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Vol. 3'3, No.3 & 4 Fall/Winter 2000 THE GREAT LAKES ENTOMOLOGIST

PUBLISHED BY

THE MICHIGAN ENTOMOLOGICAL SOCIETY THE GREAT LAKES ENTOMOLOGIST

Published by the Michigan Entomological Society

Volume 33 No. 3-4

ISSN 0090-0222

TABLE OF CONTENTS

New distribution record for the endangered crawling water beetle, Brychius hungerfordi (Coleoptera: Haliplidael ond notes on seasonal abundance and food preferences Michael Grant, Robert Vande Kopple and Bert Ebbers ...... 165

Arhyssus hirtus in Minnesota: The inland occurrence of an east coast species (Hemiptera: Heteroptera: RhopalidaeJ John D. Laftin and John Haarstad .. 169

Seasonol occurrence of the sod webworm moths (lepidoptera: Crambidoe) of Ohio Harry D. Niemczk, David 1. Shetlar, Kevin T. Power and Douglas S. Richmond. . ... 173

Leucanthiza dircella (lepidoptera: Gracillariidae): A leafminer of leatherwood, Dirca pa/us/tis Toby R. Petriee, Robert A Haack, William J. Mattson and Bruce A. Birr. . . . 187

New distribution records of ground beetles from the north-central United States (Coleoptera: Carabidae) Foster Forbes Purrington, Daniel K. Young, Kirk 1. lorsen and Jana Chin-Ting lee ...... 199

Libel/ula flavida (Odonata: libellulidoe], a dragonfly new to Ohio Tom D. Schultz. . ... 205

Distribution of first instar gypsy moths [lepidoptera: Lymantriidae] among saplings of common Great Lakes understory species 1. l. and J. A. Witter ...... 209

Comparison of two population sampling methods used in field life history studies of Mesovelia mulsonti (Heteroptera: Gerromorpha: Mesoveliidae) in southern Illinois Steven J. Taylor and J. E McPherson ...... 223

COVER PHOTO

Scudderio furcata Brunner von Wattenwyl (Tettigoniidae], the forktailed bush katydid. Photo taken in Huron Mountains, MI by M. F. O'Brien. THE MICHIGAN ENTOMOLOGICAL SOCIETY

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Copyright © 2002. The Michigan Entomological Society 2000 THE GREAT LAKES ENTOMOLOGIST 165

NEW DISTRIBUTION RECORD FOR THE ENDANGERED CRAWLING WATER BEETLE BRYCHIUS HUNGERFORDI (COLEOPTERA: HALlPLIDAE) AND NOTES ON SEASONAL ABUNDANCE AND FOOD PREFERENCES

Michael Grant 1, Robert Vande Kopple 1, and Bert Ebbers2

ABSTRACT The Federally beetle, Brychius hungerfordi, has been discov­ ered at a new location in Northern Lower Peninsula of Michigan. We also report preliminary data on a seasonal variation in relative abundance and on its possible food plants.

The crawling water beetle, Brychius hungerfordi Spangler, is known from only three sites in Michigan and from one in Ontario (Spangler 1954, Wils­ mann and Strand 1990, Roughly 1991, Strand and Spangler 1994, Keller et al. 1998). We report the discovery of a fourth location in the Northern Lower Peninsula of Michigan. We can only speculate about the beetle's natural his­ tory since it is so rare, but details concerning its abundance and food prefer­ ences are beginning to emerge.

MATERIALS AND METHODS Beetle collecting occurred in Van Hetton Creek, Montmorency County, Michigan (Twp. 31N, Sec. 5, SW K Beetles were collected with a long handled D-net (30-cm diameter) using a sweeping motion across the cur­ rent with the lip of the net held slightly above the bottom of the creek. After the initial pass the net was quickly moved back over the same area in a downstream direction, to catch beetles dislodged from the bottom during the first pass. Collecting also occurred at a large pool, located on the East Branch ofthe Maple River, Emmet County, known to support a B. hungerfordi population. In order to assess the temporal variation in beetle abundance, we sampled the pool monthly from March through December, 1999, at the middle of the month. We divided the pool into three sections, head, midsection, and tail, and sampled each section equally for a total of one hour sampling. Immedi­ ately after counting the beetles were released back into the pool from which they had been collected. Sample removal error was considered insignificant given the length oftime between sampling dates. Algal samples were collected in the field and preserved in 4% formalin for later identification in the lab.

lUniversity of Michigan Biological Station, Pellston, MI 49769. 2Great Lakes Ecosystems, P. O. Box 156, Indian River, MI 49749. 166 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 & 4

RESULTS AND DISCUSSION In July 1999, six adult beetles were captured). l'he creek drains wet­ lands, but also receives some groundwater based upon water temperature measurements made at the time of collection. The beetles were found dis­ persed along a stretch of creek several hundred meters in length and down­ stream from a pool, formed as the creek flowed from a culvert beneath a county road. This site differs from previously described locations in that the creek channel is composed of sand overlain with a thin layer of detritus (Roughley 1991, Wilsmann and Strand 1990, Strand 1989). In addition to searching for new beetle locations, we have begun study of its basic biology and natural history. Initial observations suggest a seasonal cDmponent to its relative abundance and that its food source may be filamentous green or blue-green algae. The relative abundance of beetles in the pool, as the number of adults captured per hour of sampling, is in Figure 1. The rela­ tive abundance in November is probably underestimated due to partial ice cover, which prevented a complete search of the pool. The ice was gone by De­ cember, allowing a normal search pattern to resume. The data suggest that some B. hungerfordi adults may survive through the winter, which was con­ firmed on 10 February 2000 when five adults were captured from the pool be­ neath a 24 em cover of ice. This is consistent with study of other haliplids. Hickman (1931) in laboratory studies reported that some adult haliplids sur­ vived as long as 18 months in captivity. He also reported finding some lentic species beneath ice cover. Our preliminary data also suggest that several generations may be present during a single season. The increase in relative abundance in May followed by a second increase in October suggest that a second brood of adults may emerge late in the season. If these observations are confirmed in additional studies, then an optimal period may exist in which to conduct survey work. Searching during less than optimal times would make the beetle that much more difficult to detect. ... s:. 100 90 -f ":::s 80 a.. -CG CJ 60 53 !g 44 '-J:: 40 28 ~ 28 17 Ql 20 t 9 12 I = 0 3 :::s L ... ___ ...__ ,-, 0 ..11 «" I .. 1 Mar Apr May Jun• Jul Aug Sep Oct Nov Dec 1999 Figure 1. Seasonal distribution of B. hungerfordi from E. Branch Maple River, Emmet County, Michigan. 2000 THE GREAT LAKES ENTOMOLOGIST 167

During sampling we observed that beetles were frequently found associ­ ated with dense growths of epilithic algae. Samples of cobble and rocks from areas in the pool containing beetles and from areas where they were absent were collected and brought to the lab for microscopic examination. Sites with beetles contained approximately 30% detritus with the remainder living bio­ mass. There was no clearly dominant genus of algae, but a diverse assem­ blage of Cocconeis, Oedogonium, Cymbella, Gomphonema, Navicula, Calothrix, Lyngbya, Chroococcus, Oscillatoria, Nitzschia, Frustulia, Mi­ crospora and Amphora. Samples from sites without beetles consisted of ap­ proximately 60-70% detritus with the remainder being living biomass. The living algae were 50% Audovinella violaceae red alga), 30% Epithemia (a N-fixing diatom) and 20% genera similar to beetle group. Calothrix, Lyngbya, Microspora, Oscillatoria and Oedogonium are fila­ mentous blue-green or green algae. In Hickman's feeding experiments, he re­ ported that some larva and adults of Michigan Haliplidae did well only on Spirogyra, also a filamentous green alga (Hickman 1931). It is not uncom­ mon among herbivorous insects to exhibit a limited range of foods (Matthews and Matthews 1978). Until we know the specific food plants of Brychius we can only surmise that they might be as restricted as for other haliplids and thereby place constraints on beetle distribution. Successful protection of Brychius hungerfordi depends upon expanding our knowledge of its natural history. It is our hope that this report adds to the understanding of the beetle's distribution and begins to address more fundamental questions regarding its seasonal abundance and food prefer­ ences.

ACKNOWLEDGMENTS The authors thank Dr. Rex Lowe and Aga Pinowska of the Department of Biological Science, Bowling Green State University, Ohio for their assistance in i the samples, and for the comments from two anonymous reviewers. e U.S. ish and Wildlife Service (PRT 839782) and Wildlife Di­ vision of the Michigan Department of Natural Resources (PN1304) provided sampling permits.

LITEHATURE CITED Hickman, J. R. 1931. Contribution to the biology of the Haliplidae (Coleoptera). Ann. Entomol. Soc. Am. 24: 129-142. Keller, T. A., M. Grant, B. Ebbers and R. Vande Kopple. 1998. New record for the en­ dangered crawling water beetle, Brychius hungerfordi (Coleoptera: Haliplidael in Michigan including water chemistry data. Great Lakes Entomol. 31: 137-139. Matthews, R. W. and J. R. Matthews. 1978. Insect behavior. John Wiley and Sons, New York. Roughley, R. E. 1991. Brychius hungerfordi Spangler (Coleoptera: Haliplidae), The fIrst record from Canada with notes about habitat. Coleopterists Bull. 45: 295-296. Spangler, P. J. 1954. A new species of water beetle from Michigan (Coleoptera: Halipli­ dae). Entomol. News 65: 113-117. Strand, R. M. 1989. The status of Brychius hungerfordi (Coleoptera: Haliplidael in northern Michigan. Report to the Nature Conservancy, 25 October. Strand, R. M. and P. ,L Spangler. 1994. The natural history, distribution, and larval de­ scription of Brychius hungerfordi Spangler (Coleoptera: Haliplidae). Proc. Entomol. Soc. VVash. 96: 208-213. r 168 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

Wilsmann, L. A. and R. M. Strand. 1990. A status survey of Brychius hungerfordi I (Coleoptera: Haliplidae) in Michigan. Report to the U.S. Fish and Wildlife Service. Region 3, Endangered Office. Federal Building, Fort Snelling, Twin Cities, MN, 55111. 22 August. 2000 THE GREAT LAKES ENTOMOLOGIST 169

ARHYSSUS HIRTUS (HEMIPTERA: HETEROPTERA: RHOPALIDAE) IN MINNESOTA: THE INLAND OCCURRENCE OF AN EAST COAST SPECIES

John D. Lattin 1 and John Haarstad2

ABSTRACT Arhyssus hirtus (Hemiptera: Heteroptera: Rhopalidae), is reported from the Cedar Creek Natural History Area, a Long-Term Ecological Research site, outside of Minneapolis, Minnesota where over 4000 species of arthro­ pods have been collected. This species has previously been known only from a narrow zone along the sandy edges of the Atlantic Ocean (Maryland, Massa­ chusetts, New Jersey and New York). The species is known on Hudsonia to­ mentosa at these ocean sites, but other hosts may be involved at Cedar Creek. This small species of Arhyssus occurs in both micropterous and macropterous forms, unusual for this genus. Thus far, only micropterous forms have been collected at the Minnesota site.

Corizus hirtus (Hemiptera: Heteroptera: Rhopalidae) was described by Torre-Bueno (1912) from Yaphank, Long Island, New York, based upon long­ and short-winged adults. The specimens were taken at a sandy, grassy site in pine woods in 1911. Wheeler and Henry (1984) reported it from habitats from Massachusetts, Maryland, New Jersey, and New York (as Arhyssus hirtusl. Henry (1988) summarized the taxonomic history of the species in North America. Both sexes of this small species (ca 4.5 mm) occur in macropterous and micropterous form (Torre-Eueno 1912). Chopra (1968) revised the genus Arhyssus. Later, Wheeler and Henry (1984) added information on the New ,Jersey specimens that had been collected "under Hudsonia" by H. G. Barber and "in and under Hudsonia" on Long Island by Nathan Banks. These speci­ mens are in the U. S. National Museum in Washington, D. C. Wheeler and Henry also reported on the collection nymphs and adults of A. hirtus in and under the mats ofHudsonia tomentosa Nutt. The specimens reported here are from the Cedar Creek Natural History Area, Long-Term Ecological Research (LTER) site, near Minneapolis, Min­ nesota, as part of a 25-year study of insects and other arthropods on this 2200 ha site. Over 4000 species of arthropods have been collected, chiefly by J. Haarstad, with additional species added annually. The Cedar Creek site has become one of a handful of well-known sites in North America that in­ clude the H.J. Andrews Experimental Forest, in western Oregon (about 3700 species known) and Central Plains Experimental Range, in Colorado (1649 species known), both LTER sites as well (Parsons et al. 1991, Kumar et al.

lDepartment of Entomology, Oregon State University, Corvallis, OR 97331-2907. 2Cedar Creek Natural History Area, East Bethel, MN 55005. 170 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

1976). Some very interesting species, including Arhyssus hirtus, have been collected at this Minnesota site. This bug was known formerly from a narrow range along the mid-Atlantic coast (Wheeler and Henry 1984).

COLLECTION RECORDS NEW YORK: Yaphank, Long Island (type material) (Torre-Bueno 1912); Nassau Co., Bayville, Suffolk Co., Riverhead; Smith's Point, Fire Island (Wheeler and Henry 1984). MARYLAND: Worcester Co., Assateague Island National Seashore (Wheeler and Henry 1984). :MASSACHUSETTS: Essex Co. (Wheeler and Henry 1984). All Minnesota specimens deposited in the col­ lections of the Long-Term Ecological Research Site at the Cedar Creek Nat­ ural History Area. MINNESOTA: Anoka Co., Cedar Creek Natural History Area, near East Bethel (this paper). NEW JERSEY: Ocean Co., Lakehurst (Wheeler and Henry 1984).

RESULTS All eight specimens ofA. hirtus from the Minnesota site were micropter­ ous with collection dates ranging from 24 June to 24 September, 1985-1991­ These specimens were swept from four different grassland sites, all of which contained Lechea stricta, and most contained Helianthermum bicknellii. Al­ though Hudsonia tomentosa occurs at the Minnesota location, it was not found at the specific sites where the bugs were taken. It is of interest that the xeric sandy old fields at Cedar Creek contained several other plants showing disjuncts distributions including Aristida tuberculosa and Poly­ gonella articulata. Other disjunction of insects may occur as well.

DISCUSSION Voss (1985) discussed the distribution of Hudsonia tomentosa Nutt. in Michigan (family Cistaceae) and included a map (No. 824) of its distribution. The plant, known as beach-heather or false-heather, is found on sand dune ridges in open forests of pine, oak or other trees along the coast of much of Michigan. Ownbey and Morley (1991) listed H. tomentosa from Minnesota and included a map ofits distribution. At least five other species of Cistaceae occur in Minnesota, two in the genus Helianthemum and three in Lechea, with species of both genera found at Cedar Creek and some found associated with the bug. Further study is needed to clarify the exact hosts at the Min­ nesota site. Skog and Nickerson (1973) provided additional information on Hudsonia in their study of this genus. Grigal (1985) and Woucha et aL (1995) provided detailed discussions of the unusual St. Croix River Valley and the Anoka Sand Plain in Minnesota-a landscape sure to surprise us again. Specimens of the Rhopalidae species, Arhyssus hirtus (Torre-Bueno), were found at the Cedar Creek Natural Area near East Bethel, Minnesota (a Long-Term Ecological Research site). All specimens were micropterous, al­ though long-winged adults are known from sites from Maryland, Massachu­ setts, New Jersey, and New York, along the Atlantic coast. The host plant of this bug along the Atlantic coast is Hudsonia tomentosa and although this plant is found at Cedar Creek, the bug has not yet been found on it. There, r 2000 THE GREAT LAKES ENTOMOLOGIST 171

collection sites of the bug contained Leachea stricta and/or Heliantherum bicknellii. One of us (JDL) has seen what appears to be suitable habitats for the bug in the vicinity of Pentwater, Michigan where Hudsonia tomentosa is said to occur. See Voss (1985) for the distribution of this plant in Michigan. While the presence of A. hirtus in Minnesota appears to be a major disjunction, careful field work throughout the Great Lakes region may disclose its occur­ rence at other localities.

ACKNOWLEDGMENTS Our thanks to L. Parks for careful preparation of the manuscript. Sup­ port to J.D.L. from N.S.F. grant BSR-90-11663 is gratefully acknowledged.

