Vol. 30, No.3 Fall 1997 THE GREAT LAKES ENTOMOLOGIST

PUBLISHED BY

THE MICHIGAN ENTOMOLOGICAL SOCIETY THE GREAT LAKES ENTOMOLOGIST

Published by Ihe Michigan Entomological Society

Volume 30 No, 3

ISSN 0090-0222

TABLE OF CONTENTS

Nearctic Ader;s: resurrection of A slad;ana and a revised identity for A semiannula (Lepidoptera: Tortricidae) Michael Sabourin, Ronald J. Priest and William E, Miller, , , , , " """"'" """,71

Notes on Ihe life histories of Acrosternum hilare and Cosmopepla bimaculata {Heteroptero: PentatomidaeJ in soulhern Illinois j, E, McPherson and D, L Tecic, , , , , , , , , , , , , , , , , , , , , , , " ,,',"""" "'" 79

New record of Brachycercus maculalus Berner (Ephemeroptera: Caenidae) from New York and a key to larvae of northeastern species Sleven K, Burian, Margaret A. Novak, Robert W, Bode and Lawrence Abele, , , , , , , , , , , , , 85

Hexagenia bilineata (Ephemeroptera: Ephemeridae) persists at low levels of abundance in thll Lower Fox River, Wisconsin Philip A. Cochran and Andrew p, Kinziger , , ' , , , , , ' , , , , , , ' , , , , , , , , , ' , , , , , , , , , ' 89

Evaluation of Paederus /ifforarius (Coleoptera: StaphylinidaeJ as an egg predator of Chrysoleuchia lopiaria (Lepidoptera: Pyralidae) in Wisconsin cranberry bogs Sandra Haase-Statz , , , , ' , ' , , , , , , , , , , , , , , , , , , , ' , , , , , , , , , , , , , , , , , , , , , , ' , 93

Effects of feeding by two folivorous arthropods on susceptibility of hybrid poplar clones to a foliar pathogen Kier D, Klepzig, Daniel J. Robison, Eugene B, Smalley and Kenneth F. Raffo, , , , ' " , , , , , 99

Urophora affinis and Urophora quadrifasciata (Diptera: Tephritidae) released and monitored by USDA, APHIS, PPG as biological control agents of spotted and diffuse knapweed R, F. Lang, R, D, Richard and R, W Hansen , ' , ' , , , , , , , ' , , , , , , , , , , , , , , , 105

Introduced purple loosestrife as host of native Salurniidae [Lepidoptera) James G, Barbour and Erik Kiviat , , , , , , , , , , , , , ' , , , , , , , , , , ' , , , , , , , , , , "",115

The assassin bug Zelus luridus {Heteroptera: ReduviidaeJ in Michigan's Upper Peninsula Philip A. Cochran, lames R, Hodgson and Adam A. Leisten, , , , , , , , , , , ' , ' , , , ' , , , , , , 123

COVER PHOTOGRAPH Regal fritillary. Speyeria idalia Drury (Lepidoptera: Nymphalidae) on Echinacea pal/ida, Photograph by Ann B, Swengel. THE MICHIGAN ENTOMOLOGICAL SOCIETY

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Copyright © 1997, The Michigan Entomological Society 1997 THE GREAT LAKES ENTOMOLOGIST 71

NEARCTIC ACLERfS: RESURRECTION OF A. STADfANA AND A REVISED IDENTITY FOR A SEMfANNULA (LEPIDOPTERA: TORTRICIDAEl

2 Michael Sabourin 1, Ronald J. Priest f and William E. Miller3

ABSTRACT Type study showed that Acleris stadiana (Barnes & Busck), currently considered a junior synonym ofA. semiannula (Robinson), is in fact a distinct taxon. Although superficially similar, these taxa differ markedly in genital structure. In males ofA. semiannula, the aedeagus is short, broad, and virtu­ ally straight, whereas in those of A. stadiana, it is long, thin, and sharply bent. What was known in literature as A. semiannula proved to be A. stadi­ ana. We redefine both A. semiannula and the resurrected A. stadiana.

Several years ago, abundant caterpillars were discovered feeding on the leaves of mature red eAcer rubrum L.) and silver maple CA. saccharinum L.) growing on a golf course and around residences in Isabella Co., Michigan. Reared adults were identified as Acleris sp. These adults superficially re­ sembled what was known at the time as A. semiannula (Robinson), but their genitalia differed. Our efforts to identify the species led to the findings re­ ported here. Acleris stadiana (Barnes & Busck) has been considered a junior synonym ofA. semiannula since these taxa were synonymized by McDunnough (1934). Their types were not dissected for genitalia study until now, however. We in­ vestigated the pertinent types, comparing them with the Michigan specimens reared from maple, and with what was known in literature as A. semiannula. Abbreviations for collections mentioned here are as follows: AMNH, American Museum of Natural History, New York, New York; ANSP, Academy of Natural Sciences of Philadelphia, Philadelphia, Pennsylvania; CMNH, Carnegie Museum of Natural History, Pittsburg, Pennsylvania; DP, Dennis Profant collection, Nelsonville, Ohio; EME, Essig Museum of Entomology, University of California, Berkeley; ERL, Entomol Research Laboratory collection, S. Burlington, Vermont; GJB, George J. gh collection, Portage, Michigan; GRN, Gordon R. Nielsen collection, Hinesburg, Vermont; INHS, Illinois Natural History Survey, Champaign; JDG, John D. Glaser collection, Baltimore, Maryland; JRH, J. R. Heitzman collection, Independence, Mis­ souri; I\lATH, Bryant Mather collection, Clinton, Mississippi; MSUC, Michi­ gan State University, East Lansing; RJP, Ronald J. Priest collection, East Lansing, Michigan; UMSP, University of Minnesota Entomology Museum, St. Paul; USNM, National Museum of Natural History, Washinl;,'ton, D. C.;

123476 Johnson Rd., Grantsburg, WI 54840. 25464 Jo Pass, East Lansing, MI 48823. 3Department of Entomology, University of Minnesota, St. Paul, MN 55108. ----_...._------_.----­

72 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

UWEM, University of Wisconsin, Madison; WBP, William B. Procter collec­ tion, University ofMassachusetts, Amherst. The letter n denotes the number of specimens on which a statement is based; m stands for male; f for female; and glyc. for glycerine. Collection dates are month/day/year format.

Acleris semiannula (Robinson) (Figs. 1, 3-5) Teras semiannula Robinson (1869: 282, pI. 7, fig.70 ) (Holotype: female, type # 7414, Penn., no date, genit. prep. MS 96115, forewing length 6.8 mm, in ANSP, wings shown here in Fig. 1); Zeller (1875: 212). Teras ferrugana (not Denis & Schiffermiiller 1775); Walsingham (1879: 76), Fernald (1882: 8) (in part). Acleris ferrugana (not Denis & Schiffermtiller 1775); Fernald (1902: 474) (in part), Shaw (1905: 325). Peronea ferrugana (not Denis & Schiffermtiller 1775); Meyrick (1912: 60) (in part), Forbes (1923: 487) (in part), McDunnough (1934: 321) (in part), (1939: 59) (in part), Brower (1983: 49) (in part). Acleris kearfottana (not McDunnough 1934); Obraztsov (1963: 229). Acleris tripunctana (not Hubner 1796-99); Obraztsov (1963: 223) (in part), Opheim (1964: 302-303) (in part), ?Powell (1983: 38), Godfrey et al. (1987: 30). Acleris semiannula; Godfrey et a1. (1987: 30). Acleris n. sp.; Grehan et a1. (1995: 22). Female and Male. Forewing (Fig. 1) span 12.5-16.5 mm (n=56), costal triangle present. Ground color of forewing, including costal margin within costal triangle, cinnamon drab to fuscous with brownish highlights. Costal triangle consistently dusky brown, always darker than ground, at times barely discernable depending on shade of ground. Basal patch sometimes de­ fined by a fine transverse line. Hindwing color similar to that offorewing. Male genitalia (Figs. 3, 4) (n=10). Tegumen hood-shaped with large rounded terminal lobes. Tuba analis with a large ventral projection at mid­ dle. Sacculus broad throughout its length, concave, interior ventral margin not produced. Brachiola broad. Aedeagus (Fig. 4) short, broadening toward base, virtually straight, with three equal sized cornuti. Female genitalia (Fig. 5) (n==15). Sterigma broad and flat, lateral tips sharply pointed. Anterior apophyses extending anteriorly beyond tips of sterigma. Ostium bursae circular. Ductus bursae broad caudally with a dis­ tinctive bulbous projection at middle, narrow and rugose between the bul­ bous projection and corpus bursae. Antrum wide, posterior broadly grooved longitudinally. Biology. Perhaps univoltine, as no summer adult specimens were found in pristine condition. Moths emerge in late August or early September and pass the winter. Adults were captured in Michigan in September and October resting on the upper sides of the leaves oflarval foodplants. Brown coloration and sedentary habits make the moths inconspicuous. Known larval foodplants are silver maple, red maple, and white oak (Quercus alba L.). The larvae crumple and skeletonize leaves from the under­ side, tying the lobes with silk. Only the upper leaf epidermis is left intact. Larvae and new pupae are uniform pale green. Pupation occurs in a thin co­ 1997 THE GREAT lAKES ENTOMOLOGIST 73

4

Figs. 1-8. Acleris species. 1, Wings of A. semiannula holotype female. 2, Wings ofA. stadiana holotype male. 3, Genitalia ofA. semiannula male from Isabella Co., Mich. (prep. MS 96009). 4, Aedeagus of A. semiannula male from Macomb Co., Mich. (prep. MS 96042). 5, Genitalia ofA. semiannula fe­ male from Gratiot Co., Mich. (prep. MS 96155). 6, Genitalia of A. stadiana holotype male. 7, Aedeagus of A. stadiana male from Columbia, Mo. (prep. MS 96137). 8, Genitalia of A. stadiana female from Hampton, N. H. (prep. MS 96143).

coon among webbing on the crumpled leaf. A leaf may harbor more than one pupa. The pupa extends from the cocoon during emergence. Material examined (n=56). PENNSYLVANIA: Holotype (Fig. 1). ILLI­ NOIS: Putnam Co., 1m, 09/29/1967, genit. prep. MS 96010, reared from white oak; 1m, 10/14/1968, genit. in glyc., reared from soft maple (INHS). 74 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

KENTUCKY: Barren Co., If, 08/3-411971, genit. prep. JAP 3768 (EME). MAINE: Bar Harbor, If, 09/2111937, reared from Acer rubrum (USNM). MARYLAND: Garrett Co., Garret SF, If, 0711111995, genit. in glyc. (JDG). MICHIGAN: Gratiot Co., 1m, If, 0911411991; If, 0911711991; If, 09/18/1991, genit. prep. MS 96155 (Fig. 5), all reared from Acer rubrum; Isabella Co., 4m, 13f, 09/2111981, genit. prep. MS 95077; 1m, 09/23/1981; 1m, 09/24/1981; 2m, 2f, 09/2811981, genit. prep. MS 96009 (Fig. 3); 1m, 10/1011981, genit. prep. WEM 241951, all reared from Acer saccharinum; If, 09/2211981, genit. prep. WEM 2111951; 3m, If, 10/09/1991, all on leaves ofAcer saccharinum; 3m, H, 10/09/1991, all on leaves ofAcer saccharinum (RJP); Grand Traverse Co., 1m, 07/0911960, genit. prep. MS 96063; Macomb Co., 1m, 01125/1944, genit. prep. MS 96042 (Fig. 4) (MSUC). MISSISSIPPI: Warren Co., H, 12130/1971, genit. prep. MS 96190 (MATH). NEW HAMPSHIRE: Enfield, If, 1110411978, genit. in glyc. (YPM); Hampton, If, 04/26/1909, genit. prep. Obr. 408 (AMNH). OHIO; Athens Co., Hocking College, If, 06117/1995, genit. in glyc. (DP). VER­ MONT: Brandon, Otter Creek, 1m, 10/27/1991, genit. prep. MS 96019; Colch­ ester, Sunny Hollow Nature Area, If, 10/2411991, genit. in glyc.; S. Burling­ ton, If, 07/09/1992, genit. prep. MS 96020; Guildhall, 1m, If, 10/2611991, genit. preps. MS 96017, MS 96018, 1m, 10/26/1991, genit. in glyc. (UMSP); Chittenden Co., Hinesburg, If, 10110/1991, genit. prep. MS 96205; 2f, 10/1811991, genit. prep. MS 96152c (GRN). VIRGINIA: Fairfax Co., Alexan­ dria, 1m, 05/29/1976, at light, genit. prep. JAP 4518 (EME).

