generation developing in 40-45 days in Mis­ sissippi. The population and attack density of cottonwood twig borer can differ significantly by year and location (Payne et al. 1972, Stewart and Payne 1975). Lower levels of attack were found in natural stands compared with plantations, and plantations themselves varied in attack density (Woessner and Payne 1971, Payne et al. 1972). In Texas, nearly 100% of the upper ter­ minal shoots and 50% of the lateral shoots were killed by late August or Sep­ tember (Stewart and Payne 1975). Fifteen-inch terminal shoots in Missis­ sippi were observed to contain up to 25 larvae and three different instars during late summer (Morris 1967). In this paper, we describe the phenology and infestation patterns of the cottonwood twig borer in Iowa, using weekly sampling during the host grow­ ing season over a 2-year period. Through measurements of head capsule width, we established the width range by instar, the volume oflarval feeding gallery by ins tar, and followed the population of each instar through time. Classification of active attacks (freshly bored entrance holes in terminal shoots) were made for evaluating the type of active attack most frequently containing larvae.

MATERIALS AND METHODS Research sites. Observations and measurements of cottonwood twig borer phenology and infestation patterns were conducted during the 1995 growing season (25 May - 19 August) in an agroforestry planting at the Ames Municipal Water Pollution Control Facility near Ames, Iowa. The planting design was alternating 16-m-wide and 365-m-Iong strips of herba­ ceous crops (Panicu8 uirgatum) and of hybrid poplar (Populus deltoides x P. nigra var. 'Eugenei') (Colletti et al. 1994). The hybrid poplars were planted with a 1.3 m x 2.6 m spacing in May 1992. Three plots were selected at ran­ dom within the plantation and each plot was considered one replication, for a total of three replications. Fifteen trees were designated per replication with five trees selected from each ofthe three pairs of rows in each replication. The phenology studies in 1996 (31 May--30 August) were performed at the Institute for Physical Research and Technology (IPERT) near Ames, lA, because the cottonwood twig borer population was too low at the Ames Water Pollution Control Facility site. Ten trees were selected at random in each of four plots within Populus trial plantings that contained 2-year old 'Eugenei' and other Populus clones. Phenology of cottonwood twig borer. Sampling consisted of remov­ ing 15 (1995 season) or 10 (1996 season) infested shoots of actively-growing lateral branches per plot on a weekly basis. During weeks 7 and 8 of 1995 and 5 and 6 of 1996, shoots were not collected because few active attacks could be found. Infested shoots were taken to the laboratory and dissected to collect larvae. Larvae were placed into glass vials of 70% ethyl alcohol to pre­ serve them before measuring head capsule width to determine the range 1998 THE GREAT LAKES ENTOMOLOGIST 183 within each instar. Head capsule width was measured to the nearest 0.01 millimeter at the widest area of the head, using a microscope with an ocular micrometer. Head capsule width data were analyzed using a two-way (sam­ pling date, replicate) ANOVA. To determine the instar and place where larvae overwinter on terminal shoots, 30 shoots were sampled from each of two sites (lPERT and Hinds Re­ search Farm near Ames, IA) in February 1998. Twenty-five-cm-Iong terminal and lateral shoots were collected from trees that had sustained cottonwood twig borer attacks in the 1997 growing season. The shoots were observed in a laboratory for the presence of hibernacula and dormant buds were dissected to find larvae. Volume of larval feeding galleries was measured in 1996 using the same shoots as in the larval phenology study. Only galleries that had one larva were used. Gallery widths and lengths were measured to the nearest 0.01 mm, and derived values were entered into the formula for the volume of a cylinder (7tr2h). Mean volumes of previous instars were subtracted from those of succeeding instars to determine the mean volumes of instars 3 through 5. Volume of feeding gallery data were analyzed using a two-way (sampling date, replicate) ANOVA. Infestation patterns of cottonwood twig borer. The seasonal abun­ dance of cottonwood twig borer larvae was determined by counting the num­ ber of active attacks per tree and per lateral shoot weekly (Stewart and Payne 1975). Fifteen trees in each of three replications were marked and then monitored throughout the 1995 growing season. Because of increasing tree height (2.5 - 3.5 m) and canopy complexity after the seventh week of sampling, only 10 of the 15 trees per replicate were sampled during the re­ mainder of the growing season. Seasonal abundance, as determined by the number of active attacks per selected tree, was not recorded in 1996. The number-of-active-hits data were analyzed using a three-way (sampling date, gen,eration, replicate) ANOVA. Beginning 29 July 1995, active attacks were classified further by record­ ing the following designations: (1) presence or absence of frass at the en­ trance hole of an active attack, and (2) location of active attacks (terminal growth node, growth node, or internode). In 1996, active attacks were classi­ fied as in 1995; however, attacks by first instars in the midveins of leaves also were examined. When active attack counts are recorded, a similar type of damage caused by the poplar branch borer, Oberea schaumii (LeConte) (Coleoptera: Ceram­ bycidae) must be subtracted from the active attacks. Poplar branch borer holes are distinct from cottonwood twig borer active attacks because they can be found at the bases of petioles with scarring on the outside of the entrance hole, and with wood shavings instead of red frass.

RESULTS Phenology of cottonwood twig borer. Similar to Morris' (1967) find­ ings, five instars were detected in Iowa (Table 1). First instars had head cap­ sules that varied in width from 0.16 to 0.22 mm compared with Morris (1967) who found a very nearly constant head capsule width of 0.19 mm. In addi­ tion, the range of head capsule widths for filth instars was larger during both years of the study than in Mississippi. In Iowa, more variation (greater range) in head capsule width occurred during the fourth and fifth stadia than during the second and third stadia. This greater variation corresponded 00 .J>.

Table 1. Head width of cottonwood twig borer larvae in Mississippi and Iowa and corresponding volume Head Capsule Width (mm) Gallery Mississippi' Iowa 1995 Iowa 1995 Iowa 1996 Iowa 1996 volume (mm3) :tm lnstar (n) Mean (SE) Range (n) Mean (S};~) Range (n) 1996 (SE) Q ;;0 m 1 0.19 (87) 0.19 (0.01) 0.16-0.22 (2) 0.20 (O.02) 0.16-0.22 (2) ~ };: A m 2 0.26-0.32 (172) 0.31 (0.03) 0.26-0.35 (7) 0.32 (0.02) 0.28-0.35 (8) 4.13 (1.39) (J> m Z 3 0.38-0.45 (242) 0.43 (0.03) 0.37-0.47 (36) 0.43 (0.03) 0.37-0.47 (40) 10.56 (2.28) o ?;: o 0.51-0.83 (988) 0.68 (0.09) 0.49-0.81 (147) 0.67 (0.09) 0.48-0.80 (133) 34.86 (3.97) 5 4 Q ~ 5 0.90-1.15 (604) 0.98 (0.09) 0.82-1.18 (132) 0.97 (0.09) 0.81-1.18 (154) 68.37 (6.21) *Data reported by Morris (1967). ~ w

oz w Qo .J>.

__ i .------~.'I 1998 THE GREAT LAKES ENTOMOLOGIST 185

12 1995 Q) 10 ca >... 8 .1st .!! III 2nd -0... 6 Ei3rd Q) .Q o 4th E 4 • 5th ::J Z 2 [ 0 ~ II r I I I r I _1_ _...I[,2'-----"3~-.!!4'-- ....5'--~6'--___!7_~8~ 9 10 11 12 May June July August Week (sampling began May 25)

25 1996

Q) 20 ca ...> .!! 15 -0... Q) .Q 10 E ::J Z 5 j 0 ~, ~. r f1I III I I R ~ ~ ..J 2 3 4 5 6 7 8 9 10 11 12 13 May June July August Week (sampling began May 31) Figure 1. Instar frequency of cottonwood twig borer by sampling date during the 1995 and 1996 growing seasons.

with the period of greatest increase in head capsule size of the developing larvae. The mean width of head capsules varied by sampling date in both years (P < 0.001). Based on the measurements and frequencies of head capsule widths, there were two generations per year in Iowa (Fig. 1). The first larval 186 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No.3 &4

generation was completed by the end of June and the second larval genera­ tion occurred from July through August. Because a majority oflarvae during week 1 were already in the fourth stadium (Fig. 1), second or third instars probably overwinter as in the southern United States (Stewart and Payne 1976). However, an inspection of shoots in February 1998 failed to find any hibernacula or larvae in dormant buds. Only one second instar was found and it occurred inside a hollowed area of the shoot, 20 cm below the terminal bud. The volume of feeding gallery varied by instar (F ratio = 14.5, P < 0.001, df 3). Regression analysis revealed a significant curvilinear relationship be­ tween the volume of feeding gallery (y) and head capsule width (x) (y 4.08x2.78, d.f. = 236, F ratio = 200.3, P < 0.001, r2 = 0.58, Table 1). Mean gallery volume also varied by sampling period in 1996 (F ratio = 6.8, P < 0.001, df == 10) with a greater volume of plant tissue being consumed during the early season generation (Fig. 2). Infestation patterns ofcottonwood twig borer. Corresponding to the two generations per year, monitoring of the previously marked trees during the 1995 growing season showed two peaks of active attacks (Fig. 3). Signifi­ cant effects of sampling date, generation, and replication on the number of active hits were detected (Table 2). The mean number of active attacks was larger in generation 1 (4.76 ± 0.27 SE) than in generation 2 (2.90 ± 0.23). The peak for generation 1 occurred during week 3 and the peak for generation 2 occurred during week 11. A significant linear relationship was found between the number of active attacks (y) and the number of actively growing shoots