LITERATURE CITED Chopra, N. P. 1968. A revision of the genus Arhyssus Stal. Ann. Entomol. Soc. Am. 61: 624-655. Grigal, D. F. 1985. The landscape and soils of Cedar Creek Natural History Area. Nat­ uralist 36: 8-15. Henry, T. J. 1988. Family Rhopalidae, pp. 652-64. In: T. J. Henry and R. C. Froeschner (eds.), Catalog of the Heteroptera, or true bugs, of Canada and the Continental United States. E. J. Brill, Leiden. Kumar, R., R. J. Lavigne, J. E. Lloyd and R. E. pfadt. 1976. Insects of the Central Plains Experimental Range, Pawnee National Grassland. Univ. Wyoming, Agric. Exp. Sta., Sci. Month. 32: 1-74. Ownbey, G. B. and T. Morley. 1991. Vascular plants of Minnesota: A checklist and atlas. Univ. Minnesota Press, Minneapolis. Parsons, G. L., G. Cassis, A. R. Moldenke, J. D. Lattin, N. H. Anderson, J. C. Miller, P. Hammond and T. D. Schowalter. 1991. Invertebrates of the H. J. Andrews Experi­ mental Forest, western Cascade Range, Oregon. V: An annotated list of insects and other arthropods. U. S. Dept. Agric., For. Ser., Pacific Northwest Res. Sta., Gen. Tech. Rep. P.NW-GTR-290. Skog, J. T. and N. H. Nickerson. 1973. Variation and speciation in the genus Hudsonia. Ann. Missouri Bot. Gard. 59: 454-464. Torre-Bueno, J. R. de lao 1912. A new Corizus from the northeastern United States (Hemip., Coreidae). Entomol. News 23: 217-219. Voss, E. G. 1985. Michigan flora. Part II. Cranbrook Institute Sci., Bull. 59. Wheeler, A. G., Jr. and T. J. Henry. 1984. Host plants, distribution, and description of fifth-instar nymphs of two little-known Heteroptera, Arhyssus hirtlls (Rhopalidae) and Esperanza texana (Alydidae). Fla. EntomoL 1984: 521-528. Woucha, D. S., B. C. Delaney and G. E. Nordquist. 1995. Minnesota's St. Croix River Valley and Anoka Sand Plain. Univ. Minnesota Press, Minneapolis. 2000 THE GREAT LAKES ENTOMOLOGIST 173

SEASONAL OCCURRENCE OF THE SOD WEBWORM MOTHS (LEPIDOPTERA: CRAMBIDAEI OF OHIO

Harry D. Niemczykl , David J. Shetlar2, Kevin T. Power! and Douglas S. Richmond!

ABSTRACT While nearly 100 species of sod webworms are known to occur in North America, the species complex and seasonal occurrence of these moths has been documented in relatively few states. For Ohio, there is little published record of the sod webworm species complex, and the seasonal occurrence of only a few economically important species has been documented. Using black light traps, sod web worm adult flight activity was monitored over the course of three to five years at four different locations throughout Ohio. In this paper we report the seasonal occurrence of sod web worms species captured at these locations. These data provide a historical benchmark of sod web­ worm species diversity, local abundance, and seasonal occurrence in Ohio.

Although far from complete, recent gains have been made in document­ ing the lepidopteran fauna of Ohio. Major contributions include documenta­ tion of the superfamililies Papilionoidea and Hesperiioidea (butterflies and skippers) (Iftner et al. 1992), and the moth family Noctuidae (Rings et al. 1992). The two prior groups have likely received more attention because of their relatively large size and colorful ornamentation (Scott 1986), while many of the Noctuidae are serious pests of agricultural crops (Rings et al. 1992). Conversely, many moth families are often overlooked simply because they are smaller, less visually appealing, harder to identify, and tend to be nocturnal. One such group, the sod web worm moths (Lepidoptera: Cram­ bidae), comprise a cosmopolitan family of moths formerly classified as a sub­ family of the Pyralidae. Adult sod webworms are similar to pyralid moths in having abdominal tympanal organs, and elongate, scaled labial palpi extend­ ing in front of the head to form a proboscis. However, sod webworm moths can be distinguished by their habit of holding the wings close around the body when at rest (Borror et al. 1989). The larvae of these moths feed pri­ marily on plants in the family Graminae and, as a result, several species are considered economically important pests of turfgrasses (Cobb 1995), cereal, and forage crops (Ainslee 1922). Some species of sod webworms are also known to feed on tobacco (Bohart 1947), cranberries (Franklin 1950), and coniferous nursery stock CKamm et al. 1983), Both univoltine and multivoltine species are known and, although there are exceptions (e.g., the cranberry girdler, Chrysoteuchia topiaria(Zeller», the univoltine species are generally somewhat less important economically. Ofthe six most economically important temperate sod webworm species, four

lDepartment of Entomology, The Ohio State University, Ohio Agricultural Re­ search and Development Center, Wooster, OH 4469L 2Department ofEntomology, The Ohio State University, Columbus, OH 43210. 174 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4 are multivoltine (Vittum et al. 1999). The multivoltine species are believed to be more important economically primarily because of their numbers and pro­ longed period ofactivity. While nearly 100 species of sod webworms are known in North America (Bohart 1947), the species complex and occurrence of these moths has only been reported from Florida (Ainslee 1923, 1927; Kimball 1965), Iowa (Decker 1943), New York (Forbes 1923), North Carolina (Brimley 1938, 1942), Oregon (Prescott 1965), Tennessee (Matheny and Heinrichs 1975), and Virginia (Tol­ ley and Robinson 1986). Like many of the Great Lakes States, there is little published record of the sod webworm species of Ohio. Therefore, the species complex and seasonal occurrence of this group is relatively unknown in this region. The goal of this study was to document the species complex and seasonal flight activity of the common sod webworm moths of Ohio. In this paper, we consider the univoltine and multivoltine species separately because of the more evident economic significance of the latter. Using black light trap data collected from four different locations in Ohio, we identified the species com­ plex and assembled seasonal flight activity profiles for all sod webworm species collected over the course of three to five years (16 total trap years). Supplemental data were gathered from the insect collection housed at The Ohio State University, Museum of Biological Diversity located in Columbus, OH. Together, these data provide a biological record of this common, but often overlooked group of insects, and clarify the seasonal occurrence of sev­ eral species in this part ofthe Great Lakes region.

:MATERIALS AND METHODS Collection sites and dates. Collection sites represented the northern, central, and southern latitudes of the state (Figure 1). The northern Ohio col­ lection site was established at the main campus of the Ohio Agricultural Re­ search and Development Center (OARDC) in Wooster (Wayne Co.). Wooster lies within the Walhonding watershed and data were collected from the Wooster site from May, 1978 through October, 1982 (5 years) two central Ohio collection sites were used, both within the Upper Scioto watershed. The first site was located at Milford Center, in Union County. Data were collected from the Milford Center site from May through October of 1978, 1979, and 1982 (3 years). The second central Ohio site was located at the TruGreen Chemlawn research facility in Delaware (Delaware County). Data were col­ lected from the Delaware site from May, 1986 through October, 1989 (4 years). The southern Ohio site was located at the Southern Branch of the OARDC in Ripley, OH (Brown Co.). Brown County lies within the Ohio Brush-Whiteoak watershed and data were collected from the Ripley site from May, 1978 through October, 1981 (4 years). No attempt was made to quantify the vegetation at any of the sites although all areas were predominantly sur­ rounded by a matrix of agricultural fields and forest fragments. Additionally, the Delaware site was surrounded by 40 acres of managed turfgrass which was further surrounded by forest and agricultural land. Major agricultural crops include corn, soybeans, alfalfa, and wheat in the northern and central regions, and corn, alfalfa, and tobacco in the southern region. Collecting methods and data management. Collection was per­ formed using Ellisco® light traps with 15 W blacklight tubes. One light trap was placed at each of the collection sites and was on from dusk until dawn during all collecting seasons. At the Wooster, Millford Center, and Ripley lo­ cations, one insect strip (Vapona®) was used in each trap to kill captured 2000 THE GREAT lAKES ENTOMOLOGIST 175

N + Figure 1. Map of Ohio showing location of collection sites used to assess sod webworm flight activity in Ohio (e). Collections were made at the Wooster site during 1978-82, at the Delaware site during 1986-89, at the Milford center site during 1978, 79, and 82, and at the Ripley site during 1978-81. moths and strips were replaced regularly. At the Delaware location, an open container of ethyl acetate was placed in the trap head as the killing agent and was refilled daily. Captured moths were collected daily (except week­ ends) and kept frozen until identification, sorting and counting could be per­ formed. Sod webworm adult species identifications were verified by D. C. Ferguson (Systematic Entomology Laboratory, USDA, Washington, D. C.). 176 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

Voucher. specimens were deposited in the insect collection at the OARDC. Data were organized by year, calendar date, and location, and comparisons between locations were based on the average total number of each species collected per year.

RESULTS AND DISCUSSION Univoltine species. Seven species and three genera of univoltine sod webworm moths were collected during the course of the study although there were considerable differences in local abundance. Only four of the seven uni­ voltine sod webworm species collected were regularly found at all four sites. These species include Cram bus laqueatellus Clemens, Crambus agitatellus Clemens, Chrysoteuchia topiaria (Zeller), and Agriphila vulgivagella (Clemens). Based on the average total number of moths collected per year, C. laqueatellus was generally more abundant at the Milford Center and Ripley sites than at either of the other two sites (Figure 2) Crambus agitatellus was roughly equally abundant at the Milford Center, Delaware, and Ripley sites but was much less common at the Wooster site (Figure 3). Chrysoteuchia top­ iaria was most abundant at Delaware followed in order by Milford Center, Ripley, and Wooster (Figure 4). A. vulgivagella was most abundant at the Delaware and Ripley sites followed by the Milford Center and Wooster sites. Wooster recorded the lowest abundance of all four of these common species.

w~~(1978-82) I ::1 ." IIII Delaware ;l!~.,~ ~ .. , (1986-89) 20r------. Milford Center (1978,79,82) ':1 IJ.W __ ..

~I~~o I I I (1978-81Ripley ) ~I~~J~-L_,---~~~__~--__~~ MAY JUN 15 JUL 15 AUG 15 SEP 15 OCT 15

Day ofYear Figure 2. Seasonal flight activity of Crambus laqueatellus Clemens at four locations in Ohio as determined by the average number of moths caught in black light traps on each date. •

2000 THE GREAT LAKES ENTOMOLOGIST 177

Delaware ~ ~ :1 J I (1986-89) _L, • 1~ .~~ 2~ri______~ ______'Le." -. Z .E Milford Center eli0) § 10 (1978, 79, 82) ~ 15r------~c_------__. Ripley 10 (1978-81)

oL~..1-..uIU-. MAY JUNI5 lUL 15 AUG 15 SEP 15 OCT 15

Day ofYear Figure 3. Seasonal flight activity of Crambus agitatellus Clemens at four lo­ cations in Ohio as determined by the average number of moths caught in black light traps on each date.

Wooster (1978-82)

r-. I I..~J ..,.. " CIl Delaware -S .~ 0 100 (1986-89) ;:?; .~ ~ ..... '"C0 0 ...,.Sl l ...... 0) tJ .0 " 20 Milford Center ~ ~[ (1978, 79, 82) ~ ~ 10 D o ...1. ,.,liJl&II., "- .. Ripley ~[hi~£~... (1978-81 ) MAY JUN15 JUL 15 AUG 15 SEP 15 OCT 15

Day ofYear Figure 4. Seasonal flight activity of Chrysoteuehia topiaria (Zeller) at four locations in Ohio as determined by the average number of moths caught in black light traps on each date. 178 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

J (1978-82) ~10:0IJ ______~__...aLL_w_oo~st_~~ til d~:~,

~? -_~. '2 00! I Delaware -: ~ _ .wA, (:986-89) i ~ 0 ~=-=-=-=-=-=-=-=-=-=-=-=-==-=-=-=-=-=-==-=-=-=:~~~~~~':=-=-~ Z,g 4j) I ~Milford Cent~ a:9. 200 (1978,79,82) ~ ~ L______--~--____--~~~.~4~..__~!Lu·••L-~~

(1978-8J) ~ 36:0l__~______~______~~J.-...... R~i~PI_~ __~ MAY JUN 15 JULI5 AUG 15 SEPI5 OCT 15

Day ofYear Figure 5. Seasonal flight activity of Agriphila vulgivagella (Clemens) at four locations in Ohio as determined by the average number of moths caught in black light traps on each date.

Three of the seven species collected were found regularly only at the Delaware site (Figure 5). These species included Crambus caliginosellus Clemens, Crambus luteolellus Clemens, and Agriphila ruricolella (Zeller). Based on the average total number of moths collected per year, C. caligi­ nosellus was by far the most abundant of these species followed in order by C. luteolellus, and A ruricolella. Although all three of these species may have been collected at the other sites, their numbers were extremely low. This problem could have been exacerbated by handling practices. Because sod webworm moths tend to be small and delicate, excessive handling, freezing, and shipping made many specimens impossible to identify. Future efforts di­ rected at collecting this group of moths should also explore alternative meth­ ods since blacklight traps may not be the most effective way to collect all species. There appeared to be a trend of earlier to later emergence corresponding to latitude for C. laqueatellus, C. agitatellus, and C. topiaria but notA uulgi­ uagella. This observation implies that while development and adult flight of the prior three species is mainly temperature dependent, the later species may rely more heavily on other environmental cues such as photoperiod. In­ deed, Tolley and Robinson (1986) determined that calendar date was a better flight peak predictor than degree-days for many sod webworm species in Vir­ ginia. Further research will be necessary to determine the relative impor­ tance ofvarious environmental factors in determining seasonal flight activity for each species. Based on seasonal flight activity profiles, there appear to be three dis­ 2000 THE GREAT LAKES ENTOMOLOGIST 179 10.------, Delaware ~ Jl1 (1986-89) Iii"' 5 I ~ ~~. ~ OL ______~aw·'·~L,.IU...... I.1.1.I~I~J~~J~~... , .. ____--__----______~

Delaware (1986-89)

Day ofYear Figure 6. Seasonal flight activity of Crambus luteolellus Clemens, Crarnbus caliginosellus Clemens, and Agriphila ruricolella (Zeller) at Delaware, Ohio, as determined by the average number of moths caught in black light traps on each date. tinct life-history strategies employed by these univoltine moths: early emer­ gence, mid-season emergence, and late emergence. While the majority of species (four) can be considered mid-season emergers (C. agitatellus, C. topi­ aria, C. caliginosellus, and C. luteolellus), one species, C. laqueatellus, falls into the early emergence category, and two species can be considered late season emergers (A. uulgiuagella and A. ruricolella). These different emer­ gence periods may result in a temporal partitioning ofavailable resources be­ tween similar species thereby limiting interspecific competition. Multivoltine species. Five species representing five different genera of multivoltine sod webworms were collected during the course of the study (Figs.7-11). However, numbers varied between sites and only four species were collected at all of the collection sites: Parapediasia teterrella (Zincken), Pediasia trisecta (Walker), Fissicrambus mutabilis (Clemens) and Micro­ crambus elegens (Clemens) (Table 2). Crambus praefectellus (Zincken) was notably absent from the Ripley site and was the least abundant species at both Wooster and Milford Center although it was far more abundant at the Delaware site. P. teterrella, P. trisecta, and F. mutabilis were most abundant at the Delaware site followed in decreasing order by Milford Center, Ripley, and Wooster. Although M. elegens was most abundant at the Delaware site as well, its abundance at Wooster was a close second followed by Milford Center and Ripley. Based on the average total number of moths collected per loca­ tion per year, the most abundant species overall was P. teterrella, followed in decreasing abundance by P. trisecta, F. mutabilis, lltl. elegens, and C. prae­ fectellus. 180 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 & 4

40~------, Wooster 20 (1978-82)

oL---~~"~~~~~~"'"

300200 • I Delaware

1.1''k.....I ...... L...... 10: L~+...... "-.JI., 1 JJul,J"II~."•• I~.... q~'''H''''.&~ LJ.ULI.... ~L..98_6_.8_9)_

I Milford Center .... ,,kM,JJIr.,wL,..,, ... 1"L :978,79,82) 12:0:ol____~~.. ~~~~~~~..~I,U...... h ~~~ ______40------­ 30 Ripley 20 (1978·81) 1~L---u.IM....~..~~~ MAY JUNI5 JUL IS AUG 15 SEP IS OCT IS

Date Figure 7. Seasonal flight activity of Parapediasia teterrella (Zincken) at four locations in Ohio as determined by the average number of moths caught in black light traps on each date.

150100 • Delaware ,I , 1.1 (1986-89) 5:L-~~~~JU~••~~~~.~~~~,~t.~~Lt...~II ..l.1~'H~I~~~I4LeIWU'~____ ~

::: I Milford Center SO JJ~ (1978, 79, 82) OL-____~~~~._w ..~h~~~ ______~~+~L~,L~J~~~_£I.wu.~~______~

Date Figure 8. Seasonal flight activity of Pediasia trisecta (Walker) at four loca­ tions in Ohio as determined by the average number of moths caught in black light traps on each date. 2000 THE GREAT LAKES ENTOMOLOGIST 181

w~~e~ (1978-82)j J i~[LJ'~15 Delaware 10 (1986-89)

o~-~~~~~~~~~~~~

12~·- ... ----~---- .....---....~.....------­ 8 1 I Milford Center J4 l._ .lIILM ~JI rJ J..,..II ~I • ~:978, 79, 82) S,...-·· --_.....

i ~ Ripley . 4! (1978-81)] ol----'~--L--..~~~ MAY JUN 15 JUL 15 AUG 15 SEP 15 OCT 15

Date Figure 9. Seasonal flight activity of Fissicrambus mutabilis (Clemens) at four locations in Ohio as determined by the average number of moths caught in black light traps on each date.