Acleris stadiana (Barnes & Busck), New Status (Figs. 2, 6-8) Peronea stadiana Barnes & Busck (1920: 217) (Holotype: male, Ottawa, On­ tario, 18.9.05, C. H. Young, genit. prep. MS 96122, forewing length 7.0 rom, in USNM, wings shown here in Fig. 2); Forbes (1923: 485). Peronea ferrugana (not Denis & Schiffermuller 1775); Forbes (1923: 487) (in part), McDunnough (1934: 321) (in part), Procter (1946: 3(8), Brower (1983: 49) (in part). Peronea semiannula (not Robinson 1869); McDunnough (1934: 322, figs. 328(8),332(9), (1935: 138), (1939: 59), Prentice (1965: 735), Brower (1983: 49). Acleris tripunctana (not Htibner,1796-99); Obraztsov (1963: 223) (in part), Opheim (1964: 302, pI. 1, fig. 7) (in part). Acleris semiannula (not Robinson 1869); Mackay (1962: 18, fig. 12), Obraztsov (1963: 230), Razowski (1966: 459, pI. 35, Figs. 3, 4 ), Grehan et a1. (1995: 22), Ferge (1995: 29), Grehan & Sabourin (1995: 24-25, pI. 3, Figs. 26, ?27). Female and Male. Wingspan 11.2-17.0 mm (n=89). Forewing with costal triangle. Ground color of forewing light buff to amber. Color of costal margin within costal triangle same as rest of forewing or darker. Costal tri­ angle varying from tawny to dusky brown, darker than ground, at times barely discernible depending on shade of ground. Costal triangle coloration occasionally continues from its apex to dorsal wing margin. Hindwing paler than forewing. Male genitalia (Figs. 6, 7) (n=27). Ventral margin of sacculus slightly concave, a slight interior rise at middle. Aedeagus (Fig. 7) long, thin, sharply bent, with two pairs of cornuti separated by a chitinous plate. Female genitalia (Fig. 8) (n=19). Sterigma broad and flat, lateral tips 1997 THE GREAT LAKES ENTOMOLOGIST 75 rounded and blunt. Anterior apophyses projecting anteriorly no farther than lateral tips of sterigma. Ostium bursae circular. Ductus bursae with a sclero­ tized ring around neck, broad through most of its length, narrowed at junc­ tion with corpus bursae. Antrum small, sclerotized along rim. Biology. Bivoltine. Prentice (1966) stated that A. stadiana (as A. semi­ annula) is a solitary leaf roller on birch (Betula), with pupae appearing in late June and again in late August. Mackay (1962) described the larva (as A. semiannula). Fall generation adults overwinter. Material examined (n=89). ONTARIO: Holotype (Fig. 2, 6). CON­ NECTICUT: New Haven Co., North Haven, 1£, 11106/1959; Tolland Co., Mansfield, 1m, 11115/1958 (YPM); New Haven Co., Hamden, 1£, 10/18/1968, reared from Betula populifolia, genit. in glyc. (USNM). ILLINOIS: Algon­ quin, 1£, 03/25/1985, genit. prep. MS 96145 (INHS). MAINE: Bar Harbor, 07/17f??, Mount Desert Is., 10/05!??, (WBP); Lincoln, If, OS/211??, genit. in glyc. (USNM). MARYLAND: Adelphi, 1£, 05/29/1969, genit. in glyc. (USNM). MASSACHUSETTS: Lancaster, pine barrens, 1m, 11/05/1993, genit. prep. MS 96111 (UMSP). MICHIGAN: Cheybogan Co., Ocqueoc Lk., 3f, 07/25-26/1996, at blacklight, genit. preps. MS 95068, MS 96146; 1£, 07/27/1994 (USNM); Chippewa Co., Tahquamenon SP, 1m, 09/27-29/1995, at UV light, genit. prep. MS 96207; Clinton Co., Dansville SGA, 1£, 03124/1994, genit. in glyc. (RDK). MINNESOTA (?): 1m (no locality), C, 04/10/1892, genit. prep. MS 96007 (UMSP). MISSOURI: Columbia, 1m, 06/10/1969, genit. prep. MS 96137 (Fig. 7) (JRH). NEW HAMPSHIRE: Enfield, 1m, 10/26/1978, genit. in glyc. (YPM); Hampton, 1m, 04/27/1911, genit. prep. MS 97102; 1£, 11110/1911, genit. prep. MS 96143 (Fig. 8); 1£, 11120/1904, genit. prep. MS 96159 (CMNH). NEW JERSEY: Montclair, If, 11/02/1903; 1£, 11/0111923, genit. prep. MS 97103 (CMNH); Montclair, 1£, 11102/1903 (EME). NOVA SCOTIA: Armdale, If, 04113/1948, genit. in glyc.; Kings Co., Aylesford, 1£, 10/03/1950 (YPM). PENNSYLVANIA: Finleyville, 1f, 11/201?? (CMNH). RHODE ISLAND: Elmwood, 1m, 1£, 10/16/1920, genit. preps. MS 97104, MS 96144 (CMNH). VERMONT: Addison Co., Bristol, 1m, 07/28/1991, genit. prep. MS 96030; Chittenden Co., Burlington, 1m, 0410611993, genit. in glyc., 1m, 03/27/1993, genit. prep. MS 96149; Colchester, 1f, 0411411991, genit. in glyc.; 1m, 10/25/1991, genit. prep. MS 96148; 1m, 04/20/1992, genit. prep. MS 96154e; 1f, 09/28/1992; 1m, 10/16/1993, genit. prep. MS 96107; 1m, 11/10/1993, genit. prep. MS 95070; Essex Town, Sleepy Hollow Rd., 1m, 04/04/1991, genit. in glyc.; Jericho Research Forest, 1m, 05/08/1993, genit. prep. MS 95042; S. Burlington, If, 07/1711991, genit. prep. MS 96153d; 1f, 07/2511992; 1f, 04115/1993, genit. in glyc.; Guildhall, 1m, 10/26/1991, genit. in glyc.; Rutland Co., Chittenden, 1m, 10/27/1991, genit. prep. MS 96113 (UMSP); Chittenden Co., Hinesburg, 1m, 11/0211968, genit. prep. MS 96150a; 1f, 10/3111991, genit. prep. MS 96151b (GRN); Underhill St. Pk., e1. 715m, If, 08/20/1995, genit. prep. MS 96132 (ERL). WISCONSIN: Bayfield Co., If, GMP #748050B, 08/20-09/06/1974, reared from Betula papyrifera; H, (no locality), GMP #764356B, 09117/1976, reared from Betula papyrifera; If (no locality), 09113/1976, reared (foodplant unrecorded), genit. prep. MS 96135; Polk Co., Gibson Lk., 1f, 06/29/-07/07/1960, reared (foodplant un­ recorded) (UWElVD; Oneida Co., Lk. Katherine, 1m, 04/13/1961, genit. prep. JAP 980; 2m, 0411811961, genit. prep. JAP 2769; 10m, 2f, 04121/1961, genit. preps. JAP 1418, JAP 2765, genit. in glyc.; 3m, 05/07/1961; 1m 05/22/1961; If, 06/01/1961; 2m, 07/10/1961, genit. prep. JAP 1421; 1m, 07/16/1961, genit. prep. JAP 1410; 1m, 07/20/1961; 1m, 07/2211961, genit. prep. JAP 2770; 1m, 10/08/1961; 2m, 10/0911961, genit. prep. JAP 2771; 2m, 3f, 10/16/1961; 1m, 10/19/1961; 2m, 04/22/1962; 1m, 07/04/1962 (EME); Vilas Co., W side of Car­ lin Lk., H, 07/02/1987 (GJB). 76 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

DISCUSSION Polymorphism and geographic variation in exterior traits of Acleris are often so pronounced that genitalia provide the only reliable diagnostic char­ acters. Even then, occasionally one sex only may be distinguished by geni­ talia. Acleris semiannula and A. stadiana differ from each other most notably in genital structure. The lateral tips of the sterigma are flat and sharply pointed in A. semian­ nula, whereas they are round and blunt inA. stadiana. The anterior apophy­ ses extend beyond the lateral tips of the sterigma in A. semiannula, but not inA. stadiana. The ductus bursae inA. semiannula has a bulbous projection at its middle, then narrows rugosely to the corpus bursae, whereas that in A. stadiana is broad to just before the corpus bursae (Figs. 5, 8). The aedeagus ofA. stadiana is long, narrow and sharply bent, whereas that of A. semian­ nula is short, broadening toward base and virtually straight (Figs. 4, 7). Su­ perficially, the contrast between fore- and hindwings is greater in A. stadi­ ana than in A. semiannula (Figs. 1, 2), A. semiannula varies more in the intensity of wing color, and A. stadiana varies more in wing pattern and body size. We found the A. semiannula and A. stadiana type specimens and their labels to be in excellent condition for usability. The sexes of A. semiannula were associated through numerous specimens reared by us from the same foodplant at the same locality; those of A. stadiana, by museum specimens reared from the same foodplant and locality, reared from the same foodplant at different localities, and from adults collected at the same localities. Several other Acleris species can be similar to A. semiannula and A. sta­ diana superficially. The occasional continuation of costal-triangle coloration from the triangle apex to the dorsal wing margin in some A. stadiana is like that in A. braunana (McDunnough). Paler specimens of A. stadiana match some specimens of A. comandrana (Fernald) and A. subnivana (Walker). Darker specimens of A. semiannula match some specimens ofA. implexana (Walker),A. oxycoccana (Packard), andA. kearfottana (McDunnough). We refer to A. semiannula, A. stadiana, and their look-alikes as the A. semiannula species complex. Members of the complex inc1udeA. schalleriana viburnana (Clemens), A. oxycoccana, A. subnivana, A. cervinana (Fernald), A. braunana, A. comandrana, A. implexana, A. cornana (McDunnough), A. simpliciana (Walsingham), A. negundana (Busck), A. ferrugana (Denis & Schiffermuller), A. kearfottana, A. caryosphena (Meyrick), and A. notana (Donovan). These taxa have wingspans less than 20 mm, whitish to dark brown forewing coloration, and possess a costal triangle. Some specimens can have finely reticulated wings, or costal-triangle coloration continuing from the triangle apex to the dorsal wing margin. Some species may have a useful superficial character such as the emarginate forewing costal margin in A. subnivana, but it bears emphasis that reliable identification must be based on genitalia. In Eurasia, A. notana and A. ferrugana frequently have been confused with each other (Obraztsov 1957, Bradley et al. 1973). In North America, nearctic members ofthe complex have not only been mistaken for each other, but for palaearctic counterparts. Several nearctic species of the complex are close enough to the palearctic A. notana and A. ferrugana to have been mis­ taken for them. Obraztsov's (1963) report of A. notana (as A. tripunctana) from New Hampshire is actually A. obtusana fuscana (Busck), a taxon also confused with A. notana in Eurasia (Kyrki 1982). The report by Godfrey et al. (1987) ofA. notana (as A. tripunctana) in Illinois is based on a misidenti­ fied specimen of A. semiannula. Although we were unable to locate the two 1997 THE GREAT LAKES ENTOMOLOGIST 77

Canadian specimens McDunnough (1934) identified as A. ferrugana, we have not otherwise encountered A. ferrugana or A. notana in the Nearctic. Sources useful for identifying taxa of the complex are Bentinck and Diakonoff (1968), McDunnough (1934), Razowski (1966), and Wolff (1964). Based on male genital characters, the following key separates A. semian­ nula and A. stadiana from other species of the complex.

1. Ventral margin of sacculus deeply emarginate just before cucullus ...... oxycoccana, s. uiburnana. 1.' Ventral margin of sacculus not deeply emarginate ...... 2 2. Ventral margin of sacculus bulging at middle...... caryosphena, notana, implexana, braunana, ceruinana. 2.' Ventral margin of sacculus not bulging at middle ...... 3 3. Ventral margin of sacculus straight or slightly concave ...... 4 3.' Ventral margin of sacculus markedly concave ...... 6 4. Aedeagus possessing fewer than four cornuti..... simpliciana, subniuana. 4.' Aedeagus possessing four or more cornuti...... 5 5. Aedeagus possessing four cornuti and a separate sclerite ...... stadiana. 5.' Aedeagus possessing four or more cornuti and no separate sclerite ...... cornana. 5." Aedeagus possessing five cornuti and a separate sclerite .... comandrana. 6. Aedeagus possessing three cornuti ...... semiannula. 6.' Aedeagus possessing other than three cornuti ...... ferrugana, negundana, kearfottana.

ACKNOWLEDGMENTS 'Ye thank the following for specimen loans or use of facilities: D. Azuma, G. J. Balogh, I. M. Borak, R. L. Brown, P J. Clausen, D. Furth, J. R. Grehan, J. R. Heitzman, R. W. Hodges, S. Krauth, R. D. Kriegel, B. Landry, B. Mather, K. Methven, J. S. Miller, G. R. Nielsen, M. C. Nielsen, B. R. Parker, T. :'1. Peters, E. L. Quinter, J. E. Rawlins, F. H. Rindge, F. W. Stehr, and J. H. 'Vilterding. We also thank R. W. Holzenthal for the use of photographic equipment, J. Eibling for first calling the larval damage to our attention, and PT. Dang, T. Wallenmaier, and C. E. Weyland for other assistance with the study.