120

OJ -E 100 E CI) 80 -E :::J "6 60 > ~ 40 oS? "i 20 " 0 ...l _-=----'3"---""4_2 5 6 7 8 9 10 11 12 13 May June July August Week (sampling began May 31)

Figure 2. Feeding gallery volume (mm3) of cottonwood twig borer larvae by sampling date in 1996. Bars represent mean + SE of the mean. 1998 THE GREAT lAKES ENTOMOLOGIST 187

Table 2. Three-way ANOVA (sampling date x generation x replication) of the number ofcottonwood twig borer active attacks per tree during the 1995 growing season. Source of variation d.f. F ratio Pvalue SampIing date 11 13.5 < 0.001 Generation 1 28.4 <0.001 Replicate 2 8.0 < 0.001

10 ~ .... 8 -l ~ 6 (,) CIS -CIS 4 ~ tj 2

1 ~2__~3~_4~_5~~6__~7__~8~_9~__~10~~1~1__1~2~~13 May June July August Week (sampling began May 26)

Figure 3. Number of cottonwood twig borer attacks per tree by sampling date during the 1995 growing season. Bars represent mean + SE of the mean.

(x) during week 7 in 1995 (y = -0.40 + 0.17x, d.f. = 43, F ratio 23.50, P < 0.001, r2 = 0.31). The significant difference in the number of active attacks among replicates was caused, in part, by a greater number of actively grow­ ing shoots in one of the three replicates. In general, only one active attack was observed per infested shoot (Table 3). More than 80% of infested shoots contained only one active attack and less than 2% of infested shoots contained three or more active attacks. For shoots that had more than one attack, the percentage of active attacks that contained actively feeding larvae increased toward the apex of a shoot. Of the three active attacks closest to the terminal apex, feeding larvae were found in 97.1% of the first active attack, 2.6% in the second active attack, and 0.3% in the third. Greater than 90% of active attacks had frass at the entrance hole of the twig-boring larvae. Furthermore, more than 85% of active attacks housed only one larva per feeding gallery (Table 3). 188 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4

Table 3. Frequency of active attacks per infested shoot for generation 2, 1995 and the frequency of cottonwood twig borer larvae per feeding gallery in 1995 and 1996. No. of larvae No. of active attacks Frequency per feeding Frequency (% SE) per infested shoot (%SE) gallery 1995 1996 1 83.1 (2.0) 1 93.6 (4.0) 88.2 (1.4) 2 12.9 (1.7) 2 5.4 (3.0) 11.1 (1.4) 3 2.1 (0.7) 3 1.0 (1.0) 0.7 (004) 4 0.9 (0.1) 5 0.2 (0.2) 6 0.5 (O.5)

DISCUSSION Phenology of cottonwood twig borer. The results of this study were consistent with Morris' (1967) measurements of head capsule widths and number of instars, suggesting that there is not much geographical variation in body size of cottonwood twig borer larvae. One plausible explanation for the greater range in the size of fifth instar head capsules in Iowa compared with Mississippi is the possibility that an additional instar may occur for fe­ male cottonwood twig borer. Although Morris (1967) did not report any evi­ dence of an additional instar for females, this is a common phenomenon for many insects (Coulson and Witter 1984). More work is needed to determine the occurrence of sexual dimorphism during the larval stage or the cause of the greater range in body size of larvae in Iowa Two generations of cottonwood twig borer occur annually in central Iowa based on the weekly sampling during 1995 and 1996. The shorter growing season in Iowa results in fewer generations per year compared with the southern United States. Week 1 in 1995 and week 3 in 1996 were the peaks or the start of decline of the fourth instar from the overwintering generation; therefore, future observations should commence in early May. During the in­ tergenerational period (weeks 7 and 8 of 1995 and weeks 5 and 6 of 1996), it was difficult to locate cottonwood twig borer larvae because first instars are in the midvein of the leaves. The peak of fourth and fifth ins tars by the third week of August suggests that the generation would be completed before the average first frost (September 22). After this generation deposits eggs and larval eclosion occurs, the larvae probably begin to feed and molt in leaf mid­ veins before moving to an overwintering location as second instars. The ab­ sence of any hibernacula in Iowa suggests that, because of the colder winters in Iowa compared with Mississippi, cottonwood twig borer may have evolved to overwinter inside terminal shoots or possibly in the leaf litter around the base of trees on the ground. The mean gallery volume increased in a curvilinear fashion from the sec­ ond instar to the fifth instar demonstrating that the late instars consume proportionately more than the early instars. The volume consumed increased from weeks 1 to 3 as the mean stage of instar increased. During the second generation, volume of feeding galleries was considerably less than during the first generation. This difference may have been caused by the first genera­ tion larvae having a longer feeding period and/or because of the impact of grasshopper damage during the second generation. In both years, larval feeding during the second generation was affected by damage to actively growing shoots from grasshopper feeding (Melanoplus femurrubrum DeGeer ===------=------======------~"'..

1998 THE GREAT LAKES ENTOMOLOGIST 189

and Melanoplus differentialis Thomas) (Orthoptera: Acrididae). Large popu­ lations of grasshoppers appeared in mid-July and were observed feeding on actively lVowing shoots. This grasshopper-caused damage most probably had a direct Impact on twig borer populations by killing larvae that were feeding inside the shoots or an indirect impact by reducing their food resources (e.g. fewer actively growing terminals). In addition, possible differences in the nu­ tritional value of shoots throughout the growing season may have influenced the feeding of larvae. Infestation patterns of cottonwood twig borer. The results of this study suggest that lower densities of cottonwood twig borer are found in cen­ tral Iowa than in the southern United States. In 1995, only one active attack per infested shoot was found, and, in general, only one larva per feeding gallery was found in 1995 and 1996. These findings are in contrast to the southern United States where a 15-inch terminal contained up to 25 larvae and three different instars during late summer (Morris 1967). These differ­ ences between geographic regions were probably caused by the greater num­ ber of generations per year and a longer history of cottonwood plantations in the south. In addition, Lorsban® was applied in July 1995 for controlling grasshopper populations and may have caused a direct impact on the larvae or on the natural enemy complex (Woessner and Payne 1971). The combina­ tion of this insecticide application with the grasshopper-caused damage to ac­ tively growing shoots probably resulted in the observed lower number of at­ tacks per shoot in the second generation. The positive correlation between active attacks and the number of actively growing shoots indicates that fac­ tors leading to increases or decreases in the number of actively growing ter­ minals affects cottonwood twig borer populations.

ACKNOWLEDGMENTS We thank Richard B. Hall for project leadership, Rosemarie A. Anderson, Ying Fang, Chris Feeley, Christine Hall, Roger D. Hanna, Jennifer Jarrard, and Sisi Lin for sampling assistance. The research was supported by the Oak Ridge National Laboratory, operated by Martin Marietta Energy Systems, Inc. for the U.S. Department of Energy, contract no. DE-AC05-840R21400. Journal Paper No. J-17855 of the Iowa Agriculture and Home Economics Ex­ periment Station, Ames, Iowa Project No. 2976, 3088, and 3443, and sup­ ported by Hatch Act and State of Iowa funds.

LITERATURE CITED Colletti, J. P., and eleven additional authors. 1994. An alley cropping biofuels system: operation and economics. pp. 303--310. In: Proc. Third North Am. Agrofor. Con. (J. P. Colletti and R. C. Schultz, eds.). Iowa State University, Ames, Iowa. Coulson, R. N. and J. A Witter. 1984. Forest Entomology. Wiley-Interscience, New York. G1>ettel, M. S. and B. J. R. Philog?me. 1979. Further studies on the biology of pyrrharc­ tia (isia) isabella (Lepidoptera: Arctiideael. III. The relation between head capsule width and number of instars. Can. Entomol. 111: 323-326. Morris, R. C. 1967. Biology of Gypsonoma haimbachiana (Lepidoptera: Olethreutidae), a twig borer in eastern cottonwood. Ann. Entomol. Soc. Am. 60: 423-427. Morris, R. C., T. H. Filer, J. D. Solomon, and others. 1975. Insects and diseases of cot­ tonwood. USDA For. Ser. Gen. Tech. Rep. SO-8. 37 p. Payne, T. L., R. A Woessner, and V. C. Mastro. 1972. Evaluation of cottonwood clonal 190 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4

selections for resistance to cottonwood twig borer attack. J. Econ. Entomol. 65: 1178-9. Solomon, J. D. 1995. Guide to insect borers of North American broadleaf trees and shrubs. USDA For. Ser. Agric. Handb. 706. 735 p. Stewart, J. W. and T. L. Payne. 1975. Seasonal abundance and impact of the cotton­ wood twig borer on cottonwood trees. J. Econ. Entomol. 68: 599-602. Stewart, J. W. and T. L. Payne. 1976. Overwintering habits and winter mortality of the cottonwood twig borer. Southwest. Entomol. 1: 9-12. Woessner, R. A and T. L. Payne. 1971. An assessment ofthe cottonwood twig-borer at­ tacks. Southern Conf. Forest Tree Impr. Proc. 11: 98-107. 1998 THE GREAT LAKES ENTOMOLOGIST 191

THE SEQUENTIAL RELATIONSHIP OF BODY OSCilLATIONS IN THE PAPER WASP, POLISTES FUSCATUS (HYMENOPTERA: VESPIDAE)

Bobbi J. Harding 1 and George J. Gamboa 1,2

ABSTRACT Three kinds of body oscillations by foundresses of the paper wasp, Polistes fuscatus, were analyzed from 100 h of videotapes of 17 multiple- and 16 single-foundress, preworker colonies. The three kinds of oscillations were observed to be temporally proximate only after prey returns. Their sequen­ tial occurrence was always antennal drumming (AD), abdominal wagging (AW), and lateral vibration (LV). This sequence is consistent with the hypoth­ esized communicative meanings of ADs, AWs, and LVs. In particular, ADs may signal larvae to withold salivary secretions prior to receiving a liquid meal from an adult female; oscillations AW and LV may signal larvae to se­ crete and withold saliva, respectively, Additional studies are required to pro­ vide causal evidence of the communicative meanings of ADs, AWs, and LVs.