..... ~~~~~2) I

.~...... -...... ­....lII...-~.. 1...... h..,U.l...... illIIILL__~ j ~ Delaware '­ t (1986-89) o ~ 1i 1.0 ­ .....----­...... -­..---.....-. ---....---­...... ----.~....­ Z S Milford Center

i ~ ::: L.L~Jl.. ~ I.~L • J.. IIL..I ~1~1,97~::J ;( :[ -----.~--- ....--.-...--.... Ripley (1978-81) • .,.1 Ilo,I,JJ.. •.• MAY JUN 15 JUL 15 AUG 15 SEP 15 ocr 15

Date Figure 10. Seasonal flight activity of Microcrambus elegens (Clemens) at four locatioIls in Ohio as determined by the average number of moths caught in black light traps on each date. -

182 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

Wooster (19"."J

Delaware (1986-89)

.... 4~~ ~:~~k.~; I ~I....J...L..II~d~---,I_ :L~MAY JUN 15..... JUL15 AUG 15 SEP15 OeTtS Date

Figure 11. Seasonal activity of Crambus praefectellus (Zincken) at three locations in Ohio as determined by the average number of moths caught in black light traps on each date.

TABLE 1. Average number of seven different univoltine sod web worm species col­ lected per year* at four locations in Ohio (total number collected/number of years col­ lected). Species Wooster Delaware Milford Center Ripley

~..... Agriphila vulgivagella 371 937 533 921 Agriphila ruricolella 5 Chrysoteuchia topiaria 131 2,713 1,041 1,107 Crambus agitatellus 64 1,095 300 819 Crambus caliginosellus 1,186 Crambus laqueatellus 19 108 154 256 Crambus luteolellus 251 "'Number ofyears varies with collection site; Wooster f, years, Delaware = 4 years, Milford Center =3 years, and Ripley", 4 years. 2000 THE GREAT LAKES ENTOMOLOGIST 183

Table 2. Average number of five different multivoltine sod webworrn species collected per year* at four locations in Ohio (total number collected/number ofyears collected). Species Wooster Delaware Milford Center Ripley Parapediasia teterrella 625 5,210 1,564 1,016 Pediasia trisecta 203 2,603 1,283 618 Fissicrambus mutabilis 11 241 113 87 Microcrambus elegens 116 149 92 23 Crambus praefectellus 6 196 4 *Number ofyears varies v,ith collection site; Wooster 5 years, Delaware 4 years, Milford Center 3 years, and Ripley =4 years.

Table 3. DateJLocality records for sod webworrn specimens in The Ohio State Univer­ sity insect collection-Columbus, OH. Species Location Date (month/year) ------Agriphila ruricolella (Clemens) Granville, OH August 1929 Agriphila vulgivagella, (Clemens) Granville, OH August 1929 Columbus, OH September 1942 Arequipa turbatella (Walker) Granville, OH July 1930, July 1989 Crambus alboclavellus Zeller Granville, OH June, July and August 1930 Crambus agitatellus Clemens Columbus, OH July 1898 Crambus albellus Clemens Granville, OH June 1930 Crambus caliginosellus (Clemens) Granville, OH June 1930 Crambus girardellus Clemens Granville, OH June 1930 Crambus hortellus Hubner Granville, OH June 1930 Crambus laqueatellus Clemens Columbus, OH Granville, OH June 1930 Crambus luteolellus Granville, OH June 1930 Crambus pusionellus Zeller Sugar Grove, OH July 1920 Crambus zeellus Fernald Black Hand, OH July 1930 Fissicrambus mutabilis (Clemens) Granville, OH June 1930 Microcrambus (Clemens) Granville, OH July 1929 Parapediasia teterrella (Zincken) Granville,OH August 1929 Parapediasia decorella (Zincken) Granville, OH June 1930 Columbus, OH July 1897 Pediasia trisecta (Walker) Granville,OH August 1929, June 1930 Urola nivalis (Drury) Oak Harbor, OH June 1925 Granville, OH June 1929 Oak Openings, OH July 1975 *-=no data available r

184 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

Of the five species collected, four are considered important pest species: P. teterrella, P. trisecta, F. mutabilis and C. praefectellus (Vittum et al. 1999). Knowledge ofthe seasonal flight activity of pest species is particularly impor­ tant [, .. of monitoring and intervention. Flight activity profiles for some s strate differences in the number ofgenerations occurring be­ tween sites. For instance, P. teterrella appeared to have at least a partial third generation at Wooster and Delaware while a third generation was obvi­ ous at Ripley. This same trend was true ofP. trisecta with close to a full third generation appearing in some years at Ripley. These differences in voltinism are not overly surprising and may be due to local weather patterns or micro­ climate effects. Indeed, other authors have reported similar findings. For in­ stance, P. trisecta is reported to have one generation per year in the Pacific Northwest, two generations in the Midwest, and three in Virginia (Ainslie 1927, Crawford and Harwood 1964, Robinson and Tolley 1982). Similarly, P. teterrella is reported to have two generations in Virginia and three genera­ tions in Tennessee (Robinson and Tolley 1982, Ainslie 1930). Our data indi­ cate that both P. teterrella and P. trisecta may regularly have three genera­ tions in Ohio, a fact that was previously undetermined.

ACKNOWLEDGMENTS The authors thank Keith Kennedy, Jeff Rodencal, Patrice Suleman, and Dan Chandler for technical assistance in handling, collecting and sorting moths. Funding for this work was provided, in part, by the Ohio Turfgrass Foundation, and by State and Federal funds appropriated to the Ohio Agri­ cultural Research and Development Center.

LITERATURE CITED Ainslie, G. G. 1922. Webworm injurious to cereal and forage crops and their control. USDA. Farmers Bull. 1258: 1-16. Ainslie, G. G. 1923. The Crambinae of Florida. Fla. Entomo!. 6: 49-55. Ainslie, G. G. 1927. Additions and corrections to the list of Crambinae of Florida. Fla. Entomol. 11: 12-14. Ainslie, G. G. 1930. The bluegrass webworm. USDA Tech. Bull. 173: 1-25. Bohart, R. M. 1947. Sod webworm and other lawn pests in California. Hilgardia 17: 267-308. Borror, D. J., C. A. Triplehorn and N. F. Johnson. 1989. An introduction to the study of insects, 6th ed. Saunders College, Chicago. Brimley, C. S. 1938. The insects of North Carolina. North Carolina Dept. Agric. 560 pp. C. S. 1942. Supplement to the insects of North Carolina. North Carolina Dept. Cobb, P. Sod webworms, pp. 86-88. In: R. L. Brandenburg and M. G. Villani Ceds.), Handbook of turfgrass insect pests. Entomol. Soc. Am., Lanham, MD. Crawford, C. S. and R. F. Harwood. 1964. Bionomics and control pf insects affecting Washington grass seed fields. Wash. Agric. Exp. Sta. Tech. Bull. 44: 1-25. Decker, G. A. 1943. An annotated list of Crambid moths known to occur in Iowa. Proe. Iowa Acad. Sci. 50: 337-339. Forbes, W. T. M. 1923. Lepidoptera of New York and neighboring states. Cornell Univ. Agric. Sta. Mem. 68. Franklin, J. 1950. Cranberry insects in Massachusetts. Mass. Agric. Exp. Sta. Tech. Bull. 44 .

• 2000 THE GREAT LAKES ENTOMOLOGIST 185

Iftner, D. C., J. A. Shuey and J. V. Calhoun. 1992. Butterflies and skippers of Ohio. Ohio Biological Survey, new series 9(1). Kamm, J. A, P. D. Morgan, D. L. Overhulser, L. M. McDonough, M. Triebwasser and L. N. Kline. 1983. Management practices for cranberry girdler (Lepidoptera: Pyrali­ dae) in Douglas-fIr nursery stock. J. Econ. Entomol. 76: 923-926. Kimball, C. P. 1965. The Lepidoptera of Florida; an annotated checklist. Fla. Dept. Agric. Matheny, E. L. and E. A Heinrichs. 1975. Seasonal abundance and geographical distri­ bution of sod webworms (Lepidoptera: Pyralidae: Crambinae) in Tennessee. J. Tenn. Acad. Sci. 50: 33-35. Prescott, H. W. 1965. Report on Oregon crambid moth collection by Forest Grove Sta­ tion. U.S. Dept. Agric., Agric. Res. Servo Spec. Rep. 10-227. Rings, R. w., E. H. Metzler, F. J. Arnold, and D. H. Harris. 1992. The owlet moths of Ohio (Lepidoptera: Noctuidae). Ohio Biological Survey, new series 9(2). Tolley, M. P. and W. H. Robinson. 1986. Seasonal abundance and degree-day prediction of sod webworm (Lepidoptera: Pyralidae) adult emergence in Virginia. J. Econ. Ento­ mol. 79: 400-404. Robinson, W. H., and M. P. 'fuIley. 1982. Sod web worms associated with tUrfgrass in Virginia. Am. Lav,'TI Appl. 3: 22-25 Scott, J. 1986. The butterflies of North America: A natural history and field guide. Stanford University, Stanford, CA Vittum, P. J., M. G. Villani and H. Tashiro. 1999. Turfgrass insects ofthe United States and Canada. Cornell University, Ithaca, NY. 2000 THE GREAT LAKES ENTOMOLOGIST 187

LEUCANTHIZA DIRCELLA [LEPIDOPTERA: GRACILLARIIDAE): A LEAFMINER OF LEATHERWOOD, DIRCA PALUSTRIS

Toby R. Petrice I, Robert A. HaackI, William J. Mattson2 and Bruce A. Birr2

ABSTRACT Leatherwood, Dirca palustris (Thymelaeaceae), is an understory shrub ranging throughout most of the eastern and central United States and adja­ cent Canada. During 1997-1999, we conducted studies to identifY and assess the impact of a leafminer that was causing significant damage to leather­ wood plants in eastern Gogebic County, Michigan. Leucanthiza dircella was identified as the only insect responsible for the leaf mining activity on leatherwood. In northern Michigan, L. dircella completed one generation per year. Adult moths were captured on yellow sticky panels suspended from leatherwood branches. In 1997 and 1998, most adults were captured during the first sampling period of each year: 6-12 June 1997 and 3-19 May 199B. In 1999, no moths were collected during 5-29 April but adults were collected between 30 April and 22 June 1999. In 1999, initiation of adult flight coin­ cided with D. palustris leaf flush. In 1997, leafmines were very noticeable by 30 June. The mean number of live L. dircella larvae per mine was 3.5 on 17 July 1997 and then decreased as the season progressed, with most larvae having exited the mines by late August to pupate in the soil. In late August 1997, the mean surface area of a single leaf was 17.8 cm2 and the mean sur­ face area of a single mine was 5.9 cm2. At the end of the 1997 growing sea­ son, 31% of the leatherwood leaves contained L. dircella mines, and 11% of the total leaf surface area had been mined. In 1999, only 8% of the leaves in the study area contained L. dircella mines. No leatherwood mortality was ev­ ident as a result of L. dircella leaf mining. Seven species of hymenopteran parasitoids were reared from L. dircella larvae, including one braconid in the genus Pholetesor and six eulophids in the genera Chrysocharis, Closterocerus, Pnigalio, and Sympiesis. Three coleopterans that were com­ monly observed on leatherwood plants during all years included: Glyptina brunnea (Chrysomelidae), Phyllobius oblongus (Curculionidae) and Poly­ drusus sericeus (Curculionidae).

Leatherwood, Dirca palustris 1. (Thymelaeaceae), is an understory shrub that grows to a height of 1-3 m and ranges throughout most of the eastern and central United States and adjacent Canada (Fernald 1950, Vogelmann 1953). Leatherwood is most often found growing in nutrient rich, mesic hard­ wood stands (Britton and Brown 1970, Cleland et al. 1993, Kotar et al. 1988,

lUSDA Forest Service, North Central Research Station, 1407 S. Harrison Rd., Michigan State University, East Lansing, MI 48823. 2USDA Forest Service, North Central Research Station, 5985 Highway K, Rhinelander, WI 54501. 188 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

Radford et al. 1979). The common name leatherwood is earned from the plant's tough fibrous bark, which Native Americans utilized as a source of fiber for making items such as mats, bags, cords, and rope (Whitford 1943). Leatherwood contains toxins that cause irritation and blistering when contacted by skin, and gastrointestinal discomfort when swallowed (Fuller and McClintock 1986, Muenscher 1964). Because of these properties, there has been some preliminary research exploring the potential of leatherwood extracts as deterrents for vertebrate herbivores (Ramsewak et al. 1999, Za­ sada et al. 1999). Interestingly, there are some references of leatherwood being used by early settlers for medicinal purposes (Fielder 1975). An oint­ ment was made with leatherwood bark and sarsaparilla (Aralia sp.) for treating severe skin diseases, and a tea made from the leatherwood roots was used for treating kidney problems. Moreover, a study was conducted dur­ ing the early 1900's, testing the efficacy of leather-wood extracts as a purga­ tive (Lecours 1924). Leatherwood has been planted as an ornamental shrub because its early spring bloom, tree-like appearance, and tolerance to full sun make it very suitable for shrub beds and rock gardens (Esson 1949, Tredici 1983). Tredici (1983) also noted that leatherwood is easily germinated from seeds that have been subjected to a cold period. Although uncommon throughout most of its range, when leatherwood does occur in the forest understory, it frequently grows in dense patches (Nevling 1962). This is often the case in mature hardwood forests in northern Wisconsin and the western Upper Peninsula of lVIichigan, where leatherwood may be an important understory component (Curtis 1959, Kotar et al. 1988, John C. Zasada, USDA Forest Service, Rhinelander, WI, pers. obs.). During 1997-1999, we conducted a study to identify and determine the impact of any insects that were causing significant leaf mining damage to leatherwood plants in eastern Gogebic County, Michigan.

MATERIALS AND METHODS Studies were conducted during 1997-1999 on the Ottawa National For­ est, Watersmeet Ranger District, near Taylor Lake in eastern Gogebic County, Michigan (Fig. 1). The area selected was classified as mature hard­ wood forest and was composed primarily of sugar maple (Acer saccharum), yellow birch (Betula alleghaniensis), and American basswood (']'ilia ameri­ cana). Leatherwood was abundant in the understory and had undergone heavy leaf mining damage during the early 1990's (John C. Zasada, pers. comm.). In 1997, from 6 June through 25 August, we attached one double­ sided yellow sticky panel (15 cm by 30 em) on each of six leatherwood plants, 1 to 2-m tall, to capture adult leafminers. Sticky panels were collected at two-week intervals, with the exception of the first two sample periods which were collected at six-day and four-day intervals, respectively. Each sticky panel was placed in a plastic bag, and frozen for later inspection. During each site visit, we visually examined leatherwood foliage for ovipositing in­ sects and leaf mining activity. We collected specimens of suspect insects that belonged to leaf mining families. We also collected leaf samples from ten leatherwood plants at two-week intervals beginning 17 July 1997. Each sam­ ple consisted ofone branch tip per plant, ca. 20-cm in length. Each branch tip typically had 6 to 12 leaves and included 2 to 3 year's growth. We selected a branch tip that appeared to represent the leaf mining activity currently found on each individual plant being sampled. Leaf samples were placed in­ 2000 THE GREAT lAKES ENTOMOLOGIST 189