LITERATURE CITED Barnes, W. & A. Busck. 1920. Notes and new species. Contributions to the Natural History of the Lepidoptera of North America 4: 211-248. Bradley, J. D., W. G. Tremewan, & A. Smith. 1973. British tortricoid moths, Cochylidae and Tortricidae: Tortricinae. Ray Society, London. 251 pp. Brower, A. E. 1983. A list of the Lepidoptera of Maine. Part 2, the microlepidoptera, Section 1, Limacodidae through Cossidae. Maine Agr. Exp. Sta. Tech. Bull. 109, 60 pp. Denis, J. N. C. M. & I. Schiffermiiller. 1775. Ankundigung eines systematischen Werkes von den Schmetterlingen der Wienergegend, herausgegben von einigen Lehrern am k. k. Theresianum. 323 pp. [not seen] Ferge, L. 1995. Season summary. News Lepid. Soc., June 1995, 32 pp. Fernald, C. H. 1882. A synonymical catalogue of the described Tortricidae of North America north of Mexico. Trans. Amer. Entomol. Soc. 10: 1-64. 78 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

1902 [1903]. Family Tortricidae, pp. 448-489. In: Dyar, H. G. (ed.), A list of North American Lepidoptera. U. S. N atl. Mus. Bull. 52. Forbes, W. T. M. 1923. The Lepidoptera of New York and neighboring States. Part 1. Cornell Univ. Agr. Exp. Sta. Mem. 68, 729 pp. Godfrey, G. L., E. D. Cashett, & M. O. Glenn. 1987. Microlepidoptera from the Sandy Creek and Illinois River region: an annotated checklist of the suborders Dac­ nonypha, Monotrysia, and Ditrysia (in part) (Insecta). Illinois Natural History Sur­ vey Spec. Publ. 7, 44 pp. Grehan, J. R & M. Sabourin. 1995. Maple feeding Tortricidae of the northeastern United States: guide to identification of adults. Vermont Agr. Exp. Sta. & Vermont State Misc. Pub. 117, ¥MC Res. Rep. 11,47 pp. Grehan, J.R, B. L. Parker, G. R Nielsen, D. H. Miller, J. D. Hedbor, M. Sabourin, & M. S. Griggs. 1995. Moths and butterflies of Vermont (Lepidoptera): a faunal check­ list. Vermont Agr. Exp. Sta. and Vermont State Misc. Pub. 116, VMC Bull. 1, 95 pp. Hubner, J. 1796-99. Sammlung europii.ischer Schmetterlinge, Tortrices. [not seen] Kyrki, J. 1982. Acleris obtusana, a good species (Lepidoptera, Tortricidael. Notulae En­ tomol. 62: 37-42. Mackay, M. R 1962. Larvae of the North American Tortricinae (Lepidoptera: Tortrici­ dael Can. Entomol. SuppL 28, 182 pp. McDunnough, J. 1934. The Canadian species of the tortricid genus Peronea. Can. J. Res. 11: 290-332 __. 1935. Xew Canadian eucosmids with notes (Lepidoptera). Can. Entomo!. 67: 140-149. 1939. Check list of the Lepidoptera of Canada and the United States of Amer­ ica. Part 2, microlepidoptera. Mem. South. Calif. Acad. Sci. 2, 171 pp. ;\-leyrick, E. 1912. Tortricidae, 71 pp. In: Wagner, H. (ed.), Lepidopterorum Catalogus 10. Obraztsov, N. S. 1957. Die Gattungen der palaearktischen Tortricidae. 1. Allgemeine Aufteilung der Familie und die Unterfamilien Tortricinae und Sparganothinae. 3, Fortsetzung und Schluss. Tijdschr. Entomo!. 100: 309-347. Obraztsov, N.s. 1963. Some North American moths of the genus Acleris (Lepidoptera: Tortricidae). Proc. U.S. Natl. Mus. 114: 213-217. Opheim, M. 1964. The genus Acleris Hubner, 1825. Notes on the Norweigan Tortrici­ dae 2 (Lepidoptera). Norsk Entomol. TIdsk. 12: 296-314. Powell, J. A. 1983. Tortricidae, pp. 31-41. In: R W. Hodges (ed.). Check list of the Lepi­ doptera ofAmerica north of Mexico. London. 284 pp. Prentice, RM. 1966. Forest Lepidoptera of Canada recorded by the Forest Insect Sur­ vey, microlepidoptera. Can. Dept. Forest. Pub!. 4: 543-840. Procter, W. 1946. Biological survey of the Mount Desert region, Vo!' 7. The insect fauna. 566 pp. Wistar Institute ofAnatomy and Biology, Philadelphia. Razowski, J. 1966. World fauna of the Tortricini (Lepidoptera, Tortricidae). Panstw. Wydawn. Nauk., Krakow. 576 pp. Robinson, C. T. 1869, Notes on the American Tortricidae. Trans. Am. Entomo!. Soc. 2: 261-288. Shaw, S. A. 1905. List of micro-Lepidoptera taken in Hampton, New Hampshire. Ento­ mol. News 16: 323-327. Walsingham, T. de Grey, Sixth Earl. 1879. Illustrations of typical specimens of Lepi­ doptera Heterocera in the collection of the British Museum. Pt. 4, North American Tortricidae. 76 pp. Wolff, N. L. 1964, The Lepidoptera of Greenland. Medd. om Gr0nl. 159(11),74 pp. Zeller, P. C, 1875. Beitrage zur Kenntnis der nordamericanischen Nachtfalter, beson­ ders der Microlepidopteren. Verh. Zool-Bot. Ges. Wien 25: 205-360. 1997 THE GREAT LAKES ENTOMOLOGIST 79

NOTES ON THE LIFE HISTORIES OF ACROSTERNUM HILARE AND COSMOPEPLA BIMACULATA (HETEROPTERA PENTATOMIDAE) IN SOUTHERN ILLINOIS

J. E. McPherson and D. L Tecle l

ABSTRACT The life histories ofAcrosternum hilare and CosmopepZa bimaculata were studied in southern Illinois from May 1972 to September 1974 and from Sep­ tember 1992 to June 1995. Both species were bivoltine, overwintered as adults, and became active in early spring. The subsequent generations were characterized by marked overlapping of the nymphal instars. No active adults were found after early November.

Acrosternum hilare (Say) and Cosmopepla bimaculata (Thomas) are com­ mon phytophagous stink bugs that occur over much ofAmerica north of Mex­ ico (Froeschner 1988, McPherson 1982); both are common in southern Illinois (McPherson 1982). Much has been published on their biology, including an­ nuallife cycles, food plants, laboratory rearing, immature stages, and preda­ tors and parasites (McPherson 1982). However, the life cycles have not been documented thoroughly throughout the bugs' geographic ranges. Acrosternum hilare, the green stink bug, is of major economic impor­ tance, particularly as a pest of agricultural crops (e.g., soybeans) in the mid­ western, southern, and southeastern United States (McPherson 1982). As would be expected, its annual life cycle has been studied in relation to its damage to these crops. However, it has been listed as both uni- and bivoltine, with reports of two generations per year from studies in more southern loca­ tions. For example, it has been reported as univoltine in Canada (Javahery 1990), Utah (Sorenson and Anthon 1936), central Illinois CEsselbaugh 1948), Ohio (Whitmash 1917), and Virginia (Underhill 1934); and bivoltine in Kansas (Wilde 1969), Arkansas (Miner 1966), and South Carolina (Jones and Sullivan 1982). Presuming this reported difference is correct, and reflects the geographic locations of these earlier studies, then southern Illinois would ap­ pear to be in a transition zone between these broad geographic areas. The annual life cycle of C. bimaculata is poorly understood. Information consists primarily of scattered notes, although it apparently is univoltine in Alberta, Canada (McDonald 1968). From May 1972 to September 1974, one of us (JEM) was involved in a survey of the Pentatomoidea of the LaRue-Pine Hills Ecological Area (now LaRue-Pine Hills Research Natural Area) (hereafter referred to as Pine Hills), which is located in Union Co. in southern Illinois (McPherson and Mohlenbrock 1976; also see same pUblication for description of Pine Hills). Two of the 49 species recorded were A. hilare and C. bimaculata. Both were

IDepartment of Zoology, Southern Illinois University, Carbondale, 1L 62901. 80 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3 common as adults, but low numbers of nymphs made determination of the life cycles difficult. A. hilare was listed as apparently bivoltine and C. bimac­ ulata as probably bivoltine. From September 1992 to June 1995, the Pine Hills pentatomoids were resurveyed to ascertain changes that had occurred in this fauna during the intervening years. Once again, A. hilare and C. bimaculata were found to be common as adults. However, nymphs for each species also were collected in substantial numbers. As the data from the earlier survey still were available, they were combined with those from the resurvey to gain a better under­ standing of the annual life cycle of each species. Presented here, then, are the results ofthe combined surveys, including host plants.

MATERIALS AND METHODS Collecting trips were taken weekly from September to November 1992, March to December 1993 and 1994, and May to June 1995. The March and N ovemberlDecember collecting periods were before the bugs had emerged from and after they had entered overwintering sites, respectively. Specimens were collected by hand-picking and sweeping. The disappearance of the in­ sects in the fall generally corresponded to the senescence ofhost plants. Host plants were identified primarily with keys by Mohlenbrock (1986) but also by Britton and Brown (1913), Gleason and Cronquist (1991), Jones (1963), and Mohlenbrock and Voigt (1959). The SIUC herbarium served as

APRIL

ADULT "'132

~ 1etiNSTAR c( N_12 ::I Q ~ !i 2ndIHSTAR No" ~ !i W 3rd INSTAR ~ N." W £L

4U1INSTAR N....

sthlNSTAR N.1H

Fig. 1. Field life cycle ofA. hilare. Percentage of individuals ofeach stage per sample collected during 1972-1974 and 1992-1995 seasons in Union County, IL. 1997 THE GREAT LAKES ENTOMOLOGIST 81 reference for key characters and confirmation of species identification. Letter designations following host plants are as follows: A, adults; F, feeding; C, cop­ ulation; E, eggs; N, nymphs.

RESULTS AND DISCUSSION Acrosternum hilare (Say) Adults emerged from ovenvintering sites in late April-early May and were found continuously until late October-early November (Figs. 1-2). Al­ though the subsequent nymphal instars were not found in perfect chronologi­ cal sequence, their times of occurrence, numbers collected, and associated peaks of abundance indicate that A. hilare is bivoltine in southern Illinois. Thus, its life cycle is similar to that reported for the southern United States. Host plants recorded during both surveys combined included Allium canadense L. (A), Ambrosia trifida L. (N), Ceanothus americanus L. (N), Celtis occidentalis L. (N), Cercis canadensis L. (A,N), Cornus drummondii Meyer (F,N), Elymus uirginicus L. (A), Eupatorium rugosum Houttuyn (N), Hybanthus concolor (Forster) (A), Impatiens capensis Walter (A,F,N), Impa­ tiens pallida Nuttall (A,F,N), Perilla frutescens Britton (F,N), Rhus glabra L. (A, F), Rubus allegheniensis Porter (N), Teucrium canadense L. (A), Trades­ cantia subaspera Ker (N), Ulmus rubra Muhlenberg (A), Verbascum thapsus L. (A, E, N), Verbesina alternifolia Britton (N), Vicia uillosa Roth (F,N), and Vitis aestivalis Michaux (A, N).

A""'l ..AY JUNE I JULY AUGUST HPTEMBEJl OCTOBER NOVEMBER ..50 AIlULT...... 25 50 i I 25 ,.UNSTAR !l ".12 0( 25 ::I C 50 5= 25 2ndINSTAR is 0 ~ N.e7 25 !5 50 25 !iii 3ldIHSTAR W 0 N ... II: 25 - W - c.. 50 25 ~ I 41hINSTAR N.1il1 25 50 25 51h !NSTAR 0 I N.133 25 50

Fig. 2. Field life cycle of A. hilare. Percentage in each sample of total individ­ uals of same stage collected during 1972-1974 and 1992-1995 seasons in Union County, IL. 82 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

50 25 ADULT N=157 25 50 25 lotlNSTAR ~ 0 N=2G « 25 :::l C SO • :; 25 is 2ndiNSTAR ~ N.45 U. 25 0 50 I­ Z 25 UJ 3N1INSTAR (.) N.53 a: 25 UJ Il. 50 25 4thiNSTAR 0 N.59 25 50 25 SthlNSTAR N.l09 25 50 Fig. 3. Field life cycle of C. bimaculata. Percentage of individuals of each stage per sample collected during 1972-1974 and 1992-1995 seasons in Union County, IL.

Cosmopepla bimaculata (Thomas) The annual life cycle of this species was not as clear at that found for A. hilare. Adults emerged from overwintering sites in early May and were found until mid-late October (Figs. 3-4). The subsequent occurrences of eggs and nymphs indicate this species is bivoltine. Eggs clusters were found from late May to early June on V. thapsus (n=7 ) and Geranium carolinianum L. (n= 5) and from mid-to late August on Scrophularia marilandica L. (n 18). Nymphs of the first generation were found approximately from late May through late July, primarily on V. thapsus and G. carolinianum, with a peak of fifth instars (most frequently collected of the instars) in late June-early July. Nymphs of the second generation were found approximately from early August to late September, primarily on S. marilandica and T. canadense, with a peak of fifth instars in early September. The collection of young in­ stars (lsts-3rds) concurrent or subsequent to the collection of the last fifth instars of the second generation may indicate a partial third generation but probably is the result ofsampling error. 1997 THE GREAT LAKES ENTOMOLOGIST 83

MAY JlJIIE JULY AUGUST I SE1'11:MBER OCTOBER 50 25 ADULT 0 1'1-157 25 - 50 25 1s1INSTAR 0 en N.2O «..J 25­ ::;)c 50 • :; 25 2ndiNSTAR ...... 0 is N.45 ~ - ~ 25 - II.o 50 25 !z 3rdiNSTAR ...... IJJ 0 o N=53 """II"""", ex: 25 a..IJJ 50 2S 4thiNSTAR ...... 0 N.S9 - ...... 25 - 50 25 ! 5thiNSTAR 0 .... N.1OQ 2S "'" 50 I I Fig. 4. Field life cycle of C. bimaculata. Percentage in each sample of total in­ dividuals of same stage collected during 1972-1974 and 1992-1995 seasons in Union County, IL.

Additional host plants recorded during both surveys combined included Bromus inermis Leyser (A), Bromus secalinus L. (A,F), Campsis radicans (L.) (A), Daucus carota L. (A,F), Phleum pratense L. (A), Plantago lanceolata L. (F,N), Rumex crispus L. (A,F), and Veronica arvensis L. (A,F,C).