Body oscillations have been reported in adult females of at least 15 species of wasps, and thus appear to be widespread among social wasps. The communicative meaning of these oscillations, however, is poorly understood (Savoyard et al. 1998), The species of social wasp whose oscillatory behavior has been studied most extensively is the paper wasp, Polistes fuscatus (Fabr.). Three distinct kinds of oscillations have been described in P. fuscatus: antennal drumming (AD), abdominal wagging (AW) and lateral vi­ bration (LV). ADs consist of a female positioning her head over the opening of a cell containing a larva and rapidly hitting her antennae against the rim of the cell (Pratte and Jeanne 1984). AWs consist of slow (2-7 oscillations/s), side-to-side movements of a female's gaster against the nest as she walks over brood cells (Gamboa and Dew 1981). LVs consist of a stationary female rapidly (17 oscillations/s) moving her abdomen from side-to-side against the surface of the nest producing a short «Is), audible sound (Savoyard et al. 1998). All three types of oscillations are thought to be signals by adult females to larvae (Gamboa and Dew 1981, Pratte and Jeanne 1984, Downing and Jeanne 1985, Savoyard et al. 1998). The behavioral repertoire of larvae and the kinds of signals from female adults to larvae are thought to be quite lim­ ited (Savoyard et al. 1998), Larvae receive malaxated prey and liquid food from adults and, in turn, provide saliva to adults (West-Eberhard 1969, Pratte and Jeanne 1984, Hunt 1991). Adult females may signal larvae to se­ crete or withhold saliva (Savoyard et al. 1998). Larval saliva contains sugars,

lDepartment of Biological Sciences, Oakland University, Rochester, MI 48309. 2Corresponding author. 192 THE GREAT lAKES ENTOMOLOGIST Vol. 31, No. 3 &4 amino acids, and proteins (Hunt et al. 1982) and has been postulated to play an important role in the evolution of eusociality (Hunt 1991). Although ADs, AWs, and LVs are presumed to be signals by females to larvae, their specific communicative meanings are not well understood. Savo­ yard et al. (1998) reported that the three kinds of oscillations are sometimes associated with each other. More specifically, Savoyard et al. (1998) docu­ mented that the numbers ofLVs were significantly, positively correlated with both the numbers of ADs and AWs. We examined these three kinds of oscilla­ tions to determine if there is a specific temporal and sequential relationship among them. Such information might provide additional insight into their communicative meanings.

METHODS We videotaped 17 multiple- and 16 single-foundress, preworker colonies of P. fuscatus nesting in plywood nestboxes near Rochester, Michigan. The nestboxes were suspended from wood crossbars attached to metal fenceposts. Foundresses were marked for individual identification on 17 and 19 June 1997, approximately one week prior to the beginning of videotaping. Each of five Sony TR910 Hi8 video cameras was positioned on a tripod approximately 0.9m beneath a nestbox so that the field of view consisted of the wasps on the face and side of the comb. The video, filmed at 30 frames per second with a 15X optical telephoto, provided a field of -7cm in diameter. We videotaped 17 multiple-foundress colonies from 22-27 June and 16 single-foundress colonies from 27-29 June. We completed all videotaping between -1000 h and -1730 h when ambient air temperatures were ~21°C. We completed 4 h of videotaping for each multiple-foundress colony (68 h) and 2 h of videotap­ ing for each single-foundress colony (32 h). Videotapes were viewed in a laboratory and the behavior of foundresses was recorded on a microcassette audio recorder. We recorded departures, re­ turns, items foraged, body oscillations, and their associated times for all foundresses. To increase observer reliability and to minimize observer bias, one individual (GJG) observed all videotapes and completed all transcrip­ tions.

RESULTS Pratte and Jeanne (1984) reported that ADs occurred only after wasps re­ turned with prey. Thus we examined the sequence of oscillations that fol­ lowed prey returns since this was the only context in which we observed all three kinds of oscillations. In single-foundress colonies, we recorded nine prey returns of which six were followed within 30 min by all three kinds of body oscillations. In multiple-foundress colonies, we recorded 80 prey returns of which eight were followed within 30 min by all three kinds of body oscilla­ tions. Fifteen of these oscillation sequences were interrupted by the returns of other foundresses. This may at least partly explain why single-foundress colonies had a higher proportion of prey returns that were followed by all three oscillations than did multiple-foundress colonies. When there was more than one complete sequence of oscillations for a colony, we averaged the values from multiple sequences so that each colony contributed a single datum for calculating a mean value. This resulted in a sample size of five single- and six multiple-foundress colonies for which we had at least one complete sequence of three different oscillations. We found 1998 THE GREAT LAKES ENTOMOLOGIST 193

Prey Return

ADs AD. CHEWING FEEDING INACTIVE INSPECTING AWs LVls) - • . I I I I I I I o 150 300 450 600 750 900 1050 1200

Time(s)

Figure 1. The sequence of activities performed by foundresses of Polistes {us­ catus after returning with a prey item. The seconds spent in each activity are mean values based on five single- and six multiple-foundress colonies. Prey returns and LVs are the initial and terminal events, respectively, of the se­ quence of activities.

no significant differences between single- and multiple-foundress colonies in the nine parameters discussed below (p ;::: 0.09; Mann-Whitney U Test with a Bonferroni Correction). Upon returning with a prey item, the wasp chewed the prey item (x = 312s, range = 150-735s), which became noticeably smaller as the wasp ex­ tracted fluids from the prey (Fig. 1). Female Polistes extract hemolymph from prey as they malaxate it (Hunt 1984). The wasp then distributed the prey mass to larvae (x", 51s, range = 5-100s). After feeding the prey to larvae, the wasp began performing ADs (x", 12.9 ADs, range 2-20ADs) while display­ ing inactivity between episodes ofADs (x =205s, range = 30-470s). The wasp then began to inspect cells between episodes of ADs (x = 8.6 ADs, range =: 1-29 ADs; x= 139s, range 30-405s). During cell inspections, adult wasps lowered their heads into cells containing larvae. Next, the female performed AWs (x =: 2.3 episodes of AWs, range 1-6 episodes) as she inspected and walked over cells (x = 207s, range = 5-660s). Finally, the wasp performed one to five LVs (x 2.0 LVs), which were the terminal oscillations of the se­ quence.

DISCUSSION The sequence of the three types of oscillations was invariant: for all 14 recorded sequences, the order of the oscillations was AD, AW, and LV. This se­ quence is consistent with the suggested communicative meanings hypothe­ sized for ADs, AWs, and LVs from previous studies. Pratte and Jeanne (1984) provided evidence that ADs follow the feeding of solid prey to larvae. They postulated that ADs inform larvae that they are about to receive a liquid meal from an adult and that the larvae should withhold their salivary secre­ tions. Our observations that ADs occurred after larvae were fed prey and that wasps extracted liquid from prey items shortly before performing ADs is consistent with Pratte and Jeanne's (1984) hypothesized function of ADs. Savoyard et al. (1998) speculated that AWs and LVs might signal larvae to secrete and to withhold larval salivary secretions, respectively. Our findings 194 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No.3 & 4

thatAWs always preceded LVs and that LVs were the terminal oscillations of the sequence are consistent with the hypotheses that AWs stimulate larval secretions and LVs inhibit larval secretions. It has been hypothesized that much of the adult-larval communication in social wasps is mediated by the substrate vibrations produced by body oscil­ lations (Savoyard et al. 1998). Since the durations and oscillation frequencies of ADs, AWs, and LVs are known (Pratte and Jeanne 1984; Savoyard et al. 1998), it should be possible to mechanically reproduce the three kinds of body oscillations in combs containing larvae. This, in turn, could be utilized in ex­ perimental studies to provide causal evidence of the communicative mean­ ings of ADs, AWs, and LVs.