Figure 1. County outline map of Michigan, with dots representing the ap­ proximate location of (1) the leatherwood study site in Gogebic County, ca. 46° 15' N Lat, 89° 3' W Long; (2) the nearest weather station in Watersmeet, Gogebic County, ca. 46° 17' N Lat, 89° 10' W Long; and (3) the weather sta­ tion in Stambaugh, Iron County, ca. 46° 3' N Lat, 88° 37' W Long. Darkened area on insert map indicates the location of Michigan within the United States. dividually in plastic bags, and stored on ice for later processing in the labora­ tory. On 25 August 1997, after leaf mining had ceased and most larvae had ex­ ited the mines, we collected leaf samples in a 200-m by 100-m area surround­ ing the central study site. We sampled 110 leatherwood plants, taking one branch sample from each plant that was 1 m in height or taller. Each branch sample was chosen to be representative of the leaf mining damage level for the particular plant being sampled. In the laboratory, the 1997-collected leaves that contained mines were held over a strong light, allowing the number oflarvae inside of each mine to be counted. We also recorded if mines contained holes through which larvae could have exited to pupate in the soil. In addition, we noted if leaves pos­ 190 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 & 4

sessed fungal rust spots and if the section of branch sampled had evidence of other insects such as scale insects. Mter the leaves were examined, they were photocopied so the total leaf surface area and total mined area could be determined. Photocopying allowed for a simple means of leaf "preservation" and it accurately depicted mined and non-mined areas of each leaf. Using scissors, we cut out each photocopied leaf and passed it through a leaf area meter to determine total surface area. Each paper "leaf' was measured twice and the average surface area recorded. We then removed and measured the mined area of each photocopied leaf. The same procedure was followed for the leafsamples collected from the 110 plants on 25 August 1997. The number of larvae per mine and percent of mines with exit holes were analyzed for differences among sample dates using a one-way-ANOVA (PROC GLM, SAS Institute 1990). For certain statistical analyses, leaves were first placed in 10-cm2 leaf size classes. A one-way-ANOVA (Proc GLM) was used to test for differences in percent of leaves mined and percent of leaves with rust spots among leaf size classes. Arcsin square-root transfor­ mations were performed on all percentage data prior to analyses. Tukey's Studentized Range Test was used to separate means when ANOVA was sig­ nificant. A significance level of P :s; 0.05 was set for all analyses. Leaves containing larvae were placed on moistened paper towels inside of plastic bags in 1997. We used a pin to make small holes in the upper sur­ face of the bags to reduce molding. The bags were examined at least twice weekly for insects emerging from the leaf mines. Leafminer pupae were re­ moved and placed in petri dishes on moistened paper towels. Hymenopteran parasitoids emerging from the leaf mines were placed in vials containing 70% ethyl alcohol and were later sent to specialists for identification. Leafminer pupae were overwintered at 4°C for 120 days. Adult leafminers emerging from overwintered pupae were pinned and used as reference speci­ mens for identifying insects captured on the sticky panels. The number of adult leafminers captured on each sticky panel was recorded. Voucher speci­ mens of leafminer adults and hymenopteran parasitoids are currently stored in the USDA Forest Service insect collection, Michigan State University, East Lansing, Michigan. In 1998, we placed one sticky panel (15 cm by 30 cm) on each of 20 leatherwood plants from 3 May through 3 November. Sticky panels were col­ lected on 19 May, 1 June and 7 July. We replaced sticky panels on 7 July to monitor any mid- to late-season adult leafminer activity; these traps were collected 3 November 1998. In 1999, we placed one sticky panel cm by 30 cm) on each of 12 leatherwood plants from 5 April through August and samples were col­ lected on 12 April, 22 April, 29 April, 10 May, 17 May, 27 May, 4 June, 15 June, 22 June and 4 August. The last sample period was longer than earlier periods in 1999, and was meant to monitor any mid- to late-season adult leafminer activity. On 25 August 1999, we sampled one branch from each of 22 leatherwood plants in the study area. For each branch, we recorded the number of mined and unmined leaves. Samples were collected from the same area sampled in 1997 and branches were selected following the same protocol as used in 1997. Also in 1999, leatherwood shoot and leaf development were monitored on three leatherwood plants. On each plant, we selected a branch tip in each of the four cardinal directions and marked the terminal bud ofeach with a plas­ tic tie. Beginning 29 April, we examined each marked bud and recorded cur­ rent-year's shoot length and length of the largest leaf on that shoot during each sticky panel collection through 22 June (see above). Also, beginning 10 May through 15 June, we recorded the number of leaves present on each of 2000 THE GREAT LAKES ENTOMOLOGIST 191 the marked current-year's shoots. Shoot length, leaf length, and the number of leaves present were analyzed for differences among sample dates using a mixed model ANOVA (PROC MIXED, SAS Institute 1990), with shoots within leatherwood plants assigned as a random effect of the model. Least squared means significant at the P.:$; 0.05 level were separated Tukey's Studentized Range Test. We obtained official weather records from the National Oceanic and At­ mospheric Administration (NOAA, Asheville, North Carolina), for Wa­ tersmeet, Gogebic County, Michigan, which is approximately 10 km west of the study site (Fig. 1). Occasionally, weather records were not reported by the Watersmeet weather station. When that occurred, we used data from the next closest recording station, which was Stambaugh, Iron County, Michigan, located approximately 40 km east of the study site (Fig. 1). Daily heat sums were calculated for 1997, 1998 and 1999 using the averaging method, i.e. av­ erage of daily high and low temperature subtracted from base temp, and the Baskerville-Emin method (Baskerville and Emin 1969). Heat sums for each method were calculated using base 5°C and base lOoC. Biology. All adult leafminers emerging from the overwintered pupae were identified as Leucanthiza dircella Braun (Lepidoptera: Gracillariidae) (Fig. 2) by Ronald Priest (Michigan State University, adjunct curator). This moth has been reported from Clermont County, Ohio (Braun 1914, Forbes 1920), and from Michigan (Nielsen 1998). Most L. dircella moths were captured in May and early-June during all three sample years, although the number of moths captured varied greatly between years. In 1999, adults were first captured between 30 April and 10

Figure 2. Leucanthiza dircella adult that emerged in the lab 15 July 1999 from a larva collected 21 May 1999 in Ingham County, Michigan (total wingspan approximately 6.2 mm). Specimen provided by Ronald Priest. 192 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

Table 1. Mean number of Leucanthiza dircella adults collected per yellow sticky trap (2-sided; each side 15 em by 30 em) per sampling day and heat sums for 1997, 1998, and 1999. Heat sums (start date", March 1) were calculated using the averaging method (Averaging) and Baskerville-Emin method (B-E) for the bases 10°C and 5°C. Heat sums Heat sums Sampling Julian Adults! base 10°C* base 5°C** period days trap/day Averaging B-E Averaging B-E 1997 (41 moths collected on 6 traps) 6 June-12 June 157-163 0.97 66-104 104-144 199-266 221-285 13 June-17 June 164-168 0.07 112-125 152-170 279-313 298-330 18 June-30 June 169-181 0.03 131-246 175-291 323-498 340-509 1 July-17 July 182-198 0.01 257-377 301-428 514-713 524-715 18 July-29 July 199-210 0.00 387-475 439-529 728-871 730-866

1998 (631 moths collected on 20 traps) 3 May-19 May 123-139 2.03 12-60 38-106 66-178 100-219 20 May-l June 140-152 0.16 65-83 112-145 188-245 228-290 2 June-7 July 153-188 0.01 83-2M 147-306 249-545 295-582

1999 (21 moths collected on 12 traps) 5 April-12 April 95-102 0.00 0 7-8 8-9 24-31 13 April-22 April 103-112 0.00 0 8-12 9-14 32-44 23 April-29 April 113-119 0.00 0-1 12-23 14-30 44-67 30 April-10 May 120-130 0.02 1-32 25-64 33-104 72-142 11 May-17 May 131-137 0.02 32-48 66-87 110-149 147-188 18 May-24 May 138-144 0.04 50-57 90-102 157-184 194-223 25 May-3 June 145-154 0.03 57-94 102-145 184-253 223-292 4 June-15 June 155-166 0.05 99-197 150-249 263-411 301-444 16 June-22 June 167-173 0.01 197-226 250-282 416-473 449-503 23 June-4 August 174-216 0.002 237-604 293-665 489-1066 519-1073 *Heat sums as ofend ofeach day. **Two L. dircella adults were captured during this sample period but the number cap­ tured per trap per day was < 0.01.

May when Baskerville-Emin heat sums (base lOcC; start date = 1 March) were between 25 and 64, respectively. The greatest number of L. dircella adults was captured on sticky panels between 6 to 12 June 1997 and between 3 to 19 May 1998. Trap catches were consistently low in 1999 (Table 1). Given that we captured adults during the first sampling periods in 1997 and 1998, it is likely that adult flight initiated before sticky panels were deployed during those two years. Pooling the 1997 and 1998 data, L. dircella capture rates were highest when Baskerville-Emin heat sums were between 38 and 144 (base lOoC; start date = 1 March; Table 1). For comparison, heat sums calculated with the Baskerville-Emin method base 5°C and heat sums calculated with the averaging method using bases cc and lOoC are also in­ cluded in Table 1. As expected, the mixed model ANOVA for Dirca palustris shoot lenP,th and leaf length varied significantly among sampling dates (P < 0.0001). The first evidence of flushing occurred 29 April 1999 when bundles of small leaves were found protruding from the buds. Shoot elongation was completed by 17 May and leaves were fully expanded between 24 May and 3 June 1999 (Fig. 3). The number of leaves per shoot averaged 5.8 on 10 May 1999, and 2000 THE GREAT LAKES ENTOMOLOGIST 193

a A 6 - ~ b 0 b - 0 .-:t .c III ... 4 a.CD - III CD > 'II .9! 2 ­ '0 zci 0 B 100 - ab a b ab 80 - E c g 60 .c - d CI r-=­ - c 40 - .9! 'Ii CD 20 ...J - 0 n c 60 - E a a a a §. a ;; cCI 40 - .9! 0 b - 0 .c 20 U) -

0 0

Figure 3. Mean (+ 1 SE) (A) number of Dirca palustris leaves per current­ year's shoot during the period of 10 May through 15 June 1999, (B) length of the longest leaf on a current-year's shoots during the period of 29 April through 22 June 1999, and ee) current-year's shoot length during the period 10 May through 22 June 1999 based on 12 shoots on 3 plants (4 shoots per plant). Means with the same letter are not significantly different at P s 0.05 (Tukey's Studentized Range Test). *Only a tight bundle of leaves protruded from buds on 29 April; leaf length measured from tip of longest leafto base of leafbundle; new shoots were not visible. 194 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

Table 2. Mean (±SE) number of live Leucanthiza dircella larvae found per leaf mine and the percent of leaf mines with holes (through which L. dircella larvae could exit) that were collected from leatherwood plants in Gogebic County, Michigan, during the period 17 July to 25 August 1997. Sampling No. of Live larvae Percent mines date mines per mine with holes 17 July 37 3.5 ± 0.4 a* 11 ± 5 c* 29 July 56 1.9 ± 0.2 b 21 ± 6 c 13 August 47 0.5 ± 0.1 c 68 ± 7 b 25 August 44 O.l±O.lc 100 ± 0 a *Means followed with the same letter (within columns) are not significantly different at the P :<;; 0.05 level (Tukey's Studentized Range Test). then decreased slightly as the season progressed (Fig. 3). This decrease in leaf number was primarily due to the abscission of leaf-like stipules, which were first counted as leaves during the early sampling dates. Comparing the heat sums during moth activity, i.e., number of moths col­ lected on sticky panels C!'able 1), with D. palustris phenology (Fig. 3), it ap­ that most moths were captured during the period of active D. palustris and shoot elongation. This relationship is not surprising since L. dircella is host specific to D. palustris. Emerging early during D. palustris leaf flush would allow L. dircella adults to oviposit when leaf tissues are likely soft and most nutritious (Mattson and Scriber 1987). Early season oviposition would also allow the resulting larvae to develop during the entire growing season. This synchronization is probably most important in the northernmost range ofL. dircella where development time may be limiting. Leaf mines were first obvious in 1997 when we visited the study site on 30 June. Each mine contained 1 to 10 legless larvae that formed upper sur­ face blotch mines. The mean (:!: SE) number of live larvae per mine was 3.5 (:!: 0.4, N = 37 mines) on 17 July 1997 and then decreased through the re­ mainder of the season (Table 2). Larvae began exiting mines in July, but most exited during August (Table 2). In general, larvae fed through mid­ August 1997, at which time they developed thoracic legs and exited the mines to pupate. In the laboratory, after larvae exited the mines, they moved to the corners ofthe plastic bags where they formed a translucent, silk cham­ ber and then pupated. In nature, larvae probably pupate in the soil or leaf litter. Leucanthiza dircella likely overwinters in the pupal stage, given that a cold period (4°C for 120 days) was required before adults would emerge in the laboratory. Adults emerged 7-10 days after pupae were removed from cold storage and held at room temperature (ca. 20-24°C). Leucanthiza dircella had only one generation per year in the study area in northern Michigan. Interestingly, L. dircella was reported as having two generations per year in Ohio (Braun 1914, Forbes 1920). A minimum of two generations per year likely occurs in southern Michigan, given that new L. dircella larval feeding mines were observed during both May and September on leatherwood plants in Ingham County, Michigan in 1999 (T. R. Petrice, pers. obs.). Furthermore, adults emerged in the laboratory on 14 June 1999 from larvae that were collected in Ingham County 21 May 1999 and were not subjected to a cold period. Parasitoids. In the laboratory, seven species of hymenopteran para­ sitoids emerged from L. dircella larvae in 1997. They consisted of the bra­ 2000 THE GREAT LAKES ENTOMOLOGIST 195 conid Pholetesor sp. (8 specimens); and the eulophids (Chrysocharis polita Howard (5), Closterocerus trifasciatus Westwood (6), Pnigalio maculipes Crawford (4), P minio (Walker) (2), Pnigalio sp. (3), and Sympiesis sp. (1). Only one of these species, P maculipes, was previously reported as attacking L. dircella (Krombein et al. 1979). The genera Pnigalio and Sympiesis are ec­ toparasitic, which is a common method of parasitism on leafminers since the leaf mine itself serves to protect the parasitoid larvae as well as the host in­ sect. A common behavior often associated with leafminer parasitoids is for the adult female wasps to feed on their insect hosts through the leaf tissues (Askew and Shaw 1979). This provides the adult wasp with protein, which aids in egg production. Young leafminer larvae are often the targets of this behavior, and their flattened cadavers are often found in leaf mines. Indeed, we noticed several dead larvae in mines that were in this condition in 1997. Leaf mining damage. A total of 879 leaves was collected in late August 1997 from 110 leatherwood plants and of these, 31% contained L. dircella leaf mines. Of the 275 leaves containing mines, 259 leaves contained only 1 mine (94%), 14 leaves (5%) contained 2 mines, and 2 leaves (1%) contained 3 mines. The mean leaf size was 17.8 cm2 (range = 0.9 to 66.6 cm2, N = 879 leaves) and the mean mined surface area was 5.9 cm2 (range 0.2 to 24.0 cm2, N 293 mines). Although 31% of the leaves contained mines, only 11% of the total leaf surface area for all 879 leaves was mined by L. dircella in 1997. Another interesting trend was the size of the leaves attacked by L. dir­ cella. After dividing the leaves into six size classes, leaves that were 21 cm2 in surface area or greater had the highest levels ofleaf mining (F = 70.31; df = 5, 873; P < 0.0001) (Fig. 4). Leaves that were 10 cm2 in size and smaller were seldom attacked (Fig. 4). The smallest mined leaf measured 7.5 cm2• When considering only leaves 11 cm2 and larger, more than 54% of the leaves contained mines. One explanation for this difference is that smaller leaves may not have flushed at the time when adult L. dircella were ovipositing in early summer. However this is unlikely since leatherwood has determinate growth and the number of leaves did not increase after initial leaf expansion in early spring of 1999 (Fig. 3). A more likely explanation is that female moths preferentially oviposited in larger leaves. The L dircella population declined in the study area in 1999. Only 8% of the leatherwood leaves contained mines in 1999 compared with 31% in 1997. The number of adult moths captured on sticky panels was also much lower in 1999 than in 1997 or 1998 (Table 1). Parasitism could have contributed to this decline, given the rich parasitoid fauna reared from L. dircella larvae in 1997. Braun (1914) noted high parasitism rates of L. dircella larvae. Freezing temperatures during 1998-1999 winter may have also con­ tributed to the decline of L. dircella in 1999. According to NOAA weather records, in the first half of December 1998, minimum temperatures dipped well below O°C for several days, yet there was no snow cover on the ground. When the study site was visited on 15 December 1999, snow was absent from the ground and the first several cm of soil were frozen (W. J. Mattson, pers. obs.). Leucanthiza dircella pupal mortality may have occurred as a result of freezing and cold temperatures, especially since it occurred in early winter when pupae may have not yet acclimated to their full cold hardiness. In both 1996 and 1997, NOAA weather records indicated that snow covered the ground throughout December, thus insulating the soil and leaf litter. Other damage and insects. In addition to leaf mining damage, 8% of the 879 leaves collected on 25 August 1997 possessed rust spots of an uniden­ tified fungus. Rust spots on leatherwood leaves were not apparent until 30 June 1997, at the same time leaf mines were first noted. A significantly 196 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

80

a ab ab ab 60 ­ "0 CD C 's b -­ m > CG 40 - ..!! ...c CD ~ CD Il. 20 ­

c o rI ~ , ., ..;>... ..r., .,. IS'... "0 ,.~o ,.Vo ,.Yo "\so ,. x

2 Leaf surface area (cm ) Figure 4. Mean percent (+1 SE) of leaves in six size classes that were col­ lected on 25 August 1997 from 110 leatherwood plants and had evidence of leaf mining by Leucanthiza dircella. Means with the same letter are not sig­ nificantly different at P::;; 0.05 (Tukey's Studentized .Range Test). higher percentage of leaves in the 51 cm2 and larger leaf size class possessed rust spots (27%), than did leaves in the smaller leaf size classes (F = 6.10; df 873, P < 0.0001). trunk and limbs of several leatherwood plants in the study area were infested with unidentified scale insects (Homoptera). Of the branch samples collected from the 110 plants on 25 August 1997, 3% had evidence of scale insects. Several other insects were observed on leatherwood foliage in addition to Leucanthiza dircella adults, although actual feeding was not witnessed. These included one native beetle (Coleoptera), Glyptina brunnea Horn (Chrysomelidae); and two exotic beetles, Phyllobius oblongus (L.) (Curculion­ idae) and Polydrusus sericeus Schaller (Curculionidae). Phyllobius oblongus and P. sericeus were also captured frequently on sticky panels during 1997-1999. These two weevils are generalist herbivores that are native to 2000 THE GREAT LAKES ENTOMOLOGIST 197

Europe (Mattson et al. 1994). All three of these beetles inhabit the soil as lar­ vae, feeding on plant roots. At the end of the 1999 growing season, no leatherwood mortality was evi­ dent as a result ofL. dircella. However, we did not measure possible growth loss due to several consecutive years of heavy leaf mining damage. Stem analysis of leatherwood plants in the study area revealed that annual stem elongation varied from 40-180 mm (Cynthia V. Jones, Southern Illinois Uni­ versity at Edwardsville, per. obs.). Some of this variation in annual stern elongation may be attributable to leaf mining damage by L. dircella. If leatherwood becomes a popular ornamental shrub, then control of L. dircella may need to be considered when damage becomes significant.