ACKNOWLEDGMENTS We thank Scott Bundy and Lane Loya, former graduate students, De­ partment of Zoology, Southern Illinois University at Carbondale, for their as­ sistance with field work; and the U.S.D.A. Forest Service, Harrisburg, for permission to collect in the LaRue-Pine Hills Research Natural Area. This re­ search was funded, in part, by the U.s.D.A. Forest Service (Cooperative Agreement No. 23-92-43). 84 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

LITERATURE CITED Britton, N. L. and A. Brown. 1913. An illustrated flora of the northern United States and Canada. VoL I: 1-680; Vol. II: 1-735; Vol. III: 1-637. Dover Pub., NY. (Reprinted in 1970). Esselbaugh, C. O. 1948. Notes on the bionomics of some midwestern Pentatomidae. EntomoL Amer. 28: 1-73. Froeschner, R. C. 1988. Family Pentatomidae Leach, 1915. The stink bugs, pp. 544-597. 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, NY. 958 pp. Gleason, H. A. and A. Cronquist. 1991. Manual of vascular plants of northeastern United States and Canada. New York Botanical Garden, Bronx, NY. ixxv + 910 pp. Javahery, M. 1990. Biology and ecological adaptation of the green stink bug (Hemiptera: Pentatomidael in Quebec and .Ontario. Ann. EntomoL Soc. Am. 83: 201-206. Jones, G. N. 1963. Flora of Illinois (3rd ed'). Am. Midland Nat. Monog. No.7. Univer­ sity of Notre Dame, IN. vi + 410 pp. Jones, W. A. and M. J. Sullivan. 1982. Role of host plants in population dynamics of stink bug pests of soybean in South Carolina. Environ. EntomoL 11: 867-875. McDonald, F. J. D. 1968. The life history of Cosmopepla bimaculata (Thomas) (Het­ eroptera: Pentatomidael in Alberta. Quaest. EntomoL 4: 35-38. McPherson, J. E. 1982. The Pentatomoidea (Hemiptera) of northeastern North Amer­ ica (with emphasis on the fauna of Illinois). Southern Illinois University Press, Car­ bondale and Edwardsville. ix + 240 pp. McPherson, J. E., and R. H. Mohlenbrock. 1976. A list of the Scutelleroidea of the LaRue-Pine Hills Ecological Area with notes on biology. Great Lakes EntomoL 9: 125-169. Miner, F. D. 1966. Biology and control of stink bugs on soybeans. Arkansas Agr. Exp. Sta. Bull. 708: 1-40. Mohlenbrock, R. H. 1986. Guide to vascular flora of southern Illinois. Southern Illinois University Press, Carbondale and Edwardsville. viii + 507 pp. Mohlenbrock, R. H. and J. W. Voigt. 1959. A flora of southern Illinois. Southern Illinois University Press, Carbondale. 390 pp. Sorenson, C. J. and E. W. Anthon. 1936. Preliminary studies of Acrosternum hilare (Sayl in Utah orchards. Proc. Utah Acad. Sci., Arts, and Letters 13: 229-232. Underhill, G. W. 1934. The green stinkbug. VirginiaAgr. Exp. Sta. Bull. 294: 1-26. Whitmarsh, R. D. 1917. The green soldier bug. Nezara hilaris Say. Order, Hemiptera. Family, Pentatomidae. A recent enemy in northern Ohio peach orchards. Ohio Agr. Exp. Sta. BulL 310: 519-552. Wilde, G. E. 1969. Photoperiodism in relation to development and reproduction in the green stink bug. J. Econ. EntomoL 62: 629-630. 1997 THE GREAT LAKES ENTOMOLOGIST 85

NEW RECORD OF BRACHYCERCUS MACULATUS BERNER (EPHEMEROPTERA: CAENIDAE) FROM NEW YORK AND A KEY TO LARVAE OF NORTHEASTERN SPECIES

Steven K. Burian 1, Margaret A. Novak2, Robert W. Bode2 and Lawrence Abele2

ABSTRACT Brachycercus maculatus, a member of a rare group of mayflies, is now recorded for the first time from New York State in the upper Hudson River. An illustrated key to the Brachycercus larvae of northeastern North America is provided to spur further study ofthe genus in the region.

Species of the genus Brachycercus Curtis include some of the most enig­ matic mayflies. Larvae are small with ocellar tubercles and occur among fine detritus and shifting sediments of lentic and slow-flowing lotic habitats. Al­ though current distribution records indicate that the genus is broadly dis­ tributed, larvae are rarely collected. Adults are small, short-lived, and en­ countered even less frequently than larvae. Virtually nothing is known about the swarming behavior or biology of most North American species. Taxo­ nomic studies of Brachycercus (Berner 1950, Soldan 1986, Berner and Pescador 1988), which contain little information on biology, constitute most of what is known about North American species. Brachycercus maculatus Berner is a species that was believed to have been restricted to north-central Florida, but recently it was confirmed from North Carolina (M. Pescador, pers. comm.). The purpose ofthis paper is to re­ port the discovery of B. maculatus from the upper Hudson River in New York. We also present, a regional key to the larvae of the four species of Brachycer­ cus now known to occur in New York, New England, and southern Quebec. Brachycercus maculatus Berner 1946, NEW RECORD-NEW YORK: (11 Larvae) Warren Co., Hudson River, Corinth, milepoint 214 [43 0 14' 55"N/073° 49' 57"W], 7 July 1994, R.W.Bode; (4 Larvae); Saratoga Co., Hud­ son River, Waterford, milepoint 157 [42 0 47'19"N/073° 40' 38"Wl, 7 July 1994, R.W.Bode. Brachycercus maculatus larvae were collected in Petite Ponar grab sam­ ples (APHA, 1985) taken during ambient water quality monitoring at the vil­ lage of Corinth and at Waterford on the upper Hudson River. Physical and chemical water quality parameters for both collection sites are presented in Table 1. Corinth samples containing Brachycercus larvae were obtained 20 m upstream of a hydroelectric dam and 5 m from the northeastern shore. Wa­ terford samples were taken 20 m from the eastern shore. The substrate in Ponar grab samples at both sites consisted of sand, silt, and organic material

lDept. of Biology, Southern Connecticut State University 501 Crescent St., New Haven, CT, 06515. 2New York State Dept. of Environmental Conservation, 50 Wolf Rd., Albany, NY 12233-3502. 86 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

Table 1. Physical and chemical parameters of Brachycercus sites on the Hudson River. Chemical habitat parameters recorded at the time of collection are listed with annual ranges. Annual ranges of chemical parameters were obtained from Firda et al. (1993,1994, and in press).

Site Vel. a Width Deptha Flow Temp. Spec. D.O. pH (mls) (m) (m) (m3/s) (ac) Cond.d Corinth 1.0 110 2.0 23.0 45 CO. 1-24.6) (33-57) (6.6-7.3)

Waterford 0.5 300 4.0 230.4C 26.4 133 8.1 7.8 (0.2-30.0) (98-177) (3.3-14.0) (7.0-7.8) aTaken at mid-channel. bMeasured at USGS gage at Lake Luzerne 8.0 km upstream of site. "Measured at USGS gage 5.3 km upstream of site. dCmhos/cm @ 25°C)

(mostly leaves). Specimens of B. maculatus were only found in samples that contained fine sediments with few leaves or macrophyte stems. Samples with a large amount ofplant matter contained no Brachycercus. Larvae taken at both sites ranged in length from 2.66 to 5.75 mm, with a mean of 4.43 mm. Several had black wing pads indicating that emergence was imminent at the time of collection. Among these specimens body lengths ranged from 4.58 to 5.75 mm, with a mean of 5.23 mm. Previously, adult emergence has only been reported for B. lacustris (Lyman 1955) and B. flavus (Harper and Harper 1976). Both of these species have emergences that occur in late July and continue into August, whereas B. maculatus seems to be emerging in early July. Lyman (1955) showed that B. lacustris had a prolonged emergence that peaked in early August. Harper and Harper (1976) reported a similar peak emergence for B. flavus in early August. Un­ fortunately, because no other specimens ofB. maculatus were obtained it was not possible to determine the duration or the peak of its emergence at the Hudson River sites. In Florida adults of B. maculatus have been reported in February, April, and July (Berner and Pescador 1988). Four species of Brachycercus (B. fluvus Traver, B. lacustris (Needham), B. nitidus (Traver), and B. maculatus) are now known from New York, New England, and southern Quebec. Among the these species B. nitidus is most distinctive and easily identified. However, the remaining species are very similar. Differences among diagnostic characters described by Traver (1935), Berner (1946, 1950), Burks (1953), Soldan (1986), and Berner and Pescador (1988) are subtle and may eventually be shown to be within the range of variation a single widely distributed species. But any decision on this matter will require a thorough revision of the genus. Until that occurs the following key, based on our studies and the decriptive works listed above, will aid in in­ dentifying the larvae of these taxa as they are now defined. We hope that this will spur interest and futher study ofBrachycercus in this region.

Key to Larvae ofBrachycercus of Northeastern North America 1. Pronotum with a pair of well developed lateral spines (Fig.l); body length 7.0-9.0 mm ...... B. nitidus 1997 THE GREAT LAKES ENTOMOLOGIST 87

3 dm1~ I ~~ Figures 1-7. 1-3: Head and pronotum, arrows indicate pronotal spines or ridges; 1, B. nitidus; 2, B. maculatus; 3, B. lacustris; 4-5: Left lateral view of abdomen showing posterolateral projections, segment 2 is labeled; 4, B. flavus;5, B. lacustris; 6-7: Dorsal view of right gill covers; 6, B. flavus; 7, B. lacustris. 88 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

1'. Pronotum without lateral spines, but may have a pair oftransverse ridges (Fig. 2); body length less than 7.0 mm ...... 2 2. Pronotum with well developed transverse ridges that are distinct from the lateral margins to the midline (Fig. 2) ...... E. maculatus 2'. Pronotum smooth with weakly developed transverse ridges that may ap­ pear not to meet the lateral margins (Fig. 3) ...... 3 3. Posterolateral spines on abdominal segment 2 sharp and greater than half the length of spines on segment 3 (Fig. 4); gill covers with distinct dark spots near basal articulation, spots do not extend to the intersec­ tion ofridges on gill covers (Fig. 6) ...... B. fiavus 3'. Posterolateral spines on abdominal segment 2 blunt and only about half the length ofthose on segments 3 (Fig. 5), bases of spines on segment 2 wider than those on segments 3-6; gill covers without distinct spots, but may have dark shading that extends to the intersection of ridges on gill covers (Fig. 7) ...... B. lacustris

ACKNOWLEDGMENTS We thank Dr. M. Pescador for confirming our identifications and for pro­ viding information on unpublished records of B. maculatus. We also thank Dr. WL. Peters for the loan of material from the aquatic insect collection at Florida A&M University and the New York Dept. of Conservation for their support for this paper.

LITERATURE CITED APHA. 1985. Standard methods for the examination of water and wastewater (l6th edition). American Public Health Association, Washington, D.C. 1268 pp. Berner, L. 1946. New species of Florida mayflies (Ephemeroptera). Fla. Entomo!. 28: 60-82. Berner, L. 1950. The Mayflies of Florida. 1:niv. of Florida Studies, BioI. Sci. Series no. 4,267 pp. Berner, L. and M. Pescador. 1988. The Mayflies of Florida, Revised Edition. Univ. Presses ofFlorida, Gainesville, Florida, 415 pp. Burks, B.D. 1953. The mayflies, or Ephemeroptera, of Illinois. Bull. Ill. Nat. Hist. Surv. 26: 1-216. Firda, G. D., Lumina, R, Murray, P. M., and Freeman, W. O. 1993. Water Resources Data-New York, Water Year 1993, Volume 1. Eastern New York excluding Long Is­ land. 1:. S. Geological Survey Water Data Report NY-93-1, 452 pp. Firda, G. D., Lumina, R, Murray, P. M., and Freeman, W. O. 1994. Water Resources Data-New York, Water Year 1994, Volume 1. Eastern New York excluding Long Is­ land. 1:. S. Geological Survey Water Data Report NY-94-1, 488 pp. Firda, G. D., Lumina, R, Murray, P. M., and Freeman, W. O. (In Press). Water Re­ sources Data-New York, Water Year 1995, Volume 1. Eastern New York excluding Long Island. C S. Geological Survey Water Data Report. Harper, F. and P. P. Harper. 1976. Inventaire et phenologie des Ephemeropteres du lac Saint-Louis, Quebec. Ann. Soc. ent. Quebec 21: 136-143. Lyman, F.E. 1955. Seasonal distribution and life cycles of Ephemeroptera. Ann. Ento­ mol. Soc. Amer. 48: 380-391. Soldan, T. 1986. A revision of the Caenidae with ocellar tubercles in the nymphal stage (Ephemeroptera). Acta Universitatis Carolinae-Biologica 1982-1984: 289-362. Traver, J.R 1953. Part II, Systematic. pp. 239-739 In: J.G. Needham, J.R Traver, and Y.C. Hsu (eds.), The biology of mayflies with a systematic account of North American species. Comstock Pub!. Co., Ithaca, New York. 1997 THE GREAT lAKES ENTOMOlOGIST 89

HEXAGENIA BILINEATA (EPHEMEROPTERA: EPHEMERIDAE) PERSISTS AT LOW LEVelS OF ABUNDANCE IN THE LOWER FOX RIVER, WISCONSIN

Philip A. Cochran 1 and Andrew P. Kinzigerl,2

ABSTRACT After burrowing mayflies (Hexagenia bilineata) were first noted in the vicinity of the DePere Dam on the Fox River in 1991, adults have been ob­ served in small numbers each summer since then. It is possible that the Fox River population has remained at low levels because of an Allee effect. In ad­ dition, it is possible that the population is still limited by poor environmental quality, presumably in the upper layer of sediment inhabited by the larvae. 'l\vo other relatively sensitive species associated with benthic habitat, the sea lamprey (Petromyzon marinus) and the lake sturgeon (Acipenser ful­ vescens), have been observed in the Fox River in recent years. Collectively these species provide an indication of improved environmental conditions, but it is not yet clear that any of the three have established populations ca­ pab.le of successfully reproducing in the lower Fox River on a consistent basIs.