LITERATURE CITED Downing, H.A. and R.L. Jeanne. 1985. Communication of status in the social wasp, Polistes fuscatus (Hymenoptera: Vespidae). Z. Tierpsychol. 67: 78-96. Gamboa, G.J. and H.E. Dew. 1981. Intracolonial communication by body oscillations in the paper wasp, Polistes metricus. Insectes Sociaux 28; 13-26. Hunt, J.H. 1984. Adult nourishment during larval provisioning in a primitively euso­ cial wasp, Polistes metricus Say. Insectes Sociaux 31: 452-460. Hunt, J.H. 1991. Nourishment and the evolution of the social Vespidae. pp. 426-450. In; KG. Ross and RW. Matthews (eds.). The Social Biology of Wasps. Cornell Univ. Press, Ithaca, New York. Hunt, J .H., I. Baker and H.G. Baker. 1982. Similarity ofamino acids in nectar and lar­ val saliva; the nutritional basis for trophallaxis in social wasps. Evolution 36: 1318-1322. Pratte, M. and RL. Jeanne; 1984. Antennal drumming behavior in Polistes wasps (Hy­ menoptera: Vespidae). Z. Tierpsychol. 66: 177-188. Savoyard, J.L., G.J. Gamboa, D.L.D. Cummings and RL. Foster. 1998. The commu­ nicative meaning of body oscillations in the social wasp, Polistes fuscatus (Hy­ menoptera: Vespidae). Insectes Sociaux 45; 215-230. West-Eberhard, M.J. The social biology of polistine wasps. Misc. Publ. Mus. Zool. Univ. of Mich. 140; 1-101. 1998 THE GREAT LAKES ENTOMOLOGIST 195

ADDITIONAL SIPHONAPTERA RECORDS FROM SMALL MAMMALS IN THE CENTRAL UPPER PENINSULA OF MICHIGAN

William C. Scharfl and Patrick E. Lederle2

ABSTRACT Fleas were collected from mammals during the period 1990-1992 in two upper peninsula counties. Identified specimens were compared to existing distribution records for both parasite and host. Only those records which are newly documented for county, upper peninsula or Michigan are listed. We re­ port one previously unknown flea species and five new host records for Michigan. One host record is new for the upper peninsula. In addition, seven new host/parasite combinations are recorded for two central upper peninsula counties.

Six previous studies document flea and host species distributions in the upper peninsula of Michigan and islands of the Great Lakes near the upper peninsula shoreline (Lawrence et al. 1965, Wilson and Johnson 1971, Scharf and Stewart 1980, Scharf et al. 1990, Scharf 1991, and Scharf in press). Additional comparisons of distributions were made with Timm (1975) for northern Minnesota and Benton (1980) for the northeastern United States. Despite the semblance of thorough coverage by previous studies, much of the large geographic area of the upper peninsula of Michi­ gan' and a variety of flea hosts remain unexamined. The goal of this paper is to further the knowledge of flea and host species distributions with new records from the region.

MATERIALS AND METHODS We collected 143 fleas of 12 species from 9 mammal host species in Dick­ inson and Iron Counties between 1990 and 1992. Some flea species were not new records, and are not listed here. The majority of the specimens were col­ lected by systematic trapping using Leathers live traps by PEL. He and as­ sistants collected regularly as part of long term small mammal studies in Dickinson and Iron Counties conducted by the Michigan State University Terrestrial Vertebrates Group (1986-1992). The live trapping was performed at two sites. The Pirlot Road site (T43N R29W, Sections 14, 23, and 26) was located in Dickinson County, approxi­

lEcological Inventory, 760 Kingston, Traverse City, MI 49684, email: wscharf@ traverse.com. 2SAIC, 1271 Town Center Dr., MJS 423, Las Vegas, NV 89134, email: patrick_ [email protected]. 196 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No.3 &4 mately 13 kIn east of Sagola, MI. The Michigamme site (T44N R31W, Sec­ tions 24 and 25) was located in Iron County, approximately 6 kIn northwest of Channing, MI. Previous collections of fleas from these sites in prior years are reported in Scharf et al. (1990). Small mammals were brushed and examined, and fleas were picked up with forceps or an alcohol wetted finger. All fleas were stored in 70% ethyl alcohol, and later cleared and mounted in Canada Balsam on glass slides for microscopic identification by WCS. Most Siphonaptera were identified using Holland (1949), but Benton (1983) was used in some instances. Hol­ land's nomenclature is followed except that Nearctopsylla genalis genalis (Baker) follows the accepted nomenclature of Hopkins and Rothschild (1953). The slides are in the personal collection of WCS with voucher speci­ mens of all species deposited in the University of Minnesota Entomology Collection. Mammal hosts were identified by PEL. Mammal names are according to Hamilton and Whitaker (1979), but also reflect nomenclatural revisions mandated by Jones et al. (1992).

RESULTS The following list by parasite family and species consists minimally of new county records for either host or parasite. New records for the mainland of the upper peninsula, and new records for Michigan are noted. Additional specimens that were reported in earlier works are not repeated. Following each county name, is the distribution of the sexes of the flea, the number of hosts from which the flea was collected (if it exceeded one) and collection dates to indicate phenology and temporal abundance.

CERATOPHYLLIDAE

Megabothris acerbus (Jordan) - Eastern Chipmunk, Tamias striatus (L.), Dickinson Co., 5 males, 7 females from 5 hosts 8-29 May 1991; Woodland Deer Mouse, Peromyscus maniculatus gracilis LeConte NEW RECORD FOR MICHIGAN, Iron Co., 1 male, 26 May 1991. Opisdasys pseudarctomys (Baker) - Southern Flying Squirrel, Glaucomys volans (L.), Dickinson Co., NEW RECORD FOR MICHIGAN, 5 males, 13 fe­ males, 20 February 1992. Orchopeas caedens caedens (Jordan) - G. volans, Dickinson Co. NEW RECORD FOR MICHIGAN, 1 Male, 20 February 1992. Orchopeas leucopus (Baker) - Pygmy Shrew, Sorex hoyi Baird, NEW RECORD FOR MICHIGAN, Dickinson Co. , 1 male, 3 March 1992.

LEPTOPSYLLILDAE Peromyscopsylla h. hesperomys (Baker) - Peromyscus maniculatus gra­ cilis, Dickinson Co., NEW RECORD FOR UPPER PENINSULA, 1 female, 17 June 1991. 1998 THE GREAT LAKES ENTOMOLOGIST 197

HISTRICHOPSYLLIDAE

Ctenophthalmus pseudagyrtes Baker - Masked Shrew, Sorex cinereus Kerr, Dickinson Co., 1 male, 18 February 1992; Tamias striatus" Dickinson Co. 1 male, 28 May 1991; Red Squirrel, Tamiasciurus hudsonicus (Erxleben), Iron Co., 1 male, 21 February 1990. Epitedia wenmanni (Rothschild) - Peromyscus maniculatus gracilis, Dickinson Co., 1 male, 6 February 1990; Tamiasciurus hudsonicus, Dickin­ son Co., 3 males, 18 February 1990, 1 male, 21 February 1990. Nearctopsylla genalis genalis (Baker) - Sorex hoyi, Dickinson Co., NEW RECORD FOR MICHIGAN, 1 female, 3 March 1992. Tamiophila grandis (Rothschild) - Tamias striatus, Dickinson Co., 2 fe­ males, 1 from one host 29 May 1992, and 1 from another host 20 July 1992.

DISCUSSION Both species of flying squirrels, Glaucomys, are considered to be true hosts of Opisdasys pseudarctomys (Holland 1949) and it is remarkable that this flea has not been collected previously from the center of both hosts' dis­ tribution in the northern lower peninsula (Baker 1983, Scharf and Stewart 1980, Scharf et al. 1990). G. volans has previously been collected only four times in the upper peninsula (Baker 1983), but we have each recorded its presence there independently from several sites prior and subsequent to this study. Evidence supporting a sporadic distribution of the two flying squirrel species in the upper peninsula is demonstrated by our capture of G. sabrinus only at the Michigamme site (fleas reported in Scharf et al. 1990) and G. volans only at the Pirlot Road site (fleas reported in this paper) during the entire seven years of trapping. The other new flea species from G. volans, Or­ chopeas caedens caedens, is common on other species of cavity nesting Sciuri­ dae, especially the red squirrel, in many areas of the state (Lawrence et al. 1965, Wilson and Johnson 1971, Scharf 1980, Scharfet al 1990). The two new state records from the pygmy shrew, Sorex hoYi, include the flea Nearctopsylla gena lis genalis, which exhibits host specificity for the genus Sorex and the more ubiquitous flea Orchopeas leucopus, whose true hosts appear to be mice of the genus Peromyscus (Holland 1949). In the latter case, it seems that collecting and identifying the host in the central upper peninsula was instrumental to the discovery of its parasites. The new record of Megabothiris acerbus from Peromyscus maniculatus gracilis is unremark­ able because the true host of this flea appears to be Tamias striatus, and the two mammal species undoubtedly share the same nesting habitat occasion­ ally.

ACKNOWLEDGMENTS We thank the many project assistants from Michigan State University that helped collect fleas. MSU Institutional Animal Care and Use Permit AUF #11/87-464-04, and Michigan Department of Natural Resources Collect­ ing Permits to both authors allowed the collecting described here. This work was supported, in part, by the ELF Communications System Ecological Mon­ itoring Program, Michigan State University, Subcontract D06205-93-C-006, 198 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4 administered by Donald L. Beaver. The senior author thanks Orner R. Lar­ son for early encouragement in the study of Siphonaptera.