ACKNOWLEDGMENTS We thank Robert Feldpausch, Sal Hansen, Amanda Lang, Jeremy McCal­ lion, and Therese Poland for field and lab assistance; Robert Kriegel, Mogens Nielsen, Ronald Priest and John Wilterding for Le idoptera rearing and identification assistance; Michael Schauff and Paul for Hymenoptera identifications; Robert Anderson for Coleoptera identifications; and Therese Poland, Ronald Priest, Kurt Schulz and John Zasada for their comments on an earlier draft ofthis paper.

LITERATURE CITED Askew, R. R. and M. R. Shaw. 1979. Mortality factors affecting the leaf-mining stages of Phyllonorycter (Lepidoptera: Gracillariidae) on oak and birch. 2. Biology of the parasite species. Zool. J. Linn. Soc. 67: 51-64. Baskerville, G. L. and P. Emin. 1969. Rapid estimation of heat accumulation from maximum and minimum temperatures. Ecology 50: 514-517. Braun, A. F. 1914. Notes on North American Tineina, with descriptions of new species eLepid.). EntomoL News 25: 113-117. Britton N. L. and H. A. Brown. 1970. An illustrated flora of the northern United States and Canada, voL 2. Dover, New York. Cleland, D. T., J. B. Hart, G. E. Host, K. S. Pregitzer and C. W. Ramm. 1993. Field guide: Ecological classification and inventory system of the Huron-Manistee Na­ tional Forests. USDA Forest Service, Huron-Manistee National Forests, Cadillac, MI. Curtis, J. J. 1959. The vegetation of Wisconsin: An ordination of plant communities. Univ. Wisconsin Press, Madison. J. G. 1949. Leatherwood for early spring bloom. J. New York Bot. Garden 50:

Fernald M. L. 1950. Gray's manual ofbotany, 8th ed. American Book Co., New York. Fielder, M. 1975. Plant medicine and folklore. Winchester Press, New York. Forbes, W. T. 1920. The Lepidoptera of New York and neighboring states. Cornell Univ. Press, Ithaca, NY. Fuller, T. C. and E. McClintock. 1986. Poisonous plants of California. Univ. Calif. Berkeley. Kotar, J.A. Kovach, and C.T. Locey. 1988. Field to the forest habitat types of northern Wisconsin. Department of Forestry, University of Wisconsin, Madison, WI. Krombein, K V, P. D. Hurd, Jr., D. R. Smith and B. D. Burks (eds.). 1979. Catalog of Hymenoptera in America North of Mexico, voL L Smithsonian Inst. Press, Washing­ ton, D.C. 198 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 & 4

Lecours, J. E. W. 1924. Les bois de plomp Dirca palustris. BulL Sci. PharmacoL 31: 112-116. Mattson, W. J. and J. M. Scriber. 1987. Nutritional ecology of insect folivores of woody plants: Nitrogen, water, fiber, and mineral considerations, pp. 105-146. In: F. Slan­ sky and J. Rodriguez (eds.), The nutritional ecology of insects, mites, and spiders. John Wiley and Sons, New York. Mattson, W. ,J., P. Niemela, L Millers and Y. Inguanzo. 1994. Immigrant phytophagous insects on woody plants in the United States and Canada: An annotated list. USDA For. Ser., North Central For. Exp. Sta., Gen. Tech. Rep. NC-GTR-169. Muenscher, W. C. 1964. Poisonous plants of the United States. Macmillan Co., New York. Neilsen, M. C. 1998. Preliminary list of Michigan moths: The Microlepidoptera. Newsletter Michigan Entomo!. Soc. 43(4): 1, 4-14. Nevling, L. I., ,Jr. 1962. The Thymelaeaceae in the southeastern United States. J. Arnold Arboretum 43: 428-434. Radford A. E., H. E. Ahles and C. R. Bell. 1979. Manual of the vascular flora of the Carolinas. Univ. North Carolina Press, Chapel HilL Ramsewak, R. S., 1>1. G. Nair, D. L. Dewitt, W. G. Mattson and J. C. Zasada. 1999. Phe­ nolic glycosides from Dirca palustris. J. Nat. Prod. 62: 1558-156l. SAS Institute. 1990. SAS system for personal computers, release 6.12. SAS Institute Inc., Cary, NC. Tredici, P. D. 1983. Propagating leatherwood: A lesson in humility. Arnoldia 44: 20-23. Vogelmann, H. 1953. A comparison of Dirca palustris and Dirca occidentalis (Thymelaeaceae). ASA Gray BulL 2: 77-83. Whitford, A. C. 1943. Fiber plants of the North American aborigines. J. New York Bot. Garden 44: 25-34. Zasada, J. C., W. J. Mattson, E. A. Nauertz, K. Schulz, C. V. Jones, D. S. Buckley and M. G. Nair. 1999. Dirca palustris in northern hardwood forests: Fruiting, growth, and prospects for special forest products, p. 229. In: 2nd North American Forest Ecol­ ogy Workshop, 27-30 June, Orono, ME. r

2000 THE GREAT lAKES ENTOMOLOGIST 199

NEW DISTRIBUTION RECORDS OF GROUND BEETLES FROM THE NORTH CEt'-ITRAL Ut'-IITED STATES (COLEOPTERA: CARABIDAE)

Foster Forbes Purrington 1, Daniel K. Young2, Kirk 1. larsen3 and Jana Chin-Ting lee4

ABSTRACT We report 39 ground beetles new to five states in the upper midwestern United States. These species records include 19 new to Illinois (all but one from Lake County), 11 from Iowa, three from South Dakota, eight from Wis­ consin, and two from Michigan. (Three species are new to more than one state). Enigmatically disjunct collections include the myrmecophile, Helluo­ morphoides nigripennis from western Illinois, known previously only from the Atlantic and Gulf coastal plain and piedmont, and Chlaenius amoenus, reported only from southeastern states and now from northeast Iowa.

From habitats that include agricultural zones in south central Michigan and Wisconsin, shortgrass prairie in western South Dakota, tallgrass prairies in Iowa and Wisconsin, and forested wetlands along the Lake Michi­ gan shoreline in northern Illinois we report 39 ground beetle species (Coleoptera: Carabidae) that represent 43 new state distribution records (Bousquet and Larochelle 1993, Purrington and Larsen 1997, Purrington and Maxwell 1998). Many of these records are of species that have been previously reported from one or more adjoining states and Canadian provinces. Other records are of synanthropic European introductions (Bousquet 1992) evidently expand­ ing their ranges, or of native species possibly tracking newly favorable envi­ ronments, e.g. changing climates or agricultural landscapes. A few records represent enigmatically disjunct collections of uncommon species previously known only from sites distant from the western Great Lakes region.

MATERIALS AND METHODS All specimens were obtained with standard pitfall or blacklight trapping techniques, or were hand-collected. Vouchers are kept at The Ohio State Uni­ versity-Columbus (FFP), Michigan State University-East Lansing (,JC-TL), Luther College-Decorah, Iowa (KJL), and the University of Wisconsin-

IDepartment of Entomology, The Ohio State University, 1735 Neil Avenue, Colum­ bus, OH 43210. 2Department of Entomology, University of Wisconsin, 237 Russell Laboratories, Madison, WI 53706. 3Department ofBiology, Luther College, Decorah, IA 52101-1045. 4Department of Entomology, Michigan State University, East Lansing, MI 48824­ 1115. Ir

200 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

Madison [Department of Entomology (DKY) and Department of Agronomy (Jon Simonsen: JS)]. The disposition of vouchers is noted by custodial initials in the following species account. Species introduced from Europe are noted.

RESULTS State records for Michigan. Bembidion obtusum Audinet-Serville, In­ gham Co., East Lansing, Michigan State University, Entomology Farm, 4/22/1998 and 5/7/1998, 2 specimens; 3131/1999 and 4/5/1999, 3 specimens, *introduced*, pitfall, [FFP]. Harpalus puncticeps (Stephens), Ingham Co., East Lansing: Michigan State University, Entomology Farm, September 1998; March, April and June 1999, numerous specimens, *introduced*, pitfall, [FFP, JC-TL]. State records for Illinois. Acupalpus pumilus Lindroth, Lake Co., Libertyville, Grainger Woods Forest Preserve and Elm Road Forest Preserve; Winthrop Harbor, Spring Bluff Forest July 1999, numerous speci­ mens,blacklight, [FFP]. Agonum fidele Casey, Lake Co., Libertyville, Grainger Woods Forest Preserve and Elm Road Forest Winthrop Harbor, Spring Bluff For­ est Preserve, June-August 1999, numerous specimens, pitfall and blacklight, [FFP], Agonum harrisii LeConte, Lake Co., Libertyville, Grainger Woods For­ est Preserve; Winthrop Harbor, Spring Bluff Forest Preserve, July 1999, 16 specimens, blacklight, [FFPj. Amara apricaria (Paykull), Lake Co., Winthrop Harbor, Spring Bluff Forest Preserve, 7/22/1999, 1 specimen, *introduced*, [FFP]. Amara quenseli (Schonherr), Lake Co., Winthrop Harbor, Spring Bluff Forest Preserve, 9/29/1999, 4 specimens, pitfall, [FFP], Badister grandiceps Casey, Lake Co., Libertyville, Grainger Woods For­ est Preserve; Winthrop Harbor, Spring Bluff Forest Preserve, July 1999, nu­ merous specimens, blacklight, [FFP). Badister transversus Casey, Lake Co., Libertyville, Grainger Woods For­ est Preserve and Elm Road Forest Winthrop Harbor, S Bluff Forest Preserve, June-July 1999, numerous specimens, blacklight, ], Bembidion pseudocautum Lindroth, Lake Co., Winthrop Harbor, Spring Bluff Forest Preserve, 8/3/1999,1 specimen, blacklight, [FFP). Bradycellus congener (LeConte), Lake Co., Libertyville, Grainger Woods Forest Preserve, 7/8/1999, 1 specimen; Winthrop Harbor, Spring Bluff Forest Preserve, July 1999, 2 specimens, blacklight, [FFP]. Dyschirius integer LeConte, Lake Co., Winthrop Harbor, Spring Bluff Forest Preserve, July 1999,15 specimens, blacklight, [FFP]. Elaphropus tripunctatus (Say), Lake Co., Libertyville, Grainger Woods Forest Preserve and Elm Road Forest Preserve, July 1999, 10 specimens, blacklight, [FFP]. Harpalus puncticeps (Stephens), Lake Co., Libertyville, Grainger Woods Forest Preserve and Elm Road Forest Preserve; Winthrop Harbor, Spring Bluff Forest Preserve, July-August 1999, numerous specimens, *intro­ duced*, blacklight, [FFP]. Helluomorphoides nigripennis (Dejean), Mason Co., Sand Prairie-Scrub Oak State Natural Area, 7/16/1997, 1 specimen, pitfall, [FFP], Paratachys oblitus (Casey), Lake Co., Winthrop Harbor, Spring Bluff Forest Preserve, July 1999, 8 specimens, blacklight, [FFP]. Platynus opaculus LeConte, Lake Co., Libertyville, Grainger Woods For­ est Preserve, 5/28/1999, 3 specimens, blacklight, [FFP]. -

2000 THE GREAT LAKES ENTOMOLOGIST 201

Plochionus timidus Haldeman, Lake Co., Libertyville, Grainger Woods Forest Preserve, 7/30/1999, 1 specimen, blacklight, [FFP]. Pterostichus melanarius (Illiger), Lake Co., Libertyville, Grainger Woods Forest Preserve; Winthrop Harbor, Spring Bluff Forest Preserve, 7/27/­ 811811999, numerous specimens *introduced*, pitfall, [FFP]. Pterostichus novus Straneo, Lake Co., Winthrop Harbor, Spring Bluff Forest Preserve, 8/30/1999 and 9/29/1999, 9 specimens, pitfall, [FFP]. Pterostichus tenuis (Casey), Lake Co., Winthrop Harbor, Spring Bluff Forest Preserve, 7/22/1999,1 specimen, blacklight, [FFP]. State records for Wisconsin. Agonum fidele Casey, Sauk Co., Hem­ lock Draw Preserve, The Nature Conservancy, July 1998, several specimens, pitfall, [FFP]. Amara lunicollis Schillldte, Columbia, Dane, Green, Richland, and Vernon Co., June-July 1997, 1998, numerous specimens, pitfall, [FFP, JSj. Badister {lavipes laticeps Blatchley, Green Co., 6/30/1998, 1 specimen, blacklight, [JS]. Bembidion praticola Lindroth, Juneau Co., Necedah National Wildlife Refuge, 6/11/1996, 2 specimens, pitfall, [DKY]. Chlaenius pusillus Say, Columbia Co., Arlington Research Station, July-August 1996, 5 specimens; Walworth Co., Lakeland Agricultural Sta­ tion, 8/2611996, 1 specimen, [DKY, JS]. Harpalus katiae Battom, , Sauk Co., Spring Green Preserve, 8/24/1995, several specimens, [DKY, FFP]. Harpalus puncticeps (Stephens), 1 specimen, Walworth Co., Lakeland Agricultural Station, *introduced*, 9/1/1995, [DKY]. Selenophorus gagatinus Dejean, Juneau Co., Necedah National Wildlife Refuge, 5/17/1996, 1 specimen, [DKY]. State records for Iowa. Apenes lucidulus (Dejean), Clayton Co., Men­ don Township, Effigy Mounds National Monument, South Unit Prairie, 6/27/1996,1 specimen, [KJL]. Badister neopulchellus Lindroth, Winneshiek Co., Decorah, Anderson Prairie (Luther College), 6/16/1998 and 6/26/1998, 2 specimens, blacklight, [KJLl Bembidion obscurellum (Motschulsky), Floyd Co., 6/27/1978,1 specimen blacklight, [KJL]. Bembidion praticola Lindroth, Howard Co., Chester Township, Hayden Prairie, 6/1411994, 1 specimen; 9/8/1995, 1 specimen, 7/22/1996, 2 specimens, [KJL, FFP]. Bradycellus semipubescens Lindroth, Winneshiek Co., Jackson Town­ ship, Chipera Prairie, 7/1/1996, 1 specimen, [KJL]. Chlaenius amoenus Dejean, Clayton Co., Mendon Township, Effigy Mounds National Monument, South Unit Prairie, 6/27/1996, 1 specimen, [KJL]. Harpalus affinis (Schrank), Winneshiek Co., Jackson Township, 7/22/1996, 1 specimen, *introduced*, [KJL]. Pterostichus trinarius (Casey), Allamakee Co., Fairview Township, 6/19/1995, 2 specimens, (KJL). Scaphinotus bilobus (Say), Winneshiek Co., Decorah, Hickory Ridge Woods, 9/7/1998, 1 specimen; Canoe Township, Malanaphy Springs, 10/1/1999, 1 specimen, [KJL]. Trichotichnus autumnalis (Say), Clayton Co., Mendon Township, Effigy Mounds National Monument, South Unit, 6/27/1996, 1 specimen; Win­ neshiek Co., Canoe Township, Malanaphy Springs, 9/22/1999, 1 specimen, [KJLJ. Trichotichnus vulpeculus (Say), Jasper Co., Walnut Creek National 202 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

Wildlife Refuge, 10/11/1997, 2 specimens; Winneshiek Co., Decorah, 9/5/1995, 1 specimen, [KJL]. State records for South Dakota. Bembidion mutatum Gemminger and Harold, Pennington Co., Wheaton College Science Station (Hisega), 7/10/1994,1 specimen, [KJL]. Carabus nemoralis Miiller, Pennington Co., Wheaton College Science Station (Hisega), 6/12/1996, 1 specimen, *introduced*, [KJL]. Carabus taedatus agassii LeConte, Pennington Co., Wheaton College Science Station (Hisega), 5/23/1995, 1 specimen, [KJL].

DISCUSSION Harpalus puncticeps was inadvertently introduced from Europe into coastal New York (Long Island), probably in the mid-1900's (Dietrich 1957). Davidson (1975) reported its movement 100 km up the Hudson River to Poughkeepsie (Duchess County) and from northern Vermont (Washington County). By 1989 it had been collected in Nova Scotia, Quebec, Ontario, and Ohio (Larochelle and Lariviere 1989). Here we document its continuing west­ ward range expansion with the first records from Michigan, Illinois and Wis­ consin. This species favors ruderal and old field habitats developed on grav­ elly soil (Lindroth 1968). Little is known of Platynus opaculus ecology although it evidently is ex­ tremely hygrophilous. Davidson and Bell (1977) reported many captures in wetland sites of several Vermont counties. They found adults under loose bark of dead trees, stumps and logs protruding from spring floodwaters. Bee­ tles they disturbed often descended quickly below the waterline to depths of 25 cm. where they remained motionless for several minutes before surfacing. It was also found in an abandoned beaver lodge in Quebec (Lindroth 1969). We found the recent European immigrant, Bembidion obtusum, in an agricultural setting in south central Michigan, the westernmost record for this recently introduced (Lindroth 1963) ground beetle. Our records of Harpalus katiae are the most northern for this species, the subject of a recent taxonomic review by Will (1997), who suggested it is more stenotopic than its closely related congener, H. caliginosus (F.). He noted it has moved northward along major river systems, perhaps exploiting sandy alluvial habitats. From west central Illinois we report by far the most disjunct collection of the rare putative myrmecophile, Helluomorphoides nigripennis, known pre­ viously from along the Atlantic coast from Massachusetts to Texas (Ball 1956). Several congeners in southwestern states accompany doryline army ants (Hymenoptera: Formicidae: Dorylinae), feeding on immatures (Wilson 1971), but apparently nothing is known of H. nigripennis biology (Davidson 1995). Two army ant species are recorded from Illinois: Neivamyrmex ni­ grescens (Cresson) from Adams, Pulaski and Union Counties, and N. caroli­ nensis (Emory) from Pope County; a third [N. opacithorax (Emory)] probably occurs in the state (DuBois 1988). Another very anomalous distribution record is that of a male Chlaenius amoenus from northeastern Iowa. This species, [in the emarginatus group of Bousquet and Larochelle (1993) with produced mandibles], has a less than certain taxonomy. According to Robert L. Davidson (pers. comm.), amoenus Dejean may correctly apply to another emarginatus group species that more closely resembles C. pusillus Say. This implies that the Chlaenius we found may be undescribed (the Dejean type needs to be examined). Until now it has 2000 THE GREAT LAKES ENTOMOLOGIST 203

been reported only from Mississippi, Alabama and up the Atlantic coast from Florida to Virginia (Bousquet and Larochelle 1993). We hope this report will encourage field study of ground beetle assem­ blages at more sites in the upper Midwest, especially in ecologically fragile areas such as small wetlands where building development pressures degrade habitats and erode biodiversity at high rates. For us to uncover 18 new state records for Illinois in two wetland enclaves close to metropolitan Chicago, thereby increasing the known state Carabidae fauna by 4%, should be a strong incentive for further such surveys. Some ground beetles found there (of 139 species) are rarely encountered anywhere, and may represent vulner­ able relict populations (Purrington and Davidson 2000). Additionally, we note that since 1997 the known ground beetle fauna of Iowa has been expanded more than 7.3% by adding 24 new records, all from counties in the northeastern corner of the state (Purrington and Larsen 1997). In Wisconsin, similar efforts have added a total 14 new records, in­ creasing the reported fauna by 3.7% (Purrington and Maxwell 1998).