Adults of the burrowing mayfly Hexagenia limbata (Serville) formerly achieved nuisance levels in Green Bay of Lake Michigan (Schuette 1928), but reduced water quality caused by input from the lower Fox River resulted in their complete elimination by 1969 (Harris et al. 1987). It was noteworthy, therefore, when Cochran (1992) observed adult H. bilineata (Say) in the vicinity of the DePere Dam on the lower Fox River, 12 km upstream from Green Bay, during the summer of 1991. Herein we report that H. bilineata persists in the vicinity of the DePere Dam but has not achieved the large emergences characteristic of this genus elsewhere in its range (Fremling 1960). Adult mayflies were collected during routine activities in the vicinity of the DePere Dam, including a daily commute by the senior author on foot over the Claude Allouez Bridge. In addition, student technicians in our biology de­ partment who lived during the summer in the vicinity ofthe St. Norbert Col­ lege campus (just upstream from the DePere Dam) were shown specimens of H. bilineata and asked to collect any that they observed. We have used the specific designation "Hexagenia bilineata" throughout this manuscript to refer to our collections because any specimens for which identifications could be made belonged to this species (some individuals were damaged or were not captured because they were out of reach). However, because H. limbata

IDivision of Natural Sciences, St. Norbert College, DePere, WI 54115-2099. 2Present address: Dept. of Biology, Frostburg State lJniversity, Frostburg, MD 21532. 90 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

Table 1. Collections of adult Hexagenia bilineata in the vicinity of the DePere Dam on the lower Fox River, Brown County, Wisconsin. Multiple sightings on a given date are indicated in parentheses. The 1997 season was in progress at the time of manuscript revision. Year Dates of Occurrence Total 1992 June 24 (2), July 4, July 8, July 11 5 1993 July 28, July 29 2 1994 July 8 (3), July 14, July 31 5 1995 July 1, July 17, July 20 3 1996 June 28, June 29 2 1997 July 7, July 18 (4), August 19 6

occurred in this system historically (Schuette 1928) and conceivably might return, and because some authors we have cited did not specify which species they were discussing, we have used only the generic name in those contexts in which either or both species might be involved. Adult H. bilineata were collected each summer during the years 1992-1997 (Table 1), albeit in low numbers. This period includes the "flood" year of 1993, when summer monthly mean discharges at various locations along the Fox River were among the highest on record (Holmstrom et al. 1996). Although H. bilineata has been present in the Fox River since at least 1991, it has not achieved the characteristically large emergences noted in other parts of its range. Independent evidence suggests that population den­ sity of Hexagenia in the lower river is low: only a single individual was col­ lected in 58 Ekman grabs collected over a 10 km reach of river (both up­ stream and downstream from the DePere dam) during the period 10 June-ll August, 1992 (John Rafferty, personal communication). We suggest two pos­ sible explanations for the apparent lack of a popUlation increase. These hy­ potheses are not mutually exclusive. First, the Hexagenia population in the lower Fox River may be subject to an Allee effect (Brewer 1988), by which reproduction is relatively inefficient at low population levels. Such an effect would tend to keep a newly reestab­ lished population at low numbers. Indeed, it has been suggested that the large synchronized emergences typical of many mayflies increase the proba­ bility of finding a mate (Corbet 1964) or increase survival through satiation of predators (Sweeney and Vannote 1982). We note the consistent presence near the DePere Dam of several species of birds that feed on emergent in­ sects, including a colony of cliff swallows (Petrochelidon pyrrhonota) that nest on the Claude Allouez Bridge. Second, it is possible that populations of Hexagenia in the lower Fox River are still limited by poor environmental quality, presumably at the sedi­ ment-water interface (e.g., Sullivan et al. 1983). Evidence to support this hy­ pothesis might be provided by another species, the sea lamprey (Petromyzon marinus), first reported from the Fox River in 1991 (Cochran 1994). It may be no coincidence that neither species has been subsequently observed in large numbers. Like Hexagenia, sea lampreys undergo a burrowing larval stage of relatively long duration and relatively low tolerance for environmen­ tal degradation. Krieger et al. (1996) recently described the recovery of Hexagenia spp. in western Lake Erie. That swarms of adults were not recorded until approxi­ 1997 THE GREAT LAKES ENTOMOLOGIST 91 mately two decades after water quality responded to pollution abatement may be consistent with an Allee effect. Kolar et at (1997) developed a model of recolonization of western Lake Erie that suggested that recovery to pre­ 1950s population densities would take 10-41 years if population growth was logistic, even in the absence ofsuch factors as low-oxygen events, competition with chironomids, toxic sediments, and fish predation. Addition of any of these factors to the model typically led to substantially increased recovery times. The presence-and absence--of Hexagenia elsewhere in the Green Bay drainage has been recently noted. Lillie (1995) recorded at least some indi­ viduals of one or more species in the Wolf, Menominee, and Peshtigo rivers but he found none in the Oconto or Embarrass rivers (it should be noted that his surveys were targeted heavily toward mayflies that inhabit sand-bot­ tomed habitat). Choudhury et al. (1996) specifically noted the absence of Hexagenia from the diet oflake sturgeon (Acipenser fulvescens) in Lake Win­ nebago because it is such an important component of lake sturgeon diets in other systems. They attributed the lack of Hexagenia to the highly eutrophic nature of Lake Winnebago. Kempinger (1996) found Hexagenia in 9.9% of 131 age-O lake sturgeon captured in the Wolf River. A link between Hexage­ nia and lake sturgeon, such as that suggested by Cavender (1997) in Lake Erie, is of especial interest in the present context. The lake sturgeon, like Hexagenia bilineata and the sea lamprey, is a sensitive benthic species re­ cently observed in the lower Fox River (Cochran 1995). All three species are indicators of improved water quality, but in all three cases, it is unclear ifthe benthic habitat has improved to the point that successful reproduction can occur on a consistent basis.

ACKNOWLEDGMENTS We thank Andrew Cochran, Eric Golden, and Rick Wagner for collecting some of the specimens reported herein. Some of our observations were made during field work funded by the Wisconsin Sea Grant Institute under grants from the National Sea Grant College Program, National Oceanic and Atmo­ spheric Administration, United States Department of Commerce, and the State of Wisconsin (Federal grant NA90AA-D-5G469).

LITERATURE CITED Brewer, R. 1988. The science of ecology. Saunders College Publishing, Philadelphia, Pennsylvania. 922 pp. Cavender, T.M. 1997. Status of the lake sturgeon population in Lake Erie. American Society of Ichthyologists and Herpetologists, 77th Annual Meeting, p. 94 (ab­ stract). Choudhury, A., R. Bruch, and TA Dick. 1996. Helminths and food habits of lake stur­ geon Acipenser fuluescens from the Lake Winnebago system, Wisconsin. Amer. MidI. Nat. 135:274-282. Cochran, P.A. 1992. The return of Hexagenia (Ephemeroptera: Ephemeridael to the lower Fox River. Great Lakes Entomologist 25:79-81. 1994. Occurrence and significance of sea lamprey (Petromyzon marinus) in the lower Fox River, Wisconsin. Trans. Wisc. Acad. ScL, Arts, Lett. 82:17-21. 1995. Lake sturgeon (Acipenser fulvescens) in the lower Fox River, Wisconsin. Sturgeon Quarterly 3(4):8-9. 92 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

Corbet, P.S. 1964. Temporal patterns of emergence in aquatic insects. Can. Entomol. 96:264-279. Fremling, C.R 1960. Biology of a large mayfly, Hexagenia bilineata (Say), of the Upper Mississippi River. Iowa St. Univ. Agr. Home Econ. Exp. Sta. Res. Bull. 482:842-852. Harris, H.J., P.E. Sager, C.J. Yarbrough, and H.J. Day. 1987. Evolution of water re­ source management: a Laurentian Great Lakes case study. Internat. J. Env. Studies 29:53-70. Holmstrom, B.K, D.L. Olson, and B.R Ellefson. 1996. Water resources data-Wiscon­ sin, Water Year 1995. U.S. Geological Survey Water-Data Report WI-95-1. Kempinger, J.J. 1996. Habitat, growth, and food of young lake sturgeons in the Lake Winnebago system, Wisconsin. N. Amer. J. Fish. Man. 16:102-114. Kolar, C.S., P.L. Hudson, and J.F. Savino. 1997. Conditions for the return and simula­ tion of the recovery of burrowing mayflies in western Lake Erie. Ecological Ap­ plications 7:665-676. Krieger, KA., D.W. Schloesser, B.A. Manny, C.E. Trisler, S.E. Heady, J.J.H. Ci­ borowski, and KM. Muth. 1996. Recovery of burrowing mayflies (Ephe­ meroptera: Ephemeridae: Hexagenia) in western Lake Erie. J. Great Lakes Res. 22:254-263. Lillie, RA. 1995. A survey of rare and endangered mayflies of selected rivers in Wis­ consin. Wisc. Dept. Nat. Res., Research Report 170,23 pp. Schuette, H,A. 1928. The Green Bay fly in 1819: an "extraordinary visitation." Green Bay Historical Bulletin 4(2): 1-3 (Note: this reference was inadvertently omitted from Cochran 1992). Sullivan, J.R, J.J. Delfino, C.R Buelow, and T.B. Shefry. 1983. Polychlorinated biphenyls in the fish and sediment of the Lower Fox River, Wisconsin. Bull. En­ vironm. Contam. 'lbxicol. 30:58-64. Sweeney, B.W. and RL. Vannote. 1982. Population synchrony in mayflies: a predator satiation hypothesis. Evolution 36:810-821...... _-_....._------_.­

1997 THE GREAT lAKES ENTOMOLOGIST 93

EVALUATION OF PAEDERUS UTTORARIUS (COLEOPTERA:STAPHYLINIDAE) AS AN EGG PREDATOR OF CHRYSOTEUCHIA TOPIARIA (LEPIDOPTERA: PYRALIDAEl IN WISCONSIN CRANBERRY BOGS

Sandra Haase-Statz 1

ABSTRACT A preliminary study was conducted to determine if the rove beetle, Paederus littorarius Grav., would exhibit a feeding preference for the eggs of the pyralid moth, Chrysoteuchia topiaria Zeller, a pest in Wisconsin cran­ berry bogs. Individuals were offered a choice of C. topiaria eggs or Drosophila sp. adults for four days. Total number of prey items eaten was converted to weight using a multiplier based on the mean weight of 20 indi­ viduals of each prey item, respectively. A significant preference for Drosophila adults was observed in the preference trial; however as many as 24 C. topiaria eggs in addition to Drosophila offerings were consumed by P. littorarius individuals within a 24 h period. Additionally, laboratory and field observations suggests P. littorarius is a polyphagous predator.

The cranberry girdler, Chrysoteuchia topiaria Zeller (Lepidoptera: Pyrali­ dae) is a sporadic but important pest of cranberries Vaccinium macrocarpum Aiton. Larvae attack the plant by chewing the underground stems and dis­ rupting nutrient flow. Control of this pest can be problematic for several rea­ sons. First, larvae may be overlooked because damage is not usually seen until the spring following attack. Second, the larval stage is spent hidden in the leaf litter and soil, and is difficult to monitor. Lastly, the adult, which can be monitored during its 6-8 week flight period, can not be controlled with pesicide applications because the pest's flight period coincides with host plant blossoming. Chemical control can not be used when cranberries are in blossom due to the neccessity of pollinator activity (Roberts 1983, Mahr and Moffitt 1994). . The rove beetle, Paederus littorarius Gravenhorst, is a common endemic predator of small arthorpods in North American riparian habitats. It is a polyphagous and opportunistic predator which consumes a variety of prey items in the laboratory (S. Haase-Statz unpublished data). Paederus spp. are extra-oral feeders that attack their prey and suck the hemolymph from the body, and discard the exoskeleton. They feed on all life stages ofother insects but have been shown to have a preference for eggs and early instar larvae (Ahmed 1957, Kurosa 1958, Manley 1977). However, there is no published in­ formation specifically regarding the feeding habits of North American Paederus, nor has there been any work evaluating them as predators in North American agricultural systems.

IDepartment of Entomology, University of Wisconsin, Madison, WI 53706. 94 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

Paederus spp. in Asia, the Middle East and South America have been identified as important predators of a variety of agriculturally significant phytophagous insects. For example, P. alferii Koch has been effective in con­ trolling many cotton pests, P. fuscipes Curtis is important in the control of several rice pests, and P. columbinus is effective against sugar cane pests. Additionally, Paederus spp. occur in clover, maize, peas, beans, potatoes and banana plantations (Frank and Kanamitsu 1987). The objective of this study was to conduct a preliminary evaluation of P. littorarius as a predator in an agricultural system important to Wisconsin. The cranberry system was chosen based on its riparian nature which pro­ vides suitable habitat for Paederus spp. Additionly, P. littorarius had been collected in pitfall traps placed in cranberry bogs, establishing their occur­ rence there (Coffield unpublished data). Chrysoteuchia topiaria was selected as the target pest for two reasons: (1) C. topiaria eggs are oviposited randomly on the soil surface of the bog and larvae are found in the leaf litter under the cranberry vines (Roberts, 1983). P. littorarius is commonly found in leaf litter scavenging on the soil surface where it forages; (2) C. topiaria eggs and larvae are present when adult and larval Paederus are actively foraging. The study was conducted to answer the following questions: Will P. littorarius feed on C. topiaria eggs? What is the rate of consumption? And will P. littorarius preferentially feed on C. topiaria eggs?