LITERATURE CITED Baker, R. H. 1983. Michigan Mammals. Michigan State University Press, East Lans­ ing, MI. xx + 642 pp. Benton, A. H. 1980. An atlas of the fleas of the eastern United States. Marginal Media, Fredonia, NY. xv + 177 pp. Benton, A. H. 1983. An illustrated key to the fleas of the eastern United States. Biogu­ ide No.3 Marginal Media, Fredonia, NY. iv + 34 pp. Hamilton, W. J. and J. O. Whitaker. 1979. Mammals of the Eastern United States. 2nd ed. Cornell Univ. Press, Ithaca, NY. 358 pp. Holland, G. P. 1949. The Siphonaptera of Canada. Canadian Dept Agric. Bull. 70. 358 pp. Hopkins, G. H. E. and M. Rothschild. 1953. Catalogue of the Rothschild Collection of fleas. VoL 1 Tungidae and Pulicidae. Cambridge Univ. Press. 358 pp. Jones, J. K., R. S. Hoffmann, D. W. Rice, C. Jones, R. J. Baker, and M. D. Engstrom. 1992. Revised Checklist of North American Mammals North of Mexico, 1991. Occa· sional Papers, The Museum Texas Tech University 146:1-23. Lawrence, W. H., K. L. Hays, and S. A. Graham. 1965. Arthropodous ectoparasites of some northern Michigan mammals. Occas. Papers Mus. Zool. Univ. Michigan. 639: 1-7. Scharf, W. C. 1991. Geographic distribution of Siphonaptera collected from small mam­ mals on Lake Michigan islands. Great Lakes Entomol. 24: 39-43. Scharf, W. C. (in press) Siphonaptera from migrating owls: passengers on the journey. Michigan Birds and Natural History. Scharf, W. C. and K. R. Stewart. 1980. New records of Siphonaptera from northern Michigan. Great Lakes Entomol. 13:165-167. Scharf, W. C., P. E. Lederle, and T. A. Allan. 1990. Siphonaptera from the central and eastern upper peninsula of Michigan. Great Lakes Entomol. 23: 201-203. Timm, R. M. 1975. Distribution, natural history and parasites of mammals of Cook county, Minnesota. Occas. Papers, Bell Mus. Nat. Hist., 14: 1-56. Wilson, N. and W. J. Johnson. 1971. Ectoparasites ofIsle Royale, Michigan. The Michi· gan Entomol. 4: 109-115. 1998 THE GREAT LAKES ENTOMOLOGIST 199

PSILOTRETA INDECISA AND AGARODES DISTINCTUS (TRICHOPTERA: ODONTOCERIDAE, SERICOSTOMATIDAE): NEW STATE RECORDS AND NOTES ON THE HABITAT OF THESE SPECIES IN MICHIGAN

Ethan Bright! and Doug Bidlack2

ABSTRACT The caddisfly species Psilotreta indecisa and Agarodes distinctus are re­ ported for the first time in the State of Michigan. These species appear to be widely distributed in the state based upon their collection localities. Habitat information from collection localities is given.

Psilotreta is an infrequently collected odontocerid caddisfly in which the larvae are known for constructing an incredibly strong case and often aggre­ gating in large numbers when attaching their cases to rocks for pupation (Parker and Wiggins 1987). The genus ranges from Wisconsin and southern Hudson Bay in Ontario east to Quebec and Nova Scotia, and south along the Appalachian Mountains to Tennessee, Georgia and South Carolina. Several records also are from the Coastal Plain in North Carolina, South Carolina, and northern Florida. However, no records have been published from Michigan, where one would expect to find it based on the known range of this genus. Larval and adult odontocerid caddisflies were collected in 1997 from the Huron River at Indian Springs Metropark (T3N, R8E, Sec. 3), Oakland County, Michigan, and identified as Psilotreta indecisa (Walker). We then rechecked the UMMZ-Insect Division Trichoptera collection for additional odontocerid specimens. Two larval specimens collected by S. Lievense on 18 April 1949 from the Escanaba River (T41N, R23W, Sec. 7), Delta County were found in the collection. J. W. and F. A. Leonard, who curated the speci­ mens, were not able to identify these larvae to species because larvae had not been associated with adults at that time. The Leonards probably did not in­ clude the genus in their annotated list of the Trichoptera of Michigan (1949) for this reason. The genus Psilotreta has since been revised by Parker and Wiggins (1987), who published a detailed systematic account of the genus. The specimens from the Escanaba River are undoubtedly P. indecisa and should therefore be considered the first record for the state of Michigan. Four additional larvae were collected from the Little Siskiwit River, Isle Royale (T64N, R37W, Sec. 24) on 23 May 1997. The collection localities of these spec­ imens and the overall geographic range given in Parker and Wiggins (1987) indicate that this species is widely distributed in Michigan. Psilotreta inde­

lInsect Division, Museum of Zoology, University of Michigan, Ann Arbor, MI 48109-1079. 2Headlee Labs, Dept. of Entomology, Rutgers University, New Brunswick, NJ 08901-8536. 200 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4 cisa is found principally in rime areas of cool, gravel-bottom streams and rivers, a habitat that is characteristic of the northern part of the state. This species is probably more restricted to groundwater-fed sections of streams and rivers in southern Michigan. The sericostomatid genus Agarodes is known from Minnesota, Ontario and Maine to Louisiana and Florida (Wiggins 1996). Wiggins (1996) has col­ lected Agarodes larvae from sand and gravel of Ontario lakes and rivers as well as from cool, sandy springs in the southeastern United States. This genus was also expected from Michigan because it was collected from all other states and provinces bordering the Great Lakes. Several sericostomatid larvae from Gull Lake, Kalamazoo Co., Kellogg Biological Station were kindly sent to the senior author by Dr. Richard Mer­ ritt. These were collected on 11 and 16 October 1974, by D. Hayes and W T. Kendall, respectively, and identified as Agarodes distinctus Ulmer by the se­ nior author. According to Merritt, the larvae were collected from sand and gravel in a wave-swept section of Gull Lake ca. 15-20 meters from shore. These specimens should be considered the first record for the state of Michi­ gan. Subsequently, the senior author collected larvae of A. distinctus from a wave-swept portion of Ives Lake (T51N, R28W, Sec. 4), Marquette County in the Upper Peninsula of Michigan. The substrate there is predominately large rocks interspersed with sand and gravel, similar to that described by Wig­ gins. Based on collection localities of these specimens, this species is also probably widely distributed in Michigan.

ACKNOWLEDGMENTS Thanks to Dr. Richard Merritt (Michigan State University) and Joan Martin (Huron Watershed Council) for providing specimens and information. Mark O'Brien (UMMZ) made editorial suggestions. Thanks to Dr. David Gosling of the Huron Mountain Wildlife Foundation for collection access to Ives Lake and use of research facilities of the Huron Mountain Wildlife Foundation. Specimens of P. indecisa from the Huron River are in the possession of the Huron Watershed Council, Ann Arbor, Michigan; from the Escanaba River are in the collection of the University of Michigan of Zoology, Insect Di­ vision (UMMZ); and those from Isle Royale are in the private collection of the junior author. Specimens of A. distinctus from Gull and Ives Lakes are also deposited in UMMZ-Insect Division.

LITERATURE CITED Leonard, J. W. and F. A. Leonard. 1949. An annotated list of Michigan Trichoptera. Oc­ casional Papers of the Museum of Zoology, University of Michigan 522:1-35, 1 chart. Parker, C. R. and G. B. Wiggins. 1987. Revision of the caddisfly genus Psilotreta (Tri­ choptera: Odontoceridael. Royal Ontario Museum, Life Sciences Contributions 144, vi + 55 pp. Ulmer, G. 1905. Neue und wenig bekannte aussereuropaische Trkhopteren, haupt­ sachlich aus dem Wiener Museum. Naturhistorisches Hofmuseums Wien Annalen 20: 59-98. Walker, F. 1852. Catalogue of the specimens of neuropterous insects in the collection of the British Museum. Part 1. London. 192 pp. Wiggins, G. B. 1996. Larvae of the North American Caddisfly Genera (Trichopteral. 2nd edition. University of Toronto Press, Toronto. 457 pp. 1998 THE GREAT LAKES ENTOMOLOGIST 201

NEW LARVAL VARIANTS AND DISTRIBUTIONAL RECORDS FOR PLAUDITUS CESTUS IEPHEMEROPTERA: BAETIDAE)

C. R. Lugo-Ortiz and W. P. McCafferty!

ABSTRACT The larval stage of Plauditus cestus (Ephemeroptera: Baetidae) is re­ described to incorporate mouthpart, leg, and abdominal characteristics not previously accounted for. The examination of populations not studied previ­ ously shows abdominal coloration and morphology to vary considerably. Addi­ tional distributional records from North America are provided.