ACKNOWLEDGMENTS We greatly appreciate the counsel and generous taxonomic assistance of Robert L. Davidson, Carnegie Museum of Natural History, Pittsburgh. We thank Jon Simonsen, Department of Agronomy, University of Wisconsin­ Madison and Floyd Catchpole of Hey and Associates, Inc., Libertyville, Illi­ nois for donating ground beetle specimens. For their excellent checklist that colligated authoritative North American ground beetle records and helps to stabilize the nomenclature we express to Yves Bousquet and Andre Larochelle (Agriculture Canada, Ottawa) our sincere thanks. Cathy Drake, Dave Horn, and Adrienne Smith kindly commented on an early draft of the manuscript.

LITERATURE CITED Ball, G. E. 1956. A revision of the North American species of the genus Helluomor­ pho':des Ball, 1951 (Coleoptera, Carabidae, Helluonini). Proc. EntomoL Soc. Wash. 58:67-9l. Bousquet, Y. 1992. Bembidion femoratum Sturm and Amara communis (Panzer) (Coleoptera: Carabidae) new to North America. J. N. Y. Entomol. Soc. 100; 503-509. Bousquet, Y. and A. Larochelle. 1993. Catalogue of the Geadephaga (Coleoptera: Tra­ chypachidae, Rhysodidae, Carabidae including Cicindelini) ofAmerica north of Mex­ ico. Mem. Entomol. Soc. Canada No. 167. Davidson, R. L. 1975. Harpalus (Ophonus) puncticeps Stephens (Coleoptera, Cara­ bidae) in New York and Vermont. Coleopt. Bull. 29: 256. Davidson, R. L. 1995. First Virginia records for ten species of Carabidae (Coleoptera). Banisteria 5:16-19. Davidson, R. L. and R. T. Bell. 1977. Note on the distribution and habitat of Platynus opaculus LeConte (Coleoptera: Carabidae) in Vermont. Cordulia 3: 157-158. Dietrich, H. 1957. Harpalus puncticeps Steph. on long Island, N. Y. Coleop. Bull. 11: 46. DuBois, M. B. 1988. Distribution of army ants (Hymenoptera: Formicidae) in Illinois. Entomol. News 99: 157-160. Larochelle, A. and M.-C. Lariviere. 1988. Further spreading of Harpalus (Ophonus) puncticeps (Stephens) in northeastern North America (Coleoptera: Carabidael. Coleopt. Bull. 43: 75-76. 204 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 & 4

Lindroth, C. H. 1963. The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 3. Opusc. Entomol. Suppl. 24: 201-408. Lindroth, C. H. 1968. The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 5. Opusc. Entomol. Suppl. 33: 649-944. Lindroth, C. H. 1969. The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 6. Opusc. Entomol. Suppl. 34: 945-1192. Purrington, F. F. and R. L. Davidson. 2000. New southerly distribution records for the boreal carrion beetle, Nicrophorus vespilloides (Coleoptera: Silphidae). Entomol. News 111: 355-358. Purrington, F. F. and K. J. Larsen. 1997. Records of thirteen ground beetles (Coleoptera: Carabidae) new to Iowa. J. Iowa Acad. Sci. 104: 50-51. Purrington, F. F. and J. A. Maxwell. 1998. First United States record ofDyschirius sex­ toni (Coleoptera: Carabidae). Entomol. News 109: 189-190. Will, K. W. 1997. Review of the species of the subgenus Megapangus Casey (Coleoptera: Carabidae, Harpalini, Harpalus Latreille). Cole opt. Bull. 51: 43-51. Wilson, E. O. 1971. The insect societies. Belknap, Cambridge. 2000 THE GREAT LAKES ENTOMOLOGIST 205

LlBELLULA FLAVIDA (ODONATA: UBEllUUDAEj, A DRAGONFLY NEW TO OHIO

Tom D. Schultzl

ABSTRACT Libellula flavida, a "videspread but uncommon dragonfly of southeastern and south central North America, is now recorded from Ohio. A breeding pop­ ulation was discovered in an acidic fen on the site of a sandstone quarry in southern Ohio.

Libellula flavida Rambur (Odonata: Libellulidae), the Yellow-sided Skim­ mer, is a medium-sized libellulid dragonfly with pale yellow sclerites on the sides of the adult pterothorax. Little has been written about this species, but records indicate a range extending from the mid-Atlantic coast, across the south, to the south-central plains states (Needham et al. 2000). Although there are a few records for neighboring Kentucky ~Carl Cook, pers. comm.), L. flavida had never been collected in Ohio despite extensive surveys for odonates over the last decade (Glotzhober 1995, 1999, GIotzhober et al. 1995). This paper documents the discovery of a breeding population of L. flavida in Pike County, Ohio. Two adult males were first collected by the author on 24 August 1998 at a sphagnum bog located 0.5 km north of Jackson Lake along the JacksonlPike County line (39°6.48'N, 82°47.70'W). Upon returning to the site on 24 July 1999, several adults of both sexes were collected includ­ ing a few males exhibiting immature coloration. The dragonflies were concentrated around a small sphagnum fen within a large sand mine that ceased operation in 1965 (Kritsky et al. 1999). The fen was situated on a terraced slope on the east side of the quarry. Ground­ water seeped from the base of an excavated wall of Sharon Conglomerate sandstone and flowed slowly through a sphagnum mat approximately 0.5 hectare in area. In addition to Sphagnum cuspidatum Ehrh. ex Hoffm, the following plants were common throughout the fen: hardhack (Spiraea tomen­ tosa), cattail (Typha latifolia), common arrowhead (Sagittaria latifolia), rush (,funcus elfusus and J. acuminatus), bulrush tScirpus purshianus), beakrush (Rhynchospora capiteilata), Joe-Pye-Weed (Eupatorium perfoliatum), and boneset (Eupatorium maculatum). Males of L. flavida flew patrol flights over the fen and alighted fre­ quently on stems in a manner similar to that of L. lydia (Campanella and Wolf 1974). On several occasions, males were observed to seize females in flight and copulations took place while the pairs were perched. Females were observed to oviposit exophytically by hovering over open channels of water and striking the surface intermittently ,vith their abdomens. The pH of the

IDepartment ofBiology, Denison University, Granville, OH 43023. 206 THE GREAT LAKES ENTOMOlOGIST Yol. 33, No.3 & 4 water at six oviposition sites ranged from 4.4 to 7.3 with a mean of 5.8 (S.D. 1.12). Several larvae collected from the fen were later identified as L. flavida (Eric Chapman, pers. comm.), confirming that the population is breeding at this site. Several other odonates were present but uncommon around the wetland. Single adult individuals of the dragonflies Libellula lydia Drury, Libellula luctuosa Burmeister, and Erythemis simplicollis (Say) as well as the dam­ selfly Enallagma aspersum (Hagen) were present in July 1999. During the visit to the site in August 1998, male Somatochlora tenebrosa (Say) were ob­ served patrolling territories amidst the cattails of the fen. A search of the shoreline of nearby Jackson Lake yielded numerous Libellula cyanea Fab. and Libellula incesta Hagen, but no L. flavida.

DISCUSSION Although very little has been published about the ecology of L. flavida, there is consistent anecdotal evidence that this species has an affinity for acid bogs and seepages. This species has been found at acidic bogs in Arkansas (Farris and Harp 1982), South Carolina, Virginia, and northwest Florida. In southern New Jersey, L. flavida is found exclusively at acid bogs with sphagnum, especially old cranberry bogs (Bob Barber, pers. comm.). However, L. flavida has also been collected at calcareous fens and sand bot­ tomed streams. Abandoned sandstone quarries are known to foster the development of pioneer sphagnum bog communities and serve as refugia for native bog plants in Ohio (Andreas and Host 1983). Similarly, the sandpiles of several abandoned mines in southern Ohio have been colonized over the past 50 years by tiger beetle species whose habitat is otherwise scarce in the region (Kritsky et al. 1999). It appears that these manmade habitats have provided the conditions that enabled the establishment ofL. flavida in southern Ohio. Other sandstone quarries are common throughout southern Ohio and should be investigated for the presence of L. flavida. A comparison of wetland habi­ tats where this uncommon species does and does not occur may improve our knowledge ofits ecological niche.

ACKNOWLEDGMENTS Thanks to Bob Glotzhober and Eric Chapman who read an early draft of this manuscript, and to Bob Barber, Bob Behrstock, Carl Cook, Dennis Paul­ son, Steve Roble, and Tim Vogt who shared their collection records.

LITERATURE CITED l~ndreas, B. K. and G. E. Host. 1983. Development of a Sphagnum bog on the floor of a sandstone quarry in northeastern Ohio. Ohio J. Sci. 83; 246-253. Campanella, P. J. and L. L. Wolf. 1974. Temporal leks as a mating system in a temper­ ate zone dragonfly (Odonata: Anisoptera) I: Plathemis lydia (Drury). Behaviour 51: 49-87. Farris, J. L. and G. L. Harp. 1982. Aquatic macroinvertebrates in three acid bogs on Crowley's Ridge in northeast Arkansas. Proc. Ark. Acad. of Sci. 36: 23-27. 2000 THE GREAT LAKES ENTOMOLOGIST 207

Glotzhober, R. C. 1995. The Odonata of Ohio-a preliminary report. Bull. American Odonatology 3: 1-30. Glotzhober, R. C. 1999. Three new state records of Odonata from Ohio, with additional county records. Ohio Biological Survey Notes 2: 25-33. Glotzhober, R. C., R. A. Restifo, T. E. Perry and R. W. Alrutz. 1995. New dragonfly (Odonata) species in Ohio and additions to County records. Ohio J. Sci. 95: 233-239. Kritsky, G., A. Watkins, J. Smith and N. Gallagher. 1999. Mixed of tiger beetles on sand piles ofvarious ages (Coleoptera: Cicindelidae). Cicindela 73-80. Needham, J. G., M. J. Westfall and M. L. May. 2000. Dragonflies of North America. Scientific Publishers, Gainesville, Ohio. 2000 THE GREAT LAKES ENTOMOLOGIST 209

DISTRIBUTION OF FIRST INSTAR GYPSY MOTHS (lEPIDOPTERA: lYMANTRIlDAEI AMONG SAPLINGS OF FOUR TREE SPECIES COMMON IN THE GREAT LAKES REGION

J. l. StoyenofFl,2 and J. A Witter2

ABSTRACT We examined the inter-tree distribution of first instar gypsy moth larvae under natural dispersal conditions in the field in Michigan in 1991. The study focused on saplings of northern red oak (Quercus rubra), white oak (Q. alba), red maple (Acer rubrum), and witch-hazel (Hamamelis virginiana), which are common understory components of forests in the Great Lakes re­ gion. Large-volume trees with foliage that is well-developed early in the spring should provide an excellent surface area for catching newly-hatched gypsy moth larvae, which are randomly dispersed by wind in the spring around the time of leaf flush or shortly thereafter. Comparing across tree species in our study, red maples had the largest crown volumes and well­ flushed leaf material, but very few larvae were found on these trees at both sampling times in mid-May. Despite the lack of larvae on red maple, these trees were liberally covered with gypsy moth silk, indicating that many lar­ vae landed on these plants but left rapidly when they apparently found the trees to be an unacceptable food source. Among the other three tree species, patterns of larval distribution reflected levels of host phenological develop­ ment during both sampling periods, with more insects occurring on tree species that had more advanced leaf development. The highest numbers of larvae were found on northern red oak and witch-hazel. Intermediate num­ bers of insects were present on white oak. Trends in insect distribution also paralleled patterns seen in tree phenology across slope gradients. Defoliation ratings corresponded well to measures of first instar presence for each tree species. Northern red oak and witch-hazel trees experienced more early defo­ liation on average than did red maple and white oak. For those tree species that are acceptable hosts of the gypsy moth, average level of phenological de­ velopment for a given tree, species, or forest stand at the time of larval dis­ persal can be an important predictor of plant or stand susceptibility to gypsy moth establishment and subsequent defoliation.

Gypsy moth (Lymantria dispar CL.» larvae are broadly polyphagous, feeding on 300 different host species (Leonard 1981). In many lepidopterans, choice of the larval food plant is made by the ovipositing adult female. Gypsy moth females, however, are relatively unselective about where they lay their egg masses, readily depositing them even on inedible objects such as rocks,

lCorrespondence address: Dow Gardens, Midland, MI 48640-4292. 2University of Michigan, School of Natural Resources and Environment, Ann Arbor, MI 48109-1115. i

210 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4 stumps, and recreational vehicles (Lance 1983). When larvae hatch in spring, they must find their own food. By spinning silk threads, larvae are able to balloon on the wind in an attempt to find an acceptable host plant. Studies of larval behavior have reported that all healthy larvae will undergo at least one such ballooning dispersal, and larvae will generally balloon again ifthey land on an unsuitable substrate (Lance and Barbosa 1981, Capinera and Barbosa 1976, van der Linde 1971). Typically at the time of gypsy moth first instar larval dispersal in Michigan, tree phenology varies among tree species and individual trees on a given site from tight buds to flattened, expanding leaves (Chilcote 1990, Chilcote et al. 1992). Several factors affect whether dispersing larvae will encounter a given plant. These include characteristics of the plant community, plant location relative to wind patterns and obstacles, plant size and branchiness, and the level of phenological development of foliage (Stanton 1983, Feeny 1976). Ten­ dency to remain on a particular plant once it is encountered may be affected by factors such as plant species, phenological stage of the foliage, density of larvae on the plant, and the level of food reserves available for re-dispersal attempts remaining in the larva's body. Some aspects of dispersal by first instar gypsy moth larvae have been more thoroughly investigated, such as the effects of velocity and turbulence of air currents on larval movement, dis al distances, and settling (Zlotina et al. 1999, Fosberg and Peterson 198 ylor and Reling 198B, Mason and McManus 1981, McManus 1973). The e ects of differing plant characteristics on dispersal activity and distribution of young larvae within a stand is an­ other important aspect which encompasses many factors (Naidoo and Le­ chowicz 1998, Weseloh 1998, Ticehurst and Yendol 1989, Lance and Barbosa 1981, Barbosa 1978a, van der Linde 1971). In particular, understanding has been incomplete relative to ways in which the stage of foliar phenological de­ velopment may affect gypsy moth larval distribution among trees during the ballooning first instar dispersal phase. We undertook this study to examine how first instar gypsy moth larvae are distributed among saplings of various common tree species in the field and how this distribution may be related to patterns ofhost phenological development.