METHODS To determine whether P. littorarius would feed on Chrysoteuchia eggs and if so, at what rate consumption occurred, adult beetles was placed in 5" petri dishes; one per petri dish. Each petri dish was lined with damp filter paper. Beetles were then starved for 24 h. Sixty Chrysoteuchia eggs were placed in each petri dish and the beetles were given 24 h to feed. Remaining food items were removed and counted to determine total number eaten. One replicate was completed. The average number of eggs eaten per beetle for 2 d was calculated. To determine if P. littorarius would feed preferentially on Chrysoteuchia eggs, the predators were offered a choice between the target prey and frozen Drosophila adults. Drosophila were used as the alternative prey item be­ cause they had been used to successfully rear P. littorarius and were avail­ able. This trial used ten female P. littorarius and was conducted over four days. Comparisons were based on mean consumption of each individual over 4 days. The predators were placed in individual petri dishes with wet filter paper. Each was starved initially for 24 h and subsequently starved for 6 h between feedings. Beetles were given 24 h to feed. Remaining prey items then were removed and counted. The number eaten was converted to mass using a multiplier (Drosophila =0.8253 mg; C. topiaria eggs 0.0357 mg). The multiplier for each prey item was determined by weighing 20 individuals of each prey item and calculating the mean weight per prey item. At the first feeding, two (» 1.4 mg) Drosophila and 15 (" 0.6 mg) Chrysoteuchia eggs were placed in each petri dish. Eight of the ten beetles consumed all of the prey items in the first feeding; therefore, five Drosophila (» 4.0 mg ) and 25 (» 1.0 mg) Chrysoteuchia eggs were placed in each dish for subsequent feedings. The null hypothesis that there was no preferential feeding was tested using a t-test for paired means. The statistical expert in Quattro Pro 5.0 soft­ ware (MS-DOS version by Borland International, 1994) was used to analyze the data. -

1997 THE GREAT LAKES ENTOMOLOGIST 95

Adult P. littorarius used for the trials were reared from field collected adults. C. topiaria eggs were obtained from field collected adults using the methods of Roberts (1983). Drosophila were obtained from a genetics lab at the University of Wisconsin-Madison.

RESULTS Paederus littorarius adults began feeding on C. topiaria eggs immedi­ ately after they were placed in the containers. The mean number of eggs con­ sumed in 24 h was 31.5 ± 11.16, range was 21-54 (n=lO for 2 trials). In the preference trial, ten adult female P. littorarius ate between 0-22 eggs per day (Table 1), an average of 8.93 eggs (0.319 ± 0.16 mg) (Fig. 1); and between 0-4 Drosophila (Table 1), an average of 2.54 adults (2.094 ± 0.38 mg) (Fig. 1). The t-test showed a significant preference for Drosophila (p <0.00001).

DISCUSSION In the non-preference test, Paederus adults were shown to feed readily on the eggs of C. topiaria. However, they showed a marked preference for

Table 1. Daily prey consumption by Paederus littorarius. Each beetle was provided five Drosophila and 25 Chrysoteuchia topiaria eggs simultaneously. Number ofDrosophila eaten per day Beetle 1 Day 2 3 4 Mean Mean Wt 1 2 3 4 0 2.25 (±1.5) 1.86 (±1.2) 2 2 3 3.5 3 2.88 (±0.5) 2.37 (±0.4) 3 2 4 3 4 3.25 (±0.8) 2.68 (±0.7) 4 2 3 1.5 2 2.13 (±0.5) 1.75 (±OA) 5 2 3.5 5 2 3.13 (±1.2) 2.58 (±l.O) 6 2 3 2 1 2.00 (±0.7) 1.65 (±0.6) 7 2 2 4 0 2.00 (±1.4) 1.65 (±1.2) 8 2 4 4 1 2.75 (±1.3) 2.27 (±l.1) 9 1.5 4 3 0 2.13 (±1.5) 1.93 (±1.3) 10 1.5 3 4 3 2.88 (±0.9) 2.75 (±0.7)

Number ofChrysoteuchia topiaria eggs eaten per day 1 15 20 11 16 15.50 (±3.2) 0.55 (±0.1) 2 7 0 3 0 2.50 (±2.9) 0.09 (±0.1) 3 1 1 15 6 5.75 (±5.7) 0.21 (±0.3) 4 1 1 19 7 7.00 (±7.3) 0.25 (±0.3) 5 15 0 4 0 4.75 (±6.1l 0.17 (±0.2) 6 14 0 4 13 7.75 (±5.9) 0.28 (±0.2) 7 18 14 14 22 17.00 (±3.3) 0.61 (±0.1) 8 16 7 0 1 6.00 (±6.4) 0.21 (±02) 9 18 16 4 8 11.50 (±5.7) 0.41 (±0.2) 10 14 15 8 9 11.50 (±3.0) 0.41 (±0.1) 96 THE GREAT lAKES ENTOMOLOGIST Vol. 30, No.3

4

T

3 - ..-., en I E '-' c:: o a - T § 2 en - c:: o () c:: ca (!) ~ 1 l j T ;r if

jI [II ~ o 1 \ ~ ~ tW L­ t n~ 1 2 3 4 5 6 7 8 9 10

Beetles

I ===:J Chrysoteuchia I I Drosophila I

Figure 1. Consumption ofDrosophila adults and Chrysoteuchia topiaria eggs, expressed in milligrams of prey over a period of four days. Each beetle was provided with five Drosophila and 25 C. topia ria eggs daily.

Drosophila adults when given a choice. There are several possible explana­ tions for this strong showing of preference: (1) P. littorarius truly prefer Drosophila over all other prey items; (2) they prefer small dipteran prey over other types; (3) there is an energy benefit to taking a few large prey items over several small prey items; or (4) the individual predators had developed an artificial preference for Drosophila from having been reared and main­ tained on this food source. Additional information about the feeding habits of P. littorarius was 1997 THE GREAT LAKES ENTOMOLOGIST 97 gathered during rearing studies. Field collected adults were maintained on a variety of food sources which included: small dipteran adults (Chironomidae and Cecidomyiidae), Collembola and hemipterans (Cicadellidae and Cercopi­ dae) reared adventitiously from soil obtained at Paederus collection sites; early instar larvae of Plodia interpunctella Hubner (Pyralidae) and Psuedo­ plusia includens Walker (Noctuidae); various large dipteran larvae (Cal­ liphoridae, Sarcopha 'dae, Muscidae); and a variety of other odd offerings (eg. roast beef, ch cheese, and bran muffin). thus supporting the as­ sumption that P. littorarius is a polyphagous predator. It should be noted, however, that P. littorarius did not thrive on many of these prey items and Drosophila was the only prey item that has been successfully used to rear Wisconsin Paederus spp. in the lab (S. Haase-Statz, unpublished data). It is unknown if P. littorarius contributes to the control of C. topiaria. It is probably an important part of the natural enemy complex for this pest. Field observation and collection data for P. littoranus indicate that they are found in a variety of moist habitats, including wet meadows, marshes, bogs, roadside ditches and leaf litter such as are found within and surrounding cranberry beds. As stated earlier, a few adult P. littorarius were collected in pitfall traps placed within cranberry beds located near Warrens, Wisconsin, in 1991 (Cockfield, unpubL). I surveyed eight commercial cranberry beds lo­ cated on three differenct cranberry farms. Although I was not successful in locating any additional specimens in the beds, I did collect Paederus littorar­ ius in grass adjacent to beds and along streams and drainage ditches in the vacinity of commercial cranberry production. Based on collection data, it seems reasonable to assume that P. littorar­ ius is more likely to be found in unmowed grassy areas around cranberry beds than in the beds themselves. Kamm et aL (1983) concluded that the cranberry girdler prefers grassy areas surrounding cultivated areas, and leave that habitat when it becomes undesirable. Therefore, P. littorarius could be useful in regulating cranberry girdler populations in source habi­ tats.

ACKNOWLEDGMENTS I thank Dr. Daniel Mahr for advice on this project, Jane Soijka for infor­ mation regarding cranberry girdler outbreaks, and all the growers who al­ lowed access to their property. This study was supported in part by Hatch Grant WS-03570 to Dr. Daniel KYoung.

LITERATURE CITED Ahmed, M. K 1957. Life-history and feeding habits of Paederus alfierii Koch (Coleoptera: Staphylinidae). Bull. Soc. Entomol. Egypte 41: 129-143. Frank, J. H., and Kanamitsu. K 1987. Paederus, sensu lato (Coleoptera: Staphylin­ idae): natural history and medical importance. J. Med. Entomol. 24 (2): 155-19l. 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 nursey stock. J. Econ. Entomol. 76: 923-926. Kurosa, K 1958. Studies on poisonous beetles. III. Studies on the life history of Paederus fuscipes Curtis (Staphylinidae). Japan. J. Sanit. Zool. 9: 245-76. :".!ahr, S. E. R. and L. J. Moffitt. 1994. Biologic and Economic Assessment of Pesticide Usage on Cranberry. NAPIAP Report Number 2-CA-94. 98 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

Manley, G. V. 1977. Paederus fuscipes (Coleoptera: Staphylinidae): a predator of rice fields in West Malaysia. Entomophaga 22: 47-59. Roberts, S. L. 1983. Studies of the Cranberry Girdler Chrysoteuchia topiaria in Wis­ consin. M.S. thesis, Univeristy ofWisconsin-Madison. Roberts, S. L., and D. L. Mahr. 1986. Development of Cranberry Girdler, Chrysoteuchia topiaria (Lepidoptera: Pyralidael in relation to temperature. Great Lakes Entomol. 19 (2): 85-90. 1997 THE GREAT LAKES ENTOMOLOGIST 99

EFFECTS OF FEEDING BY TWO FOLIVOROUS ARTHROPODS ON SUSCEPTIBILITY OF HYBRID POPLAR CLONES TO A FOLIAR PATHOGEN

Kier D. Klepzigl,2, Daniel J. Robison 2,3, Eugene B. Smalleyl and Kenneth F. Raffa 2

ABSTRACT We investigated variation in folivore-induced effects on subsequent plant suitability to a foliar pathogen. We used a leaf disk assay to expose three clones of hybrid poplar, NC11382, NE332 and NM6, to colonization by a leaf spot pathogen, Septoria musiua. Undamaged leaf disks of NE332 were the most resistant to S. musiua, followed by NM6 and NC1l382, respectively. To test the effects of prior herbivory on subsequent susceptibility to this fungal pathogen, we inoculated S. musiua on leafdisks taken from leaves which had been exposed to feeding by Tetranychus mites or cottonwood leaf beetles. Prior activity by mites and cottonwood leaf beetle affected the subsequent susceptibility ofclones NC 11382 and NE332 to S. musiua.

Herbivory may have many effects on host plants. One such effect, in­ duced plant resistance to herbivores and pathogens, may result from bio­ chemical and physiological changes initiated by contact with the invading herbivore (e.g., Clausen et a1. 1989). Although herbivores and plant pathogens may be confronted with and affected by similar induced defenses (Schultz 1983, Krischik et al. 1991, Hammerschmidt 1993, Klepzig et al. 1996), these interactions have largely been studied separately. Studies exam· ining reciprocal effects between herbivores and plant pathogens are con­ ducted even less frequently, perhaps due to the complexity of these multi­ species interactions (Wargo and Houston 1974, Karban et a1. 1987, Krischik et a1. 1991, Klepzig et al. 1996). In particular, we know little about how folio vore feeding affects subsequent susceptibility to foliar pathogens. The purpose of this study was to consider how prior herbivory by folivo­ rous arthropods and host plant clone interact to affect subsequent suitability to a foliar pathogen. Our study system consisted of two folivorous arthro­ pods-Tetranychus spp. (Acari: Tetranychidae) mites and cottonwood leaf beetles, Chrysomela scripta Fabr. (Coleoptera: Chrysomelidae), three hybrid

IDepartment of Plant Pathology, University of Wisconsin-Madison, Madison, WI 53706. 2Department of Entomology, University of Wisconsin-Madison, Madison, WI 53706. 3Current address: Hardwood Research Cooperative, Jordan Hall - Box 8008, North Carolina State University, Raleigh, NC 27695. Correspondence: K.D. Klepzig, USDA Forest Service, Southern Research Station, 2500 Shreveport Hwy, Pineville, LA 71360. 100 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3 poplar clones (Populus spp.), and one fungal foliar pathogen-Septoria mu­ siva Peck.

MATERIALS AND METHODS We grew plants from dormant hardwood cuttings from coppiced field grown hybrid Populus clones NC11382 (P. nigra X P. berolinensis), NE332 (P. simonii X P. berolinensis) and NM6 (P. nigra X P. maximowiczii) (Robison and Raffa 1994, 1996) at the University of Wisconsin Arlington Experiment Station. We stored the cuttings (5 cm long) frozen until use, and planted them in Redi-Earth Peat-Lite® soil mix. We regularly flood irrigated and fer­ tilized with 15 g Osmocote® slow-release fertilizer (7-16-12, plus micronutri­ ents), plants and grew them in a glasshouse at 16:8 L:D photoperiod, 18-29°C and 25-80% relative humidity. We assayed leaves from three types of plants. Because, within clones, variability is greater between leaves of the same physiological age than it is between trees (Robison and Raffa 1997), we removed leaves 3 and 4 (down­ ward from the apical fully expanded leaf) from each plant for use in assays the same day. We collected leaves from control (undamaged) plants of all three clones, exhibiting no visible signs or symptoms of arthropod damage or disease. We collected leaves from clones 11382 and NE332 that exhibited ex­ tensive mite feeding damage (approximately 30-50% damage on each leaf), or exhibited C. scripta feeding damage (approximately 10% defoliation on each plant). We followed the S. musiva colonization assays developed by Ostry and Skilling (1988). Due to our incomplete understanding of the mechanisms un­ derlying any systemic responses within poplar, and because multiple infec­ tions may occur on the same leaf we followed the method cited above and re­ moved multiple leaf disks from each leaf and used leaf disk as our experimental unit. Leaf disks (18 mm diameter) were placed abaxial side up in wells cut into 2% water agar in glass petri plates. Each petri plate con­ tained 6 disks taken from the same clone and treatment (n == 4 plates per clone/treatment combination). We inoculated five leaf disks in each chamber with 0.1 ml of a spore suspension (1 x 106 conidia/mI), and the sixth disk with 0.1 ml of sterile distilled water. The assay plates were then incubated in a growth chamber at 24°C under continuous light. We measured areas of the resulting necrotic lesions using a transparent dot grid. The effects of treat­ ments, clones, and time since inoculation on lesion size were analyzed using a repeated measures ANOVA, followed by Fisher's protected least squares means comparisons where appropriate (Abacus Concepts 1989).