Provonsha and McCafferty (1982) originally described Pseudocloeon ces­ tum from larvae and male and female adults from Indiana populations. Mc­ Cafferty and Waltz (1990) later transferred the species to Barbaetis Waltz and McCafferty. Lugo-Ortiz and McCafferty (1998) recently erected the genus Plauditus for all North American species assigned to Baetis Leach that lack hind wings or hindwingpads as larvae; P. cestus demonstrates its nat­ ural affinities with this genus and was placed within it. The original species description of P. cestus was relatively incomplete with respect to the larval stage. It did not cover several characters that are now known to be impor­ tant, and did not take into account atypical individual variability that has been discovered recently in additional populations from eastern North Amer­ ica. We herein provide a redescription of the larva of P. cestus that will allow its comparative assessment within Plauditus and that will also allow atypi­ cal geographic populations to be diagnosed correctly. We additionally provide distributional records that considerably extend the known range of the species in North America. Except where otherwise noted, the material exam­ ined is housed in the Purdue Entomological Research Collection, West Lafayette, Indiana.

Plauditus cestus (Provonsha and McCafferty) Larva. Body length: 3.5-4.5 mm; caudal filaments length: 1.7-2.1 mm. Head: Coloration cream to medium yellow-brown, with medium brown to red-brown specks and marki . Labrum (Fig. 1) with branched, marginal setae, submarginal pair oflo ne, simple setae, and row oflong, fine, sim­ ple setae near middle. Hypop arynx as in Figure 2. Left mandible (Fig. 3) with six denticles. Right mandible (Fig. 4) with seven denticles. Maxillae (Fig. 5) with palp segment 1 not reaching apex of galealaciniae; palp segment 2 slightly longer than segment 1, with minute, fme, simple setae scattered

1 Department of Entomology, Purdue University, West Lafayette, IN 47907. ~~------

202 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No.3 & 4

1

5 2

6

9 10 11 Figures 1-11. Plauditus cestus, larva. 1. Labrum. 2. Hypopharynx. 3. Left mandible. 4. Right mandible. 5. Left maxilla. 6. Labium (left-ventral; right­ dorsal). 7. Right foreleg. 8. Tarsal claw. 9. Abdominal color pattern (common). 10. Abdominal color pattern (mature South Carolina specimen). 11. Abdomi­ nal color pattern (immature South Carolina specimen). 1998 THE GREAT LAKES ENTOMOLOGIST 203 over surface; three short, fine, simple setae near medial hump, and one short, fine, simple seta on opposing face. Labium (Fig. 6) with palp segment 2 with three to four short, fine, simple setae dorsally; palp segment 3 with abundant short, fine, simple setae scattered over surface. Thorax: Coloration yellow-brown to light brown, with medium brown to red-brown speckling scatteresl over surface. Hindwingpads absent. Legs (Fig. 7) cream; femora with faint red-brown to pale brown band subproximally; tarsal claws (Fig. 8) with one row of 10-11 dentieles. Abdomen (Figs. 9-11): Ground coloration pale to medium yellow-brown, with medium brown to red-brown speckling; terga 1 and 10 usually pale to medium yellow-brown; terga 2-4 and 6-10 usually with paired submedial medium brown to red-brown specks; segment 5 usually dark brown dorsally and ventrally; tergum 9 usually medium yellow-brown to light brown. Sterna sometimes with small subdermal medium to dark brown spots medially. Cerci with medium to dark brown band slightly beyond middle. Medial cau­ dal filament cream, with four to five short segments. Material examined: Canada. Ontario Prov.: Nottawasaga R., at Simcoe Side Rd. 10, 16-VI-1997, R. W. Griffths and C. Jones, two larvae. USA. Idaho: Madison Co., Henry's Fork, nr. Rexburg, VIII-1993, three larvae (deposited in the Colorado State University Collection, Fort Collins); Kentucky: Carter Co., Big Sinking Cr., 1 mi upstream from confluence with Little Sandy R., 31­ V-1978; South Carolina: Edgefield Co., double branch of Horn Cr., at SR-52, 3-VI-1997, J. F. Napolitano, two larvae (mouthparts and forelegs of one larva mounted on slide [medium: Eupara1]); Greenville Co., Big Durbin Cr., at S­ 23-191, about 2 mi N of Fountain Inn, 18-VII-1979, J. B. Glover, two larvae; Spartanburg Co., Maple Cr., at Co. Rd. 644, 22-VIII-1983, J. B. Glover, larva; Union Co., Henry Cr., at Co. Rd. 29, 13-VI-1983, J. B. Glover, two larvae; York Co., Allison Cr., at SC Hwy 49, about 7 mi SE of Clover, 12-VI-1980, J. B. Glover, six larvae (one larva mounted on slide [medium: Euparal]); York Co., Calabash Cr., at S-46-414, 2.5 mi SE of Clover, J. B. Glover, three larvae; Vermont: Windhams Co., North Brook at west edge of Wilmington, 20-VI­ 1976, W. P. McCafferty, A. V. Provonsha, M. Minno, larva. Species variability: Larvae of P. cestus show considerable variation in ab­ dominal coloration. The most common pattern is shown in Fig. 9 (see also Provonsha and McCafferty [1982]: Fig. 8), where segment 5 is uniformly dark. However, we have seen a mature larval specimen from South Carolina that is distinct in having more extensive speckling and markings dorsally (Fig. 10) and a pale sternum 5 that is only laterally darkened. The abdomen of other mature larval specimens from South Carolina is distinctive in that it is uniformly cream and lacks all banding. One of the immature larval speci­ mens from South Carolina is even more striking because tergum 5 is mostly cream with dark brown distolateral markings and segment 9 is uniformly dark brown (banded) (Fig. 11). In addition, some specimens from South Car­ olina have sterna with small subdermal medial spots, whereas most speci­ mens lack those spots. Because P. cestus co-occurs with P. gloveri McCafferty and Waltz in South Carolina (McCafferty and Waltz 1998), morphological dif­ ferences rather than color pattern should be used to distinguish their larvae. We have also discovered that the appearance of the abdominal shape of P. cestus can vary significantly. The most commonly collected specimens have a somewhat dorsoventrally compressed abdomen (Figs. 9, 11; see also Provon­ sha and McCafferty [1982]: Fig. 8). However, the abdomen of some specimens may appear distended and thus more slender and fusiform (Fig. 10). As far as we know, this distention occurs at the time of fixation and therefore may be an artifact ofpreservation. We have seen similar shape distortion in some 204 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No. 3 & 4 other species of Plauditus. Observation of live material is now required to confirm this. Habitat: Provonsha and McCafferty (1982) indicated that larvae of P. ces­ tus are found in pea gravel in shallow waters (3-35 cm) of third to fourth order streams with slow to moderate currents. The abdominal and cercal banding of larval P. cestus break up their body form against the backdrop of such of a substrate, making them extremely difficult to see in the field. Their small size requires a very small mesh to capture by kickscreen. Distribution: Plauditus cestus was previously known from Arkansas, Col­ orado, Illinois, Indiana, Iowa, and Missouri (Provonsha and McCafferty 1982, McCafferty et al. 1993, Klubertanz 1995, Sarver and Kondratieff 1997). The new records given herein (see material examined above) indicate that the species is quite widespread in North America.

ACKNOWLEDGMENTS We thank B. C. Kondratieff (Fort Collins, Colorado), R. W. Griffiths (Lon­ don, Ontario), and J. F. Napolitano (Columbia, South Carolina) for the dona­ tion or loan of some of the specimens used in this study. We also thank A. V. Provonsha (West Lafayette, Indiana) for the larval abdominal drawings. This paper has been assigned Purdue Agricultural Research Program Journal No. 15657.

LITERATURE CITED Klubertanz, T. H. 1995. Survey ofIowa mayflies. J. Kansas Entomol. Soc. 68: 20-26. Lugo-Ortiz, C. R. and W. P. McCafferty. 1998. A new North American genus of Baetidae (Ephemeroptera) and key to Baetis complex genera. Entomol. News 109: 343-353. McCafferty, W. P. and R. D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera) of North and Middle America. Trans. Am. Entomol. Soc. 116: 769-799. McCafferty, W. P. and R. D. Waltz. 1998. New species of the small minnow mayfly genus Plauditus (Ephemeroptera: Baetidae). Entomol. News 109: 354-356. McCafferty, W. P., R. S. Durfee and B. C. Kondratieff. 1993. Colorado mayflies (Ephemeroptera): an annotated inventory. Southwest. Natural. 38: 252-274. Provonsha, A. V. and W. P. McCafferty. 1982. New species and previously undescribed larvae of North American Ephemeroptera. J. Kansas Entomol. Soc. 55: 23-33. Sarver, R. and B. C. Kondratieff. 1997. Survey of Missouri mayflies with the first de­ scription of adults of Stenonema bednariki (Ephemeroptera: Heptageniidae). J. Kansas Entomol. Soc. 20: 132-140. 1998 THE GREAT LAKES ENTOMOLOGIST 205

DISTRIBUTION OF HETAERINA TlTIA {ODONATA: CALOPTERYGIDAEl IN THE EASTERN GREAT LAKES REGION

Paul D. Pratt1 and Paul M. Catling2

ABSTRACT The lower Thames and Sydenham Rivers in southwestern Ontario have well established populations of Hetaerina titia that represent the northern range limit of the species. Although first discovered in 1985, these popula­ tions are not necessarily recently established. Adults appear from mid-Au­ gust to early September and are most often seen around trees and shrubs overhanging moving water.