MATERIALS AND METHODS Study site and trees. This study was performed in 1991 at a moder­ ately mesic site dominated by 15 to 20 year old trees of northern red oak (Quercus rubra), white oak (Q. alba), bigtooth aspen (Populus grandiden­ tata), trembling aspen (P. tremuloides), red maple (Acer rubrum), and witch­ hazel (Hamamelis virginiana) in Kalkaska County, Michigan (T25N, RBW, S2B). The vegetation in this area tended to be somewhat open. The site was situated on a slope, so that study trees fell into one of three locations: lower slope (elevation of 353 m), midslope, or hilltop (elevation of 385 m) areas. Natural gypsy moth egg masses were moderately abundant throughout the site. Five minute walks through the site (Eggen and Abrahamson 1983) in each of the three slope locations yielded egg mass counts of 100, 134, and 143 in upper, mid, and lower slope areas, respectively. The plant material selected for study consisted of 19 saplings of northern red oak and 20 saplings each of white oak, red maple, and witch-hazel. All study species were represented in each of the three slope locations, except in the midslope area where northern red oak trees of the appropriate size and exposure (see below) were not available. Neither aspen species present at the -

2000 THE GREAT LAKES ENTOMOLOGIST 211

site was used in this study due to lack of trees of the appropriate size and ex­ posure in the necessary locations. We chose these particular tree species for study because they are com­ mon throughout the Great Lakes region and represent a range of foliage con­ ditions at the time of gypsy moth dispersal from tight buds to flattened, ex­ panding leaves, as well as a range of suitability for gypsy moth larvae from low to high (Chilcote 1990, Chilcote et al. 1992). We used understory-sized trees both to increase ease of sampling and because research by Ticehurst and Yendol (1989) has demonstrated that more than 80% of all early instar gypsy moth larvae may be found in the lower canopy, understory, and forest floor during both day and night. Because tree volume is an important component of bow apparent any given plant is to food-seeking herbivores, we attempted to choose experimen­ tal trees that were naturally of similar sizes (Lance 1983). Some pruning was done during April 1991 on almost all study trees from the ground or from ladders to remove portions of branches that extended beyond the height that could easily be viewed from a 2.5 m ladder (average total tree height after pruning: 2.35 ± 0.26 m for witch-hazel, 2.82 0.29 m for northern red oak, 2.88 ± 0.30 m for white oak, and 3.19 ± 0.34 m for red maple). No sample trees were pruned heavily, however. After all pruning was completed, the di­ ameter of each tree crown was measured in two directions and averaged, and the crown height and total tree height were each estimated to the nearest 0.3 m. The crown height and average crown diameter were used in the equation ofvolume for a cone to estimate the volume for each tree crown. Trees selected for use in the study were in relatively exposed locations so that larvae could readily blow onto them but could not crawl or drop onto the experimental trees from other overhanging trees. In a few cases, portions of nearby large trees were pruned away to avoid blockage of study trees and to help equalize amounts of exposure. Before eggs began hatching in May in the area of the study site, the study trees were inspected for gypsy moth egg masses, which were scraped off when encountered. Also, circles of forest floor at least 2 m in diameter around each sample tree were cleared of all woody debris and leaf litter prior to egg hatch. This helped isolate study trees from sources of potential colonizers in the forest floor litter, since egg masses may be found there as well. These measures were taken so that caterpillars en­ countered on the trees during the course of the experiment could reasonably be expected to have reached the trees through aerial dispersal activity, rather than being present simply because a given tree was in direct contact with or in close proximity to a large number ofegg masses. Sampling procedures. Daily monitoring revealed that peak egg hatch occurred at the study site on 12 May 1991. The weather was very warm and dry (12 May maximum: 27°C, minimum: l6°C), and larvae began to disperse soon after hatch. By the following day, a large portion of the larvae were moving about among potential host plants at all levels ofthe slope. We began the first sample of the study trees during mid-morning of 13 May. Each tree was given an overall rating of leaf development according to the phenological scoring system employed by Chilcote (1990) (Table 1), and a complete count was made of all larvae located anywhere on the tree from the groundline to the tips of the branches. To be counted, larvae had to be in di­ rect contact with the bark or leaves of the tree; larvae suspended from branches by silk strands were not included in the counts. Upper portions of the trees were inspected with the help of 2.5 m ladders, which allowed us to easily view all branches. As larvae were counted, they were removed from the tree and killed. The first sample of all 79 trees was completed on 14 May. A second sample was planned to take place near the end of major gypsy -

212 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No. 3 & 4

Table 1. Phenological scoring system for budbreak and leaf expansion of hardwood trees based on Chilcote (1990). Score Foliar Conditions 1 Winter condition; buds tigbt. 2 Buds swelled but no expansion of scales. 3 Buds and scales expanding. 4 Scale expansion greater than 2 mm. 5 Leaf material showing at tip of bud. 6 Leaves exposed beyond scales but blades not flat. 7 Leaves exposed and blades flattened. 8 Leaves expanding and soft. 9 Leaves expanding and tough. 10 Leaves fully expanded and tough; dark and leathery. moth egg hatch and dispersal at the field site in order to confirm patterns seen at the first sample. Warm temperatures and steadily increasing degree day accumulations during mid-May (Fig. 1) facilitated a rapid hatch. Obser­ vations of egg masses in the area indicated that most eggs were hatched by 16 May. Therefore, trees were sampled a second time on 16-17 May, using the same procedures and following the same pattern through the site. On 18 May, percent defoliation of each study tree was scored as one of five classes based on visual estimation of all foliage on each tree (1 = defoliation < 10%,2 11-25% defoliation, 3 26-50% defoliation, 4 51-75% defoliation, and 5 =76-100% defoliation). Statistical procedures. Statistical analyses were performed with the General Linear Models procedure in SAS (SAS 1996). A two-factor analysis of variance (ANOVA) design was used, which included factors of tree species, slope location, and the interaction ofthese factors. The sampling unit was in­ dividual trees (20 for each species, except 19 for northern red oak). Assumptions were tested using plots ofresiduals versus predicted values, normal probability plots, stem-and-Ieaf plots, and skewness and kurtosis co­ efficients. Where necessary, log transformations were performed on the data before analysis to more closely meet assumptions of normality and homo­ geneity. Scheffe's multiple comparison procedure was used in the evaluation of the data. This procedure was employed for tests of all pair-wise comparisons because its relatively low power reduces the risk of Type I errors occurring in the inferences made. An experiment-wise ex of 0.05 was used.

RESULTS Average tree volume differed significantly among tree species (F = 4.13; df = 3, 68; p = 0.01). Red maple trees had the greatest average volume, al­ though their volume was significantly different only from northern red oak trees with Scheffe's multiple comparison procedure (Table 2). Volume was not significantly affected by slope location (F =0.60; df =2,68; P 0.55) or the in­ teraction of species and slope location (F = 0.79; df = 5,68; p 0.56). Average tree phenology on the first sampling date (13-14 May) was sig­ nificantly affected by species (F 25.07; df 3, 68; P 0.0001) and slope lo­ cation (F 12.26; df 2,68; p = 0.0001). Leaf flush ofwhite oak trees was de­ layed compared to the other tree species (Table 2). White oaks had an r 2000 THE GREAT LAKES ENTOMOLOGIST 213 a. 35

30 ·····Daily max. .... I. --Daily min. • ••• • • • • • - ••I •••••••••• 25 • I· I •• • • I 20 •· ..­• • •I I• • • 0 • • • 0 15 : : : • •• 10 .: ~ • •• 5 - 0 -5 1 6 11 16 21 26 31 May 1991 b. 500 • •• • • - 400 --Base 7.t'C • • • • -- Base 10.O"C •• .-• • 300 •...... •- ,­ .-- 200 r~ •••••••• •••• I····· 1II"" __ fiIIIIl' 100 , '" ~-----.-~ - '" '" o 1 6 11 16 21 26 31

May 1991 Figure 1. Weather data recorded at the National Oceanic and Atmospheric Administration weather station in Kalkaska, Michigan, USA l'V .j::...

Table 2. Plant characteristics and presence of first instar gypsy moth larvae on four tree species at two sampling times during May 1991. Species Variable Red maple Whiteoak Witch-hazel N. red oak -i I N 20 20 20 19 m B Mean crown volume (m ) 1.31 ± 0.14 a 0.96 ± 0.11 ab 0.90 ± 0.09 ab 0.80 ± 0.09 b Gl ;:0 m 1 (13-14 May 1991): ~ phenological ratinga 6.8:!: 0.1 ab 5.0:!: 0.2 c 6.3:l:: 0.2 b 6.8:l:: 0.1 a s;: Mean larvae/tree 51.5 :!: 7.7 b 80.7 ± 18.6 b 424.4 ± 50.7 a 643.2 :!: 60.1 a A 3 C m Mean larvae/m of crown 44.6:!: 7.1 104.6:!: 33.7 c 536.3 :!: 72.9 b 994.8:!: 120.9 a (fl m Sample 2 (16-17 May 1991): Z Mean phenological ratinga 7.4±0.lb 6.6 ± 0.2 c 7.7:l:: 0.1 ab 8.0:!: 0.03 a o Mean larvae/tree 14.5:l:: 2.2 C 207.7:!: 31.4 b 567.1 :!: 45.2 a 587.5 :!: 70.4 a 3: Mean larvae/m3 of crown 13.2 :!: 2.6 c 238.6 :l:: 33.6 b 781.8:!: 94.2 a 901.4 ± 130.5 a o 5 Mean defoliation ratingb 1.0 ± 0.0 c 1.1:!:0.lc 3.1:l:: 0.1 b 3.7:!: 0.2 a Gl Means within each row followed by the same letter do not differ significantly with Scheffe's multiple comparison procedure. ~ al = Winter condition buds; 2 Buds swelled, no scale expansion; 3 Buds and scales expanding; 4 Scale expansion> 2mm; 5 = Leaf material at tip of bud; 6 Leaves exposed beyond scales, blades not flat; 7 Leaves exposed, blades flat; 8 = Leaves expanding and soft; 9 =Leaves expanding and tough; 10 Leaves fully mature. (Modified from Chilcote (1990).) 9: bl defoliation < 10%, 2 =11-25% defoliation, 3 =26-50% defoliation, 4 =51-75% defoliation, and 5 =76-100% defoliation. w w Z 9 w 90 .j::... 2000 THE GREAT LAKES ENTOMOLOGIST 215

average phenology of stage 5 (leaf material just showing at the tip ofthe bud) as compared to averages of stage 6 (leaves exposed but not flattened) to stage 7 (leaves exposed and flattened) for the other species. In terms of slope loca­ tion, leaf flush of lower slope trees was significantly delayed compared with midslope and upper slope trees (lower slope = 5.7 ± 0.2 a; midslope=6.3 ± 0.2 b; upper slope=6.7 ± 0.1 b; different letters indicate mean values significantly different with Scheffe's multiple comparison procedure). The interaction of tree species and slope location was also significant in relation to tree phenol­ ogy (F = 3.45; df 5,68; p = 0.01). Average number of larvae per tree at the first sampling period also was significantly affected by tree species (F = 61.58; df = 3, 68; p = 0.0001). At that time, northern red oak and witch-hazel trees had significantly more lar­ vae present on them on average than did trees of either white oak or red maple (Table 2). Slope location significantly affected number of larvae pre­ sent on trees as well (F = 10.94; df 2,68; p = 0.0001), with an increase in number oflarvae per tree as one progressed up the slope (lower slope = 127.6 ± 31.5 larvae/tree a; midslope 256.0 ± 54.9Iarvae/tree b; upper slope=467.5 ± 59.0 larvae/tree c). The interaction of species and slope location was also significant (F = 2.97; df = 5,68; p 0.02). Examining insect presence on the basis of number of larvae per unit vol­ ume of tree crown revealed results that were very similar to the results for absolute number of larvae per tree. Average number of larvae per unit vol­ ume of tree crown at the first sampling period was significantly affected by tree species (F 67.20; df::: 3, 68; P ::: 0.0001). Northern red oak had the greatest average number of larvae per unit volume followed by witch-hazel, and the figures were significantly larger on each of these species than on ei­ ther white oak or red maple, which were not significantly different from one another (Table 2). Slope location also significantly affected average number of larvae per unit volume of tree crown (F = 6.11; df = 2,68; p 0.004), with increasing values as one progressed up the slope (lower slope = 207.3 ± 71.6 larvae/unit volume a; midslope=328.4 ± 70.5 larvae/unit volume b; upper slope::: 640.6 ± 97.7 larvae/unit volume c). The interaction of species and slope location was significant as well CF = 3.47; df = 5,68; p 0.01). At the second sampling period (16-17 May), average tree phenology was once again significantly affected by tree species (F 21.73; df = 3,68; p 0.0001), with white oak trees still significantly behind the others in terms of average phenological stage (Table 2). Biologically, however, the average phe­ nological condition of the white oak trees indicated that they were beginning to be more suitable as food for the caterpillars at this point. White oak trees now had an average phenology of stage 6 to 7 (leaves fully exposed beyond bud scales with leaf blades not yet flat to blades exposed and flattening), as compared to the average white oak phenology at the first sampling period of a stage 5 (leaf material just showing at the tip of the bud). Slope location also significantly affected average phenology at the second sample date (F 12.75; df::: 2, 68; P 0.0001). Mean phenological stage, averaged across all trees in the study, increased as one progressed up the slope (lower slope = 7.0 ± 0.1 a; midslope = 7.4 ± 0.1 b; upper slope=7.8 ± 0.1 c). The interaction of species and slope location was also significant (F = 2.34; df 5, 68; P = 0.05). Average absolute number of larvae present on the study trees at the sec­ ond sampling period was significantly affected by tree species (F = 128.60; df = 3, 68; P 0.0001). Northern red oak and witch-hazel trees had the greatest numbers of larvae per tree on average, and the numbers on these species were significantly greater than the numbers on white oak (Table 2). Num­ bers of larvae on red maple were significantly lower than numbers of larvae on any other tree species. At this second sampling period, average numbers 216 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4 of larvae per tl'ee were not significantly affected by slope location (F = 2.25; df 2, 68; p ::: 0.11) or an interaction of species and slope location (F = 1.87; df= 5, 68; p ::: 0.11). Results based on number of larvae per unit volume of tree crown at the second sampling period were once again very similar to the results for ab­ solute numbers of larvae per tree. Average number of larvae per unit volume of tree crown was significantly affected by tree species (F =156.92; df::: 3, 68; p 0.0001), with values being greatest on northern red oak and witch-hazel, which were not significantly different from each other (Table 2). These were followed by the significantly lower figure on white oak and finally the aver­ age number of larvae per unit volume of tree crown on red maple, which was significantly lowest of all species. Average number of larvae per unit volume oftree crown was not significantly affected by slope location (F ::: 0.75; df = 2, 68; p ::: 0.48) but was affected by the interaction of species and slope location (F =2.80; df= 5, 68; p 0.02). Average defoliation levels were significantly affected by tree species (F 124.67; df = 3, 68; p = 0.0001) and ranged from <10% to >50% (Table 2). High defoliation levels were due to the large number of gypsy moth caterpillars per tree on certain species and the small size of leaves early in the season. Slope location also significantly affected defoliation levels (F ::: 10.27; df = 2, 68; P ::: 0.0001). Effects of slope location on defoliation followed the general pattern seen for tree phenological development and numbers of larvae pre­ sent, with lower defoliation occurring on the lower slope and increasing defo­ liation as one moved up the slope (lower slope=1.5 ± 0.1 a; midslope=2.0 ± 0.3 b; upper slope = 2.9 ± 0.2 c; where class 1 = defoliation < 10%, class 2 = 11··25% defoliation, class 3 26-50% defoliation, as described above). Tree species and slope location significantly interacted to affect defoliation as well (F =3.18; df=5, 68; P == 0.01).