RESULTS The leaf disk area colonized by S. musiva varied significantly by clone, and clone X treatment interaction (Table la). On leaf disks from undamaged plants, areas colonized by S. musiva varied significantly by clone (Table 1b), and were consistently largest on clone NC11382. Lesions were generally smallest on NE332, and intermediate on NM6 (Figure 1). Clonal effects on amount of fungal colonization were also detected. Sig­ nificant necrotic lesions formed on NC11382 by day 19, but not until days 26 and 33 on clones NM6 and NE332, respectively. By the experiment's end le­ sions on NM6 occupied up to 69% of the leaf disk surface area, nearly equiva­ lent to those on NC11382. Uninoculated control disks (N=4 for each treat­ 1997 THE GREAT lAKES ENTOMOLOGIST 101

Table 1. Analyses of variance for size of S. musiva lesions measured after inoculation of hybrid poplar leaf disks. CAl Two-way, repeated-measures ANOVA of clone (NC11382 and NE332) by treatment (mite-damaged, beetle-damaged and undamaged) effects. (B) One-way, repeated-measures ANOVA of clonal (NC11382, NE332 and NM6) effects (undamaged tissues). Greenhouse-Gilisser (G-G) and Hunyh-Feldt (H-Fl estimated p- values are given. A. Source df SS MS F P G-G H-F clone 1 64.78 64.78 52.28 .0001 treatment 2 4.43 2.22 1.79 .1714 clone "treatment 2 6.91 3.45 2.79 .0659 Subject (Group) 114 141.27 1.24 time 5 9.64 1.93 6.86 .0001 .0002 .0001 time * clone 5 29.61 5.92 21.09 .0001 .0001 .0001 time * treatment 10 2.78 0.28 0.99 .4501 .4307 .4336 time * clone" treatment 10 8.40 0.84 2.99 .0011 .0076 .0061 time'" Subject (Group) 570 160.07 0.28

B. Source df SS )'1S F P G-G H-F clone 1 20.11 20.11 11.26 .0014 subject 58 103.62 1.79 time 5 40.71 8.14 26.53 .0001 .0001 .0001 time'" clone 5 6.15 1.23 4.01 .0016 .0108 .0091 time" subject 290 88.99 0.31

ment within each clone) did not develop any necrotic lesions during the course of this experiment. The relationship between prior herbivory and subsequent fungal growth varied between clones. Lesions on leaf disks from undamaged leaves of clone NC11382 began to form after 2 weeks, but did not form on trees with prior herbivory by either mites or beetles until about 23 days. Lesions were larger than those formed on leaf disks from mite- or C. scripta damaged leaves until 26 days, and were significantly larger until 33 days post inoculation (Figure la). Average lesion size was significantly larger on mite damaged than on C. scripta damaged leaf disks at 33 days post inoculation. By the end of the ex­ periment (37 days post inoculation), up to 91% of the leaf disk surface had become necrotic. Using the curve fitting option in SuperANOVA (Abacus Con­ cepts 1989), we found significant linear relationships between lesion size and time since inoculation for undamaged (y =0.08 x - 0.86, r2 =0.96), mite dam­ a¥ed (y = 0.12 x -1.98, r2 = 0.92), and C. scripta damaged (y = 0.09 x -1.60, r = 0.93) leaf disks of clone NC 11382. Lesions began to form earlier on inoculated leaf disks from NE332 trees with prior herbivory than on undamaged trees (Figure Ib). At 37 days post inoculation, fungal colonized areas accounted for approximately 54% of the surface area. The average final colonized area on NE332 leaf disks was ap­ proximately equal to the average colonized area on leaf disks from clone NC11382 eleven days previous. Using the curve fitting option in SuperA­ 102 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

A.

25-.------, Prior Feedin& NCl1382 ____ None 2.0

1.5 ..•. ~.... Tetranychus 1.0

05 ..·e·. . C. scripta

10 15 20 25 30 35 40 B. 2.5..,....------, NE332 2.0 ---- None

1.5

.•.. ~.... Tetranychus

-• -e- - • C. scripta

10 15 20 25 30 35 40 c. 2.5,------. NM6 ---- None 2.0

1.5

1.0

0.5

0.0 +---1.,.."""'.:;::..-.1.-,---.----,---1 10 15 20 25 30 35 40 Time (days) Figure 1. Mean (± s.e.) size of S. musiua lesions on leaf disks removed from undamaged, Tetranychus mite damaged and cottonwood leaf beetle (C. scripta) damaged hybrid poplar trees at various sampling times. Data ana­ lyzed by repeated measures ANOVA. (A) Clone NC11382, (B) Clone NE332, (C) Clone NM6. 1997 THE GREAT LAKES ENTOMOLOGIST 103

NOVA (Abacus Concepts 1989), we found significant exponential relation­ ships between lesion size and time since inoculation on undamaged [y=(F10-7)*100.19X , r 2=0.94J, mite dama~ed [y=( 1*10-3)* 10008X, r2=0.95J, and C. scripta damaged [y=(2*10-4)B10o.11X, r =0.89] leaf disks of clone NE332.

DISCUSSION Undamaged tissue from clone NC11382 was more susceptible to S. mu­ siva than was similar tissue from NE332. The two clones also differed in the patterns of response with a more pronounced effect in clone NE332 than in clone NC11382. We cannot say for certain at this time, however, whether the alterations we sa\v in host susceptibility to a pathogenic fungus represent physiological induction due to herbivory. Other possible explanations may in­ volve alterations in phylloplane flora (Wilson 1995). On undamaged plants from all three clones lesions spread progressively faster on clone ::lli332, ~M6, and NC11382, respectively. These relative lev­ els of leaf symptom severity differ somewhat from field rankings of severity of stem cankers caused by the same fungus, in which clone NM6 was the most, clone NE332 was intermediate, and clone NC11382 was the least, re­ sistant (Robison and Raffa 1996). The interactions of hybrid poplar with its suite of potential plant para­ sites are highly variable. Each inducing agent and subsequent herbivore may elicit different responses depending upon the host genotype. Although a uni­ form theory on inducibility among artificial clones is not likely (Robison and Raffa 1994), the variation that is present can be very useful for tree improve­ ment and clonal deployment strategies. Moreover, inducibility needs to be an important component in understanding pest impacts and population changes in managed and natural ecosystems (Haukioja and Neuvonen 1988).

ACKNOWLEDGMENTS We thank Anthony Cina for technical assistance. The S. musiva culture was kindly supplied by M. Ostry, USDA Forest Service, St. Paul, MN. This research was supported by McIntire-Stennis project WIS 03014, the Wiscon­ sin Department of Natural Resources, and the University ofWisconsin-Mad i­ son College ofAgricultural and Life Sciences.

LITERATURE CITED Abacus Concepts. 1989. SuperANOVA. Abacus Concepts, Inc., Berkeley, CA Clausen, T. P, P. B. Reichardt, J. P. Bryant, R A. Werner, K. Post and K. Frisby. 1989. Chemical model for short-term induction in quaking aspen (Populus tremuloides) fo­ against herbivores. J. Chern. EcoL 15:2335-2346. Hammerschmidt, R 1993. The nature and generation of systemic signals induced by pathogens, arthropod herbivores, and wounds. Adv. Plant Pathol. 10:307-337. Haukioja, E., and S. Neuvonen. 1988. The autumnal moth Epirrita autumnata in Fennoscandia. In: A. A. Berryman red], Dynamics of forest insect populations. Plenum, New York pp 163-178 Karban, R, R Adamchak and W. C. Schnathorst. 1987. Induced resistance and inter­ specific competition between spider mites and a vascular wilt fungus. Science 235:678-680. 104 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

Klepzig, K. D., E. B. Smalley and K. F. Raffa. 1996. Combined chemical defenses against an insect-fungal complex. J. Chem. Ecol. 22:1367-1388. Krischik, V. A., R. W. Goth and P. Barbosa. 1991. Generalized plant defense: effects on multiple species. Oecologia 85:562-571. Ostry, M. E. and D. D. Skilling. 1988. Somatic variation in resistance of Populus to Septaria musiva. Plant Dis. 72:724-727. Robison, D. J. and K. F. Raffa. 1994. Characterization of hybrid poplar clones for resis­ tance to the forest tent caterpillar. For. Sci. 40:686-714. Robison, D.J., and K. F. Raffa. 1996. Productivity, drought tolerance, and pest status of hybrid Populus: Tree improvement and silvicultural implications. Biomass and Bioenergy: Robison, D.J., and K. F. Raffa. 1997. Effects of constitutive and induced traits ofhybrid poplars on forest tent caterpillar feeding and population ecology. For. Sci. 43:252-267. Schultz, J. C. 1983. Impact of variable plant chemical defenses on insect susceptibility to parasites, predators, and diseases. Symp. Amer. Chem. Soc. 208:37-55. Wargo, P. M., and D. R. Houston. 1974. Infection of defoliated sugar maple trees by Armillaria mellea. Phytopathology 64:817-822. Wilson, D. 1995. Fungal endophytes which invade insect galls: Insect pathogens, be­ nign saprophytes, or fungal inquilines? Oecologia 103:255-260. 1997 THE GREAT LAKES ENTOMOLOGIST 105

UROPHORA AFFINIS AND U. QUADRIFASCIATA (DIPTERA: TEPHRITIDAE) RELEASED AND MONITORED BY USDA, APHIS, PPQ AS BIOLOGICAL CONTROL AGENTS OF SPOTIED AND DIFFUSE KNAPWEED

R. F. lang,] R. D. Richard] and R. W. Hansen]

ABSTRACT USDA, APHIS, PPQ has distributed the seedhead gall flies Urophora affinis and U. quadrifasciata (Diptera: Tephritidae) as classical biological agents of the introduced weeds spotted and diffuse knapweed (Centaurea maculosa and C. diffusa, respectively) (Asteraceae) in the United States. From 1987 to 1996, Urophora spp. have been released in 97 counties in 14 midwestern and western states. Established populations of U. affinis and U. quadrifasciata are confirmed in 85 and 95 counties, respectively, among all 14 states. These include the first reports of successful establishment of Urophora spp. in Arizona (two counties), Colorado (eight counties), Michi­ gan (one county), Minnesota (six counties), Nebraska (four counties), Nevada (two counties), North Dakota (one county), South Dakota (four coun­ ties), Utah (three counties), and Wisconsin (two counties). The first con­ firmed establishment of U. quadrifasciata in Indiana and Michigan is also reported.

Spotted knapweed (Centaurea maculosa Lam.) and diffuse knapweed (C. diffusa Lam.) (Asteraceae) are plants native to Eurasia that have become widespread weeds in North America. Spotted knapweed occurs across south­ ern Canada, the northern United States, and throughout most ofthe western US, while diffuse knapweed occurs primarily in the western US and south­ western Canada (United States Department of Agriculture 1971). Knap­ weeds are adapted to a range ofhabitats and soil types, but appear especially well-suited to relatively dry sites (Watson & RBnney 1974). They are aggres­ sive competitors that invade non-cultivated areas and displace native plants and forage grasses (Gardner 1990, Harris & Cranston 1979, Hirsch & Leitch 1996, Watson & Renney 1974). In Montana, more than 2 million acres are in­ fested by spotted, diffuse, and Russian (Centaurea repens L.) knapweeds, causing economic losses that exceed $42 million (US) annually (Hirsch & Leitch 1996). Spotted knapweed is a short-lived perennial, while diffuse knapweed is a biennial species. Both species reproduce by seed. C. maculosa seeds are dis­ persed by shattering ofthe seedhead (Watson & Renney 1974), and C. diffusa seeds are dispersed when above-ground parts of mature plants break off and tumble with the wind (Strang, et al. 1979). Vehicles, animals, and contami­

lUSDA, APHIS, PPQ, Biocontrol Facility, Montana State Univ., Bozeman, MT 59717. 106 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3 nated hay and crop seed aid in long-distance seed dispersal for both species (Mass 1989, Wallander, et al. 1995). Because spotted and diffuse knapweeds depend on seeds for reproduc­ tion, European seedhead-feeding insects have been released as classical bio­ logical control agents in the US and Canada. Among these agents are Urophora affinis Frauenfield and U. quadrifasciata (Meigen) (Diptera: Tephritidae). Adult females ofboth species oviposit under the bracts of devel­ oping knapweed flower heads. Newly-hatched larvae burrow into the head and begin feeding on developing seeds and receptacle tissue. Knapweed plants form hard, lignified galls around U. affinis larvae, and thin, papery galls around U. quadrifasciata larvae CRees & Story 1991, Zwolfer 1970). Each nonparasitized spotted knapweed seedhead produces an average of 12.6 viable seeds (Harris 1980). Urophora affinis and U. quadrifasciata galls re­ duce spotted knapweed seed production by 2.4 and 1.9 seeds, respectively, per seedhead (Harris 1980). Galls also form a metabolic sink and can reduce the number of flower heads produced by diffuse knapweed, but generally not spotted knapweed (Harris 1980). Generally, Urophora spp. do not appear suf­ ficient to reduce spotted or diffuse knapweed density in North America (Muller-Scharer 1993). Other biological control agents that contribute more directly to plant mortality will be required (Muller-Scharer 1993). The first North American releases of U. affinis and U. quadrifasciata were made in British Columbia, Canada, in 1970 (Harris 1980). In the United States, U. affinis was first released in 1973 in Montana and Oregon, with later releases in California, Idaho, and Washington (Maddox 1979). By 1978, U. affinis was released in the eastern US and is now established in New York, Pennsylvania, and Virginia (Wheeler & Stoops 1996). Urophora quadrifasciata was not released in the western United States but immi­ grated from Canadian populations, with establishment reported in Idaho in 1980 (Gillespie 1983) and Montana in 1981 (Story 1985). Established popula­ tions were later detected in California, Oregon, and Washington. Urophora quadrifasciata was also released in Quebec, Canada in 1979 and in Massa­ chusetts and New York in 1983 (Wheeler 1995, Wheeler & Stoops 1996). It is now established in at least 11 eastern states (Hoebeke 1993, Wheeler 1995). The United States Department of Agriculture, Animal Plant Health In­ spection Service, Plant Protection and Quarantine (hereafter referred to as APHIS) began a national redistribution program for knapweed biological control agents in 1987, with U. affinis and U. quadrifasciata releases begin­ ning in 1987 and 1989, respectively. The purpose ofthis paper is to document releases and subsequent establishment of U. affinis and U. quadrifasciata by the APHIS program.