Beatty and Beatty (1971) reported Hetaerina tWa (Drury), the Smoky Rubyspot, from four counties in Pennsylvania noting that it reached its northern limit in the state. It is unknown in New York State (Donnelly 1992). Although Glotzhober (1995) noted that the most recent Ohio record was in 1957, two additional sites were recorded in 1995 (R.C. Glotzhober pers. comm.). In Michigan Hetaerina tWa was known from a 1927 record from Oakland Co., where Byers (1927) found two males and one female. It was considered to be "probably an occasional adventive in Michigan" by Kor­ mondy (1958). In 1982 H. tWa was discovered in Livingston County, Michi­ gan where it is still present in 1998 (Weichsel1998). Hetaerina titia was not listed for Canada by Westfall & May (1996), and was first reported from Ontario the same year in Pratt's (1996) check­ list of Ontario Odonata. Considering that except for the single extant Michigan site, the other occurrences in the surrounding area are all south of Canada (Fig. 1), it seems unlikely to occur in Ontario. However it was discovered on the lower Thames River by Paul Pratt and Jo Barten in 1985. In 1987 and 1991 it was found to be frequent or abundant at six localities along the Thames River. It was also found at one site on the Sydenham River near Alvinston in 1991. In 1997 we found it at two of the previously discovered sites on the Thames River and at an additional locality further upstream (Fig. 2). All of the Thames River locations are situated along a 35 km portion of the river between Thamesville and London. These locations represent the only reports for Canada and thus are at the northern known range limit. Based on the Ontario occurrences, it seems likely that H. tWa may be more common and well established in the eastern Great Lakes region than

17100 Matchette Road, LaSalle, Ontario, N9J 2S3 CANADA 22326 Scrivens Drive, RR 3 Metcalfe, Ontario, KOA 2PO CANADA 206 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No.3 &4

62°30' 82°15' 82' 61'45' 81'30'

Figure 1. Ontario distribution of the Smoky Rubyspot, Hetaerina titia.

current knowledge indicates. The earliest and latest dates for Ontario are 16 August and 4 September. We have seen no distinct peaks in population num­ bers during this period. In Ohio the flight period extends from 22 Aug. to 8 Oct. (Glotzhober 1996). Although the discovery of Smoky Rubyspot in southwestern Ontario is recent, it may have been long established there though overlooked in the field. Adults are more often associated with trees and shrubs overhanging moving water than H. americana (Fabr.) where the two occur together, and it is more likely to be seen from the water than from the shore. Associated dam­ selflies include Argia apicalis (Say), A moesta (Hagen), Enallagma exsulans (Hagen) and Hetaerina americana. H. tWa from the Great Lakes region are the tricolor form with mostly clear wings. The distinctly black-tipped hindwings and relatively long and dark abdomen are characteristic of males. The sides of the thorax of the female are more pale than metallic, instead of more metallic than pale as in H. americana, and the abdomen of the the female H. titia is much darker. 1998 THE GREAT LAKES ENTOMOLOGIST 207

86° 84° 82° 78° 76° 74' Hetaerina titia .... Ontario collections II • literature reports o 100 Miles

74° Figure 2. Distribution of Hetaerina titia in the eastern Great Lakes Region.

ACKNOWLEDGMENTS We thank Mark O'Brien, Bob Glotzhober and Karen Cedar for their com­ ments on an earlier draft of this paper.

LITERATURE CITED Beatty, AF. and G.H. Beatty. 1971. The distribution of Pennsylvania Odonata. Penn. Acad. Sci. 45: 145-167. Byers, C.F. 1927. An annotated list of the Odonata of Michigan. Occ. Papers Mus. Zool. Univ. Mich. 183: 1-16. Donnelly, T.W. 1992. The Odonata of New York State. Bull. Amer. Odonatology 1(1): 1-27. Kormondy. E.J. 1958. Catalogue of the Odonata of Michigan. Univ. of Michigan, Mu­ seum of Zoology, Misc. Publ. 104: 1--43. Glotzhober, R.C. 1995. The Odonata of Ohio-a preliminary report. Bull. Amer. Odona­ tology 3(1): 1-30. Glotzhober, R.C. 1996. Ohio Odonata Survey (http://mcnet.marietta.eduJ-odonata /species/odospec.html) 208 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No.3 & 4

Pratt, P. D. 1996. Checklist of the Odonata ofOntario. Ontario Insects 2(1): 9-12. Weichsel, J.I. 1998. Hetaerina titia IS in Michigan!. Williamsonia 2(3): 1. Westfall, M.W. and M.L. May. 1996. Damselflies of North America. Scientific Publish­ ers, Gainesville. 649 p. 1998 THE GREAT LAKES ENTOMOLOGIST 209

GLEANING ON COREIDAE (HETEROPTERA) BY TACHOPTERYX THOREYI (ODONATA: PETALURIDAEl

R.D. Woltz 1

Tachopteryx thoreyi (Hagen in Selys) is an uncommon or rare, fen and seep dwelling dragonfly noted for its large size, restricted habitat require­ ments, and characteristic habit of perching on tree trunks. While performing insect sampling and habitat assessments in a fen at Mounds State Park, Anderson, Indiana, I observed a male T. thoreyi land on a tree trunk and capture an insect on the bark of the tree, 1 June 1998, at approximately 1430 hr. Upon approaching and capturing the dragonfly, I found that it had captured and held in its grasp a large leaf-footed bug, prob­ ably Acanthocephala terminalis (Dallas) (Heteroptera: Coreidae). The drag­ onfly was caught by hand and photographed with its prey still in its grasp. Unfortunately, the dragonfly released it prey during the photograph and the prey was not able to be retrieved from the tall grasses for species Verification. Probable identification of the coreid was kindly provided by AI Wheeler, Jr., Clemson University, based on the above mentioned photograph and verbal descriptions of the species. By the time I captured the dragonfly, it had over­ turned the coreid and begun to feed on the coreid at approximately the ante­ rior metasternal suture (Le., between the mesosternum and the metaster­ num). It is of interest that although the defensive scent of the coreid was obvious, this apparently did not seem to deter the feeding by T. thoreyi in this case. Report of this feeding behavior in T. thoreyi is noteworthy for several rea­ sons. Gleaning (the taking of resting prey) in dragonflies and damselflies has been reported by Corbet (1963, page 148). However, this behavior has not previously been reported in the Petaluridae or in T. thoreyi specifically. The majority of data that has been published on the feeding of T. thoreyi includes many large and small insect species, e.g., Dunkle (1981) reports butterflies, moths, beetles, and other odonates, Williamson (1900) reported T. thoreyi feeding on moths, and Montgomery (1931) reported T. thoreyi feeding on Calopteryx (Odonata) (also reported by Dunkle (1981», moths, and in Mont­ gomery (1932) the ant species, Camponotus pennsylvanicus DeGeer. Feed­ ing on coreidae by T. thoreyi has not been previously reported.

ACKNOWLEDGMENTS I thank Paul Rothrock, Taylor University, for taking the photograph of dragonfly and pfey. I thank AI Wheeler, Jr., Clemson University, for assis­ tance in identitying, to the degree possible, the coreid in the above mentioned photograph. I thank Tom Swinford, Indiana Department of Natural Re­

1 IDNR, Division of Entomology and Plant Pathology, 402 West Washington, Room W-290, Indianapolis, IN 46204. 210 THE GREAT lAKES ENTOMOLOGIST Vol. 31, No. 3 & 4 sources, Jim Curry, Franklin College, and Ken Tennessen, Florence, Al­ abama, for critical comments on an early draft of this note.

LITERATURE CITED Corbet, P.S. 1963. A biology of dragonflies. Quadrangle Books, Inc., Chicago. 247 pp. Dunkle, S.W. 1981. The ecology and behavior of Tachopteryx thoreyi (Hagen) (Anisoptera: Petaluridae). Odonatologica 103: 189-199. Montgomery, B.E. 1931. Records of Indiana dragonflies, V, 1930. Proc. Indiana Acad. Science. 40: 347-349. Montgomery, B.E. 1932. Records of Indiana dragonflies, VI, 1931. Proc. Indiana Acad. Science 41: 449-454. Williamson, E.B. 1900. On the habits of Tachopteryx thoreyi (Order: Odonata). Ento­ 'J; mol. News 11: 398-399. I 1998 THE GREAT lAKES ENTOMOLOGIST 211

ENALLAGMA ANNA, A DAMSELFLY NEW TO THE GREAT LAKES REGION (ODONATA: COENAGRIONIDAE)

Mark F. O'Brien 1 and Paul D. Pratt2

ABSTRACT Enallagma anna, a predominantly western North America damselfly, is now recorded from southwestern Michigan and southwestern Ontario for the first time.