DISCUSSION Work by others has shown that oaks are highly favored host plants of the gypsy moth, but maples are not very acceptable or suitable hosts, which is perhaps due to the presence of toxic/deterrent chemicals in maple foliage (Barbosa et al. 1990, Martinat and Barbosa 1987, Lance 1983, Barbosa and Greenblatt 1979, Barbosa et al. 1979, Barbosa 1978a, 1978b). However, red maples often tend to be among the earlier flushing tree species in the Great Lakes region and in our study ranked highest in terms of average volume. Large-volume trees that flush early and have relatively large leaves should provide an excellent surface area for catching insects that are randomly dis­ persed by the wind, and one might expect to find high numbers of larvae on such trees. Despite this, the number of larvae present on red maple trees in our study was very low at both sampling times, both in terms of absolute numbers and on the basis of mean larvae per m3 of crown. A qualitative ex­ amination revealed that red maple buds and branch tips were often covered with more insect silk than was true for trees of other species except for white oaks with very delayed phenology, on which the amount of silk appeared sim­ ilar to that seen on red maple. The large amount of silk observed on red maple trees indicates that they were probably encountered by many larvae. However, since we found very few larvae remaining on the trees it is likely that larvae may have re-dispersed in search of different host materiaL Be" cause few larvae were located on this species, it suffered very little defolia­ tion on average during the study period. Research by others has demon­ strated that young gypsy moth larvae do disperse rapidly and with high 2000 THE GREAT LAKES ENTOMOLOGIST 217 frequency from unacceptable substrates (Martinat and Barbosa 1987, Lance 1983, Capinera and Barbosa 1976, van der Linde 1971, Leonard 1967). While there are other factors that also come into play when insects choose among relatively favored host species, match of optimal host phenol­ ogy to insect activity is one important factor. An appropriate level of pheno­ logical development means that adequate leaf material in suitable condition for feeding is available (Stoyenoff et al. 1994a, Raupp et al. 1988). Level of phenological development is also related to other important characteristics of the foliage, such as nutrient and water content, toughness, and levels of de­ fensive compounds (Hunter and Lechowicz 1992, Hough and Pimentel 1978, Feeny 1976). Many workers have found that plant phenology is a major fac­ tor affecting insect host use (Parry et al. 1998, Kolb and Teulon 1992, Craw­ ley and Akhteruzzaman 1988, Larsson and Ohmart 1988, Raupp and Denno 1983, Mitter et al. 1979, Schweitzer 1979, Witter and Waisanen 1978, Holli­ day 1977, Feeny 1970, Greenbank 1956). For those species that are acceptable hosts of the gypsy moth, average level of phenological development seen on a given tree, species, or forest stand at the time of larval dispersal may be one important predictor of plant or stand susceptibility to insect establishment and subsequent defoliation. We found that numbers of insects present on acceptable host species tended to strongly follow patterns seen in plant phenology, with more insects occur­ ring on tree species that had advanced leaf development. For instance, at the first sampling period, northern red oaks had on average leaf material free of the bud scales and flattening out, witch-hazels had leaf material free of the bud scales but on average more folded, and white oaks had much leaf mater­ ial still surrounded by bud scales with only some tightly folded leaf material showing at the tips of the buds. Northern red oaks at this sample had a high number of larvae present on them on average, witch-hazels had a slightly lower number, and white oaks had a much lower number. This was true both in terms of absolute numbers of larvae per tree and number of larvae per unit ofcrown volume. In the case of the significant increase in insect numbers on white oak be­ tween the first and second sampling times, the phenological changes that took place over this time period greatly altered the usefulness of these plants to young gypsy moth caterpillars. However, while this phenological change may be very important, it is inadequate to elevate numbers of insects found on white oak to the level found on the other acceptable host species. At stage 6, white oak leaves are stHl covered by many dense trichomes, and the pres­ ence oftrichomes has been shown to inhibit feeding in some cases of plant-in­ sect interactions (Agrawal 1999, Zvereva et al. 1998, Dix et al. 1996, Kanno 1996, Oghiakhe 1995, Wright and Giliomee 1992, Gross and Price 1988, Khan et al. 1986, Hardin 1979, Johnson 1975, Levin 1973). This factor may be coming into play here. By contrast, northern red oak trees at the second sample date were about 1.5 stages more advanced than white oak on average and had expanding, soft, young leaves that had shed any trichome covering. While it is possible that insect numbers were higher on northern red oak be­ cause the more phenologically advanced northern red oak leaves had larger flattened leaf surfaces available for catching larvae than did white oak, we repeatedly observed that the lower-phenology white oak trees were covered with more caterpillar silk than were the northern red oak trees, indicating that large numbers of larvae were also encountering the white oak trees but some were leaving in search of more highly acceptable food sources. Average defoliation levels observed for the various tree species in this study indicate that northern red oak and witch-hazel were very acceptable food sources to the larvae, and they ate large fractions of the leafmaterial on 218 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 & 4 these species. Since white oak leaves were on average less developed and smaller than leaves of northern red oak and witch-hazel by the date defolia­ tion was evaluated, it would have been possible for the white oaks to score higher in terms of defoliation rating even if they were fed on less, due to the fact that white oak trees had less expanded foliage available to begin with (Valentine 1983). Such was not the case, however. Despite larger amounts of foliage present, northern red oaks and witch-hazels scored significantly higher in terms of average defoliation rating than did white oaks in this study. Both white oaks and red maples escaped significant defoliation early on in the season due to various characteristics that led to these plants being less acceptable to young gypsy moth larvae. Foliage of preferred host species has been shown by others to generally retain higher numbers of gypsy moth larvae than foliage of less preferred species (Lance 1983, Lance and Barbosa 1981, Barbosa 1978a). Based on av­ erage number of insects present and average defoliation levels seen in the field in this study, witch-hazel appears to be a very acceptable host species for young gypsy moth larvae, almost as much so as northern red oak. Larvae may use species such as witch-hazel and northern red oak at a time when other species such as white oak or aspens are phenologically behind and are not as acceptable to the insects. The insects can later move on to aspens and white oak as their leaves flush and their acceptability increases (Stoyenoff et al. 1994b, 1994c). Presence of acceptable hosts that are phenologically well timed with caterpillar hatch and early dispersal can therefore increase over­ all defoliation potential of a stand throughout the season and heighten per­ formance of the gypsy moth in an area.

ACKNOWLEDGMENTS This work was supported in part with funds provided to the University of Michigan by the USDA McIntire-Stennis Program. We thank the Michigan DNR, Forest Management Division, for providing research housing. Anony­ mous reviewers provided helpful comments and insight.

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2000 THE GREAT LAKES ENTOMOLOGIST 219

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COMPARISON OF TWO POPULATION SAMPLING METHODS USED IN FiElD LIFE HISTORY STUDIES OF MESOVEUA MULSANTI [HETEROPTERA: GERROMORPHA: MESOVELIIDAEj IN SOUTHERN ILLINOIS

Sleven J. Taylor1 and J. E. McPherson2

ABSTRACT A field life history study of Mesovelia mulsanti was conducted in south­ ern Illinois, the results of which are compared with those from an earlier study also conducted in southern Illinois. The two studies differed in the collecting techniques used (quadrat sampler versus aquatic net). Results of the present study give a clearer picture of the life history of this insect be­ cause the quadrat sampler collected representative samples of nymphs and adults more effectively than the aquatic net and, thus, the quadrat samples more accurately represented the actual chronology of the annual genera­ tions.

The water treader Mesovelia mulsanti White occurs from Newfound­ land, Nova Scotia, New Brunswick (Scudder 1987), New York, and Massa­ chusetts south to Florida, and west to British Columbia and California; it also has been reported from Mexico to Argentina, the West Indies, and the Hawaiian Islands (Smith 1988). It occurs throughout much of Illinois (Tay­ lor 1996). Mesovelia mulsanti is epipleustonic, inhabiting shaded and unshaded areas, especially standing water with duckweed and algae (McPherson 1988, Taylor 1996). It is predaceous, feeding primarily on insects on the water sur­ face and, possibly, on small organisms that come to the surface from below (McPherson 1988). This insect's life history has been studied in southern Illinois (Gal­ breath 1973, 1975, 1976a, 1976b; McPherson 1988) and elsewhere (e.g., Hoffmann 1932; Hungerford 1917, 1920; Lanciani 1987). However, there is some question about the number of generations per year. In southern Illi­ nois, for example, Galbreath (1973, 1975) stated that this species has five and possibly more generations per year, whereas McPherson (1988) found only three generations and a partial fourth, with eggs apparently overwin­ tering. McPherson's (1988) field study was conducted in the La Rue-Pine Hills Ecological Area (now La Rue-Pine Hills Research Natural Area) in Union

lCenter for Biodiversity, Illinois Natural History Survey, 607 East Peabody Drive, Champaign, Illinois 61820-6970. 2Department of Zoology, Southern Illinois University at Carbondale, Carbondale, Illinois 62901-6501. 224 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

County from 1983 to 1986. Specimens were collected with a 0.33 m-wide aquatic net at approximately weekly intervals during the active season (April to November). Data from the four years were combined to gain a better understanding of the annual life cycle. However, there is some question about the reliability of his data because of problems inherent in the use of an aquatic net (i.e., smaller specimens are more likely to be overlooked) and be­ cause the study was part of a larger qualitative study of both the semi­ aquatic and aquatic Heteroptera. For this larger study, sweeps collected ma­ terial below and on the surface of the water simultaneously and included aquatic vegetation, detritus, and insects. There was no attempt to limit sweeps to the surface of the water and, therefore, detecting the earlier in­ stars of M. mulsanti (1st instars averaged less than 1.00 mm in length [un­ published data]) in samples containing much organic debris was difficult. As would be expected, the number of adults collected represented an artificially high percentage of the total number of individuals collected during the study (2,239/5,092; 44%). This paper results from a larger study of the biology of the Gerromorpha in Illinois (Taylor 1996). As a part ofthat project, we studied the life histories and microhabitat distributions of several species (including M. mulsanti) during 1989 and 1990 at President's Pond on the campus of Southern Illinois University at Carbondale, Jackson County, Illinois. President's Pond is roughly triangular, with a surface area of approximately 0.29 hectare (0.72 acre) (see Taylor [1996] and Taylor and McPherson [1998a, 1998b, 1999] for detailed description of pond). The collecting technique for President's Pond was designed s cally to sample all gerromorphan instars, regardless of size. Because erson's (1988) study was conducted only 25 km southwest of President's Pond, we were able to compare our results and collecting techniques with his. There­ fore, the purposes of this paper are to present the results of our study of the annual life cycle of M. mulsanti at President's Pond, compare the efficacy of the collecting technique with that ofMcPherson (1988) for sampling nymphal and adult stages, and examine the influence of these techniques on our abil­ ity to distinguish between generations.

MATERIALS AND METHODS Samples were collected weekly from 18 March to 25 November 1989 and, because of overcollecting in 1989, biweekly from 11 February to 2 December 1990. Sampling was limited to an area along the eastern shore because (1) the cattails along the western shoreline prevented use of the quadrat sam­ pler (see below); (2) the riprap shoreline of the southern border was unnat­ ural and, often, disturbed by fishermen; and (3) the water surface along the eastern shore, which was a mosaic of open water, duckweeds, and emlerlzerlt stems, supported a diverse gerromorphan fauna. Four 60 m transects were made parallel to a relatively uniform section of the eastern margin at 0, 0.5, 1.0, and 1.5 m from the shoreline. Each sample was collected with a floating quadrat sampler (0.25 x 0.25 x 0.05 m) (see Taylor 1996), with four replicates placed randomly along each transect; the resulting 16 quadrat samples were pooled, providing a broad sampling of the habitat. Prior to each sample, the collector (SJT) stood for approxi­ mately 3 minutes to allow the insects to acclimate to the disturbance; then, the sampler was placed on the surface of the water. Epipluestonic arthro­ pods were removed with a fine mesh nylon net (0.42 mm diam. mesh open­ ings), preserved in alcohol, and sorted in the laboratory by instar, and, ad­ -

2000 THE GREAT LAKES ENTOMOLOGIST 225

Figure 1. Percent of individuals in each stage per sample of Mesovelia mul­ santi collected at President's Pond, Southern Illinois University at Carbon­ dale campus, Jackson County, during 1989. Beginning and end points of each shaded area represent sample dates preceding and following collection of specimens, respectively. ditionally for 5th instars and adults, by sex and wing morph (apterous and macropterous). Galbreath's (1975) intermediate morph was not distin­ guished from the apterous morph in our study. Nymphal instars were dis­ tinguished by tergal and genitalic features (see McPherson and Taylor 1993).

RESULTS AND DISCUSSION Mesovelia mulsanti apparently overwinters as eggs. Combining data from 1989 and 1990, first instars were found from late April through mid­ September, second instars from mid-April through late September, third in­ stars from late April through late September, fourth instars from late April 226 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

MAR APRIL MAY JUNE JULY AUG SEPT. OCT NOV. 50

5 ADULT .... a (N=214) t-- 2 5

50)-

2 5 1ST a INSTAR ~- ... 5 (N=536) ......

50)-

2 5 2ND a INSTAR ...-... --_._.

5 4TH a INSTAR ...... - - 5 (N=127) 2

501-

5 5TH a INSTAR ~ r--- r- 5 (N=117) -- 50 '1 Figure 2. Percent in each sample of total individuals of same stage of Mesovelia mulsanti collected at President's Pond, Southern Illinois Univer­ sity at Carbondale campus, Jackson County, during 1989. Beginning and end points of each shaded area represent sample dates preceding and following collection of specimens, respectively. through late September, fifth instars from late April through late September, and adults from early May through early November (Figs. 1-4). The number of generations per year was difficult to determine although apparently there were four or five. Hoffmann (1932), under uncontrolled lab­ oratory conditions, reported nymphal development averaged 20.02 days; whereas Lanciani (1987), under controlled conditions (28oC, 12L:12D pho­ toperiod), reported it averaged 13.38 days. Because these insects are active from April to November in Illinois (McPherson 1988, Taylor 1996), and devel­ opmental time varies with changes in temperature (Galbreath 1975), the short generation time reported by Lanciani (1987) suggests that the active season is of sufficient length to allow four or five generations per year in southern Illinois. McPherson (1988) reported three generations and a partial fourth. Al­ r 2000 THE GREAT LAKES ENTOMOLOGIST 227

(/) 0; :::J ~ '6 .£: '0 1:

Figure 3. Percent of individuals in each stage per sample of Mesovelia muZ­ santi collected at President's Pond, Southern Illinois University at Carbon­ dale campus, Jackson County, during 1990. Beginning and end points of each shaded area represent sample dates preceding and following collection of specimens, respectively.

though it might seem that his study would have been more thorough because it was conducted over four years rather than two, voltinism was determined by combining the data across all years into a single year because the num­ bers of early instars were low. In addition, sample dates varied slightly from year to year, and, therefore, samples from the four years were lumped into weekly samples (JEM, unpublished data). Therefore, it is likely that the peaks corresponding to generations were obscured. As discussed earlier, McPherson (1988) collected aquatic and semiaquatic Heteroptera with an aquatic net, whereas the present study used a quadrat sampler. Because sampling was limited to the epipleuston in the present study, samples were relatively free of organic debris compared to those col­ lected in McPherson's (1988) study. Comparisons of the numbers of nymphal instars and the adults showed that the quadrat sampler collected representa­ 228 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 & 4

Figure 4. Percent in each sample of total individuals of same stage of Mesovelia mulsanti collected at President's Pond, Southern Illinois Univer­ sity at Carbondale campus, Jackson County, 1990. Beginning and end points of each shaded area represent sample dates preceding and following collection of specimens, respectively.

tive samples of all stages more effectively than the aquatic net (Fig. 5). Al­ though the results of the present study were not dramatically different from those of McPherson (1988), the quadrat samples more accurately represent the actual chronology of the annual generations and provide higher confi­ dence in our conclusions.

ACKNOWLEDGMENTS We thank R. E. DeWalt and M. J. Wetzel, Illinois Natural History Survey, for their critical reviews. R A. Brandon, J. A. Beatty, B. M. Burr, Department of Zoology; and D. Ugent, Department of Plant Biology, Southern Illinois University at Carbondale, reviewed an earlier version ofthis manuscript. 2000 THE GREAT LAKES ENTOMOLOGIST 229

A B Adult 5th Instar 4th Instar 3rd Instar 2nd Instar 1st Instar

Figure 5. Life stage distribution (percent of individuals) for MesoveZia muZ­ santi. A-specimens collected with quadrate sampler in 1989 at President's Pond, Southern Illinois University at Carbondale campus, Jackson County. B-specimens collected with aquatic net from 1983 to 1986 at La Rue-Pine Hills Research Natural Area, Union County (McPherson 1988).

LITERATURE CITED Galbreath, J. E. 1973. Diapause in Mesouelia mulsanti (Hemiptera: Mesoveliidae). J. Kansas Entomol. Soc. 46: 224-233. Galbreath, J. E. 1975. Thoracic polymorphism in Mesouelia mulsanti (Hemiptera: Mesoveliidae). Univ. Kansas Sci. Bull. 50: 457-482. Galbreath, J. E. 1976a. The effect of the age of the female on diapause in Mesouelia mulsanti (Hemiptera: Mesoveliidae). J. Kansas Entomol. Soc. 49: 27-3l. Galbreath, J. E. 1976b. Reproduction in Mesouelia mulsanti (Hemiptera: Mesoveli­ idae). Trans. Illinois State Acad. Sci. 69: 91-99. Hoffmann, C. H. 1932. The biology of three North American species of Mesouelia (Hemiptera-Mesoveliidae). Can. Entomol. 64: 88-95,113-120,126-134. Hungerford, H. B. 1917. The life-history of Mesouelia mulsanti White. Psyche 24: 73-84. Hungerford, H. B. 1920. The biology and ecology of aquatic and semiaquatic Hemiptera. Univ. Kansas Sci. Bull. 11: 3-328 (1919). Lanciani, C. A. 1987. Rearing immature Mesouelia mulsanti (Hemiptera: Mesoveliidae) on a substratum of duckweed. Fla. Entomol. 70: 286-288. McPherson, J. E. 1988. Life history of Mesouelia mulsanti (Hemiptera: Mesoveliidae) in southern Illinois. Great Lakes Entomol. 21: 19-23. McPherson, J. E. and S. J. Taylor. 1993. Distinguishing nymphal instars of Mesouelia mulsanti (Heteroptera: Mesoveliidae). Great Lakes Entomol. 26: 233-236. Scudder, G. G. E. 1987. Aquatic and semiaquatic Hemiptera of peatlands and marshes in Canada. Mem. Entomol. Soc. Canada 140: 65-98. Smith, C. L. 1988. Family Mesoveliidae Douglas and Scott, 1867, pp. 247-248. In: T. J. Henry and R. C. Froeschner (eds.) Catalog of the Heteroptera, or true bugs, of Canada and the Continental United States. E. J. Brill, New York. Taylor, S. J. 1996. Habitat preferences, species assemblages, and resource partitioning by Gerromorpha (Insecta: Heteroptera) in southern Illinois, with a faunal list and 230 THE GREAT LAKES ENTOMOLOGIST Vol. 33, No.3 &4

keys to species of the state. Ph.D. Dissertation, Department of Zoology, Southern Illi­ nois University at Carbondale. Taylor, S. J. and J. E. McPherson. 1998a. Voltinism in Neogerris hesione (Heteroptera: Gerridae) in southern Illinois. EntomoL News 109: 233-239. Taylor, S. J. and J. E. McPherson. 1998b. Voltinism in Merragata brunnea (Het­ eroptera: Gerromorpha: Hebridae) in southern minois. Fla. Entomo!. 81: 509-515. Taylor, S. J. and J. E. McPherson. 1999. Morphological variation and polyvoltinism of Microvelia pulchella (Heteroptera: Veliidae) in southern Illinois, USA. Acta Soci­ etaris Zoologicae Bohemicae 63: 237-249. -

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