Enallagma anna Williamson, the River Bluet, is a striking blue and black coenagrionid damselfly first described by Williamson (1900) from Wyoming. Most records are from the United States, where the species ranges from the north-central plains states west to California, Arizona, and Oregon. Westfall and May (1996) also lists Illinois, Wisconsin and the province of Alberta, which updates Walker (1953), as he did not list this species for Canada. Westfall and May (1996) describe E. anna as a "... very robust species, mostly from relatively arid western highlands." Provansha (1975) provided behavioral notes from Utah, and found larvae in small to medium streams with moderate flow at elevations of 4200 ­ 7000 ft. We report on the discovery of two new populations of E. anna from Cass County, Michigan and Essex County, Ontario (Canada) during fieldwork con­ ducted in 1998. The Ontario record extends the known range of E. anna by 380 km eastward. The first author collected E. anna in Cass Co., MICHIGAN on 21 June 1998 in vegetation bordering the Dowagiac River upstream from McKenzie Hwy. (41.997°N x 85.982°W). A total of four males and one female were collected along the river, which is a slow-flowing clear stream about 6 m wide at the point of collection. The bottom substrate is gravelly/sandy with areas of siltation in the aquatic weedbeds. The bright blue abdomen of the robust (ca. 33 mm) male is quite noticeable against vegetation and over the water. The second author collected E. anna in Essex Co., ONTARIO on 21 June 1998 from emergent vegetation in Big Creek in Amherstburg (41.1101oN x 83.0825°W). Despite a lengthy search only a single male was found along this narrow, slow flowing, muddy stream. Enallagma civile (Hagen) was the most common damselfly noted at this location.

lInsect Division, Museum of Zoology, University of Michigan, Ann Arbor, MI 48109-1079. email:[email protected]. 27100 Matchette Road, LaSalle, Ontario, N9J 2S3, CANADA. email: prairie@net­ core.ca 212 THE GREAT LAKES ENTOMOLOGIST Vol. 31, No.3 & 4

DISCUSSION It is unknown how long E. anna has been a resident of Michigan or On­ tario. However, the species has not been reported from Ohio, and that state has been actively surveyed for the past decade (Glotzhober 1995). Kormondy (1958, 1962) did not list the species for Michigan, and it has not appeared in any publications on the Indiana fauna. According to the Illinois State Mu­ seum's web site, E. anna is known from Boone, Cook, JoDavies, and McHenry Counties (http://www.museum.state.il.us/research/entomology/ od_cnty4species.html). Cook Co., Illinois would therefore harbor the closest known population to Cass Co., MI. The population along the Dowagiac River seemed to be substantial, as numerous adults that were also probably E. anna were seen along a 100 m segment of the river. Whether these populations are indicative of recent spo­ radic founding events or slow range expansion as the result of changing cli­ matic patterns or changes in water quality and quantity is less clear. Although an eastern species, E. civile has been less common in the north­ eastern United States and Canada. Catling (1996) has documented that species' expansion into southern Ontario, where it has become the common Enallagma in southwestern Ontario within the last 30 years. At the Amher­ stburg site, E. civile is an associate of E. anna and is very similar in color pattern and size, and therefore, E. anna could easily be overlooked from a distance. In contrast, Enallagma basidens Calvert has been gradually extending its range into the northeast from the southwest, and this has been well-docu­ mented over the past 70 years (O'Brien 1997). However, E. basidens prefers lentic habitats that are man-made or disturbed, not clear streams, and dis­ turbed habitats have certainly increased during this century, thereby favor­ ing species (such as E. basidens and E. civile) that can take advantage of those situations. We are certain that as surveys of the fauna of the Great Lakes region be­ come more systematic and intensified, new records of this sort are more likely to appear.

ACKNOWLEDGMENTS We thank Nick Donnelly for verifying the E. anna specimens and for re­ viewing the manuscript. The senior author thanks Laura Krueger and Bill Westrate for their assistance and good company on the June 21 Michigan Odonata Survey field trip. The Cass County specimens are deposited in the University of Michigan Museum of Zoology and bear the following numbers: MOS0018003, MOS0018021, MOS0018057, and MOS0018386. The Essex County specimen is in the collection of the Paul Pratt, and will eventually be housed in the Canadian National Collection.

LITERATURE CITED Catling, P. M. 1996. Evidence for the recent northward spread of Enallagma civile (Zy­ goptera: Coenagrionidael in southern Ontario. Proc. Entomol. Soc. Ontario 127: 131-133. Glotzhober, R. C. 1995. The Odonata of Ohio-A preliminary report. Bull. of American Odonatology 3{l):1-30. 1998 THE GREAT LAKES ENTOMOLOGIST 213

Kormondy, E. J. 1958. A catalogue of the Odonata of Michigan. Misc. Pub!. Museum of Zoology, Univ. of Michigan. 104:1-43. Kormondy, E. J. 1962. New records of Michigan Odonata. Entomo!. News 73:191-194. Provansha, A. W. 1975. The Zygoptera (Odonata) of Utah with notes on their biology. Great Basin Naturalist. 35:379-390. O'Brien, M. F. 1997. Enallagma basidens (Odonata: Coenagrionidae) expands it range into Michigan. Great Lakes EntomoL 30(4):181-183. Walker, E. M. 1953. The Odonata of Canada and Alaska, Vol. 1. University of Toronto Press: Toronto, Ontario. 292 pp. Westfall, M. J., Jr. and M. L. May. 1996. Damselflies of North America. Scientific Pub­ lishers, Gainesville, FL 650 pp. Williamson, E. B. 1900. Notes on a few Wyoming dragonflies (Order Odonata). Ento­ mol. News. 11(5):453-458, 1 pI. NEW EDITOR OF THE GREAT LAKES ENTOMOLOGIST

I am taking this opportunity to thank the many authors, reviewers and Michigan Entomological Society board members for their support over the past 11 years. I am proud that I have been able to serve the Society in the capacity as journal editor during this period. However, the time is right for me to go on to other in­ terests and pass the editorship on to someone else. I am pleased to announce that Dr. Randall Cooper will assume the editorial duties of the Great Lakes Ento­ mologist, effective February 15, 1999. Randy Cooper has an M.S. in Entomology from Ohio State (1981) and a Ph.D. in Entomology from the University of Georgia (1986). He has stepped for­ ward to take on a job that can be quite challenging at times, and I believe that he will be an excellent person for the job. I have tried to make the transition as smooth as possible, and Randy will start work on the spring issue (Volume 32, Number 1). I have supplied the reviewers' addresses, manuscript database and other materials so that he can continue work on the journal and process incom­ ing manuscripts as quickly as possible. Obviously, some manuscripts are still in the review stage and others are in various stages of readiness, so do not be alarmed if there is a delay in processing your manuscript. We appreciate your patience during this transition period. I hope that you will continue to support the journal and Randy's efforts by providing him with manuscripts and by offering your services as a potential re­ viewer (if you have not already reviewed manuscripts for me). The journal has a wide variety of articies, and we need a diverse assemblage of reviewers to ad­ equately peer-review manuscripts.

The address of the new editor of the Great Lakes Entomologist is:

Dr. Randall Cooper 16672 152nd Avenue Spring Lake, MI 49546 email: ren:z:[email protected]

It has been a pleasure working with authors and referees over the past decade. I also thank Vivian Bradbury of Sans Serif Inc., (the typesetters of the Great Lakes Entomologist) for being so very helpful to us and improving the quality of the journal in a number of ways. The journal will continue its tradition of providing a quality product and an outlet for papers focused primarily on the Great Lakes Region. With your continued support of the Society, the new editor will steer the Great Lakes Entomologist into the new century and into its 33rd year of publication. Mark O'Brien February, 1999 INSTRUCTIONS FOR AUTHORS

SUBJECTS Papers dealing with almost any aspect of entomology will be considered for publication in The Great Lakes Entomologist. Appropriate subjects are those of interest to professional and amateur entomologists in the North Central States and Canada, as well as general papers and revisions di­ rected to a larger audience while retaining an interest to readers in our geographic area. All manuscripts are refereed by two reviewers, except for short notes, which are reviewed ex­ ternally at the discretion of the Editor. REQUIREMENTS Manuscripts must be typed, double-spaced, with 1" margins on 8 112 x 11" or equivalent size paper, and submitted in triplicate. Authors submitting only one coPy will be asked to send two more as the editor will not make the copies. Please italicize or underlme only those words that are to be italicized. Use subheadings sparingly. 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So long as these symbols are clearly seen in the manuscript, adjustments can be made in the copy sent to the print­ ers, Authors should use italics, rather than underline when submitting manuscripts on disk. PAGE CHARGES Papers published in The Great Lakes Entomologist are subject to a page charge of $35.00 per published page. Members ofthe Society, who are authors without funds from grants, institutions, or industry, and are unable to pay costs from personal funds, may apply to the Society for financial assistance. Application for subsidy must be made at the time a manuscript ill mitially sub· mitted for publication. Authors will receive page proof, together with an order blank for sepa­ rates. Extensive changes to the proof by the author will be billed at a rate of $1.00 per line. COVER ARTWORK Cover art or photographs are desired for upcoming issues. They are published free of charge. We only require that they be suitably mounted as described above, and that the subject be identi­ fied as accurately as possible. EDITOR'S ADDRESS All manuscripts for The Great Lakes Entomologist should be sent to the Editor, Mark F. O'Brien, Insect Division, Museum of Zoology, The University of Michigan, Ann Arbor, MI, 48109­ 1079, USA OTHER BUSINESS Other correspondence should be directed to the Secretary, Michigan Entomological Society, clo Dept. of Entomology, Michigan State University, East Lansing, MI 48824-1115.