MATERIALS AND METHODS Potential release sites for biocontrol agents on spotted and diffuse knap­ weed are located and sampled by state cooperators. Prior to release, these knapweed infestations are sampled between October and May with a collec­ tion of 200 randomly selected seedheads. '!\vo seedheads per knapweed plant are collected at intervals of 3-5 feet. The seedheads are placed in a plastic bag and shipped to the Bozeman Biocontrol Facility in Bozeman, Montana. Fifty of the 200 seedheads are randomly selected and examined under a dis­ secting microscope to record the number of U. affinis or U. quadrifasciata galls present. To collect Urophora spp. adults for redistribution, spotted knapweed seedheads were collected in April and May from overwintered plants in 1997 THE GREAT LAKES ENTOMOLOGIST 107

southwestern Montana. Seedheads were placed on large screen trays inside screened field cages approximately 3 x 3 x 2.1 m in size. In Bozeman, U. affi­ nis and U. quadrifasciata adults emerged from seedheads in June and July, and were collected daily from cage walls with an insect vacuum. Groups of 500 flies of both species were placed in 0.95-1 paper containers, to which some excelsior was added as a resting substrate. At least 1000 adults for each release were shipped to cooperators for release within knapweed infes­ tations. The release point was marked with a permanent stake. At each release site, Urophora spp. establishment was monitored by col­ lecting knapweed seedheads between November and May. Two seedheads were collected from randomly selected plants, beginning at the release point and proceeding in expanding circles, until 200 seedheads were collected. Seedheads were sent to the Bozeman laboratory, where 50 of the 200 col­ lected heads were randomly selected and examined under a dissecting micro­ scope. The number of U. affinis and U. quadrifasciata galls observed in each seedhead was recorded. Galls are readily separated, as U. affinis galls are hard and lignified and formed from receptacle tissue (Zwi:ilfer 1970), while U. quadrifasciata galls are thin and papery and formed in the ovaries CRees & Story 1991). One or both Urophora spp. were considered established at a site when galls were present for at least two years after the original release or the population had infested 10% of the knapweed seedheads. Urophora spp. populations were considered collectable when the average gall density ex­ ceeded 1.5 galls per seedhead.

RESULTS AND DISCUSSION The status of Urophora affinis and U. quadrifasciata populations re­ leased in 14 states by APHIS and cooperating personnel is summarized in Table 1. Urophora affinis has been released in 89 counties in 13 states. Of the 85 counties in which post-release sampling was conducted, U. affinis is established in 74 (87%). Urophora quadrifasciata has been released in 46 counties in 12 states, and established populations are present in 43 of the 46 counties (94%). In addition, established U. quadrifasciata populations were present in three counties in Michigan and one county in Indiana before this species was released in these states (Table 1). Both species are established in 32 of the 46 counties (70%) in which they have been released together. These records describe the first reported establishment of U. affinis pop­ ulations in the following states and counties: Arizona (Coconino), Colorado (Boulder, Douglas, LaPlata, Larimer, and Montrose), Michigan (Isabella), Minnesota (Becker, Beltrami, Clearwater, Ottertail, Polk, and Washington), Nevada (White Pine), North Dakota (Kidder), South Dakota (Pennington and Todd), Utah (Wasatch and Weber), and Wisconsin (Washburn and Waukesha) (Table 1 and Table 2). Urophora quadrifasciata was recovered for the first time in Arizona (Coconino), Colorado (Araphaoe, Boulder, Douglas, Jefferson, Larimer, and Montrose), Indiana (Elkhart), Michigan (Delta, Isabella, Dick­ inson, and Menominee), Minnesota (Becker, Beltrami, Clearwater, Otter Tail, Polk, and Washington), Nebraska (Antelope, Holt, Pierce, and Rock), Nevada (White Pine), North Dakota (Kidder), South Dakota (Pennington, Shannon, Todd, and Tripp), Utah (Wasatch and Weber), and Wisconsin (Washburn and Waukesha) (Table 1 and Table 2). Several U. quadrifasciata recoveries from Indiana (Elkhart), Michigan (Delta, Dickson, and Menominee), Minnesota (Washington), and Wyoming (Albany, Bighorn, Carbon, Crook, Laramie, and Sheridan) were detected in prerelease samples (Table 1 and Table 2). Urophora quadrifasciata adults disperse more rapidly than U. affinis 108 THE GREAT lAKES ENTOMOLOGIST Vol. 30, No.3

Table 1. Status of Urophora affinis and U. quadrifasciata populations in the western and midwestern United States. Abbreviation Key: E = Established; NE= Not Estab­ lished; NR= No Release; R = Recovered; ?= Unknown. U. affinis U. quadrifasciata State Yr. rel. Status Yr. recov. Yr. reI. Status Yr. recov. AZ Coconino 1992-1994 Gila 1994-1995 R 1996 1994-1995 NE CO Arapahoe 1990 NE 1990 E 1992 Boulder 1990-1992 E 1992 1990-1992 E 1992 Douglas 1989-1993 E 1992 1989-1990 E 1992 EI Paso 1990 NE 1990 NE Jefferson 1992 NE 1992 E 1993 La Plata 1992 E 1993 1992 R 1997 Larimer 1990 E 1992 1990 E 1992 Montrose 1993 E 1995 1993 E 1995 ID Adams NR E 1987 NR E 1987 Benewah 1987-1989 E 1987 NR E 1987 Bingham 1987-1989 ? NR E 1987 Blaine 1987-1989 ? NR E 1987 Boise 1987-1989 E NR E 1987 Bonner 1987 E 1987 NR E 1987 Bonnville 1987-1988 ? NR E 1987 Butte 1989 E 1987 NR E 1987 Camas NR E 1987 NR E 1987 Cassia 1988-1989 ? NR E 1987 Clark 1987-1989 ? NR E 1987 Clearwater 1987 ? NR E 1987 Custer 1987-1989 ? NR E 1987 Elmore 1988 ? NR E 1987 Idaho 1987 ? NR E 1987 Jefferson 1987-1989 ? NR E 1987 Kootenai 1987 E 1987 NR E 1987 Latah 1987 ? NR E 1987 Lemhi 1987 E 1987 NR E 1987 Lincoln 1987 E 1987 NR E 1987 Madison 1989 ? NR E 1987 01ez Perce 1987 ? NR E 1987 Power 1989 ? NR E 1987 Shoshone 1987-1989 E 1987 NR E 1987 Twin Falls 1987-1989 ? NR E 1987 IN Elkhart 1997 NE 1997 E 1994 MI Isabella 1994 E 1995 1994 E 1995 Delta NR NE NR E 1995 Dickinson NR NE NR E 1995 Menominee NR NE NR E 1994 MN Anoka 01R NE NR E 1993 Becker 1990-1993 E 1991 1990-1993 E 1991 Beltrami 1992 E 1993 1992 E 1995 Chisago NR E 01R E 1993 Clearwater 1991-1993 E 1992 1991-1993 E 1992 Otter Tail 1991-1993 E 1992 1991-1993 E 1992 Polk 1992-1993 E 1993 1992-1993 E 1993 Washington 1990 E 1991 1990 E 1989 MT BigHorn 1990 E 1988 1990 E 1988 Broadwater 1987-1989 E 1994 NR E 1994 Carbon 1988-1991 E 1988 1991 E 1988 Deer Lodge 1987 E 1987 NR E 1987 Fergus 1987-1990 ? 1990 E 1990 1997 THE GREAT lAKES ENTOMOLOGIST 109

Table 1. Continued. U. affinis U. quadrifasciata State Yr. reL Status Yr. recov. Yr. reL Status Yr. recov. 1987-1989 1987-1990 E 1987 1990 E 1987 Glacier 1987-1989 E 1988 NR E 1988 Granite 1987 E 1991 NR E 1988 Hill 1989 ? NR E 1988 Jefferson 1989 E 1991 NR E 1998 Lake 1987-1989 ? NR E 1988 Lewis & Clark1987-1990 E 1988 1990 E 1988 Liberty 1989 ? NR E 1988 Madison 1989 ? NR E 1988 Missoula 1987-1989 E 1987 NR E 1987 Musselshell 1989 ? NR E 1988 Park 1987-1992 E 1991 1992 E 1991 Pondera 1987-1989 E 1991 NR E 1988 Powder River 1988 E 1988 NR E 1988 Powell 1987-1989 E 1991 NR E 1988 Ravalli 1989 ? NR E 1988 Rosebud 1988-1989 E 1988 NR E 1988 Saunders NR E 1988 NR E 1988 Silver Bow 1987 ? NR E 1988 Sweet Grass1989-1992 E 1988 NR E 1988 Teton 1987-1989 E 1988 NR E 1988 Toole 1989 ? NR E 1988 Wheatland 1992 E 1991 1992 E 1991 NE Antelope 1990 R 1991 1990 E 1991 Holt 1991-1992 E 1991 1991-1992 E 1990 Knox NR E 1992 NR R 1991 Pierce 1994 R 1991 1994 E 1991 Rock 1990 R 1991 1990 E 1991 NY Washoe 1993 NE 1993 NE White Pine 1994-1995 E 1996 1994-1995 E 1996 ND Kidder 1990 R 1991 1990 R 1991 SD Davis NR NE NR E 1996 Pennington 1989-1992 E 1991 1989-1992 E 1991 Shannon 1994 NE 1994 E 1995 Todd 1991-1992 E 1993 1991-1992 E 1992 Tripp 1989 NE 1991 1989 E 1991 UT Grand 1991 NE 1991 NE Wasatch 1993-1994 E 1995 1993-1994 E 1992 Weber 1992-1994 E 1992 1992-1994 E 1992 WI St. Croix NR NE NR R 1992 Washburn 1991 E 1993 1991 E 1992 Waukesha 1991 E 1992 1991 E 1993 WY Albany NR NE NR NR 1994 BigHorn NR NE NR NR 1994 Carbon NR NE lIi'R NR 1994 Crook NR NE NR E 1994 Johnson 1993-1994 E 1994 1993-1994 E 1994 Laramie NR NE NR NR 1994 Lincoln 1995 E 1995 1995 E 1995 Natrona 1991-1992 E 1991 1991-1992 E 1991 Saunders NR NE NR E 1994 Teton 1990 E 1995 1990 E 1995 Uinta 1995 NE 1995 E 1995 Wheatland NR E 1991 NR E 1991 110 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

Table 2. Urophora affinis and U. quadrifasciata infestation rates of spotted and diffuse knapweed seedheads. Percent infestation by year State County and insect 1991 1992 1993 1994 1995 1996 1997 AZ Coconino U. affini 2 2 U. quadrifasciata 4 2 Gila U. affinis 6 U. quadrifasciata -0­ CO Arapahoe U. af{tnis -0­ U. quadrifasciata 4 Boulder U. affinis 26 U. quadrifasciata 6 Douglas U. affinis 44 24 U. quadrifasciata 42 68 EI Paso U. affinis -0­ U. quadrifasciata -0­ Jefferson U. affinis -0­ U. quadrifasciata 48 LaPlata U. affinis 6 2 18 U. quadrifasciata -0­ 2 -0­ Larimer U. affinis 48 U. quadrifasciata 20 Montrose U. affinis 4 34 U. quadrifasciata 6 4 IN Elkart U. affinis -0­ -0­ -0­ U. quadrifasciata 2 10 48 MI Isabella U. affinis 14 80 U. quadrifasciata 2 6 Delta U. affinis -0­ U. quadrifasciata 10 Dickinson U. affinis -0­ U. quadrifasciata 30 Menominee U. affinis -0­ -0­ U. quadrifasciata 36 MN Becker U. af{tnis 26 8 32 26 4 U. quadrifasciata 32 -0­ 8 10 34 Beltrami U. affinis 58 42 68 68 U. quadrifasciata -0­ -0­ 8 4 Clearwater U. affinis 20 32 38 34 68 1997 THE GREAT LAKES ENTOMOLOGIST 111

Table 2. Continued Percent infestation by year State County and insect 1991 1992 1993 1994 1995 1996 1997 U. -0­ -0­ 6 2 12 Otter U. affinis 12 42 28 54 60 U. quadrifasciata 2 22 64 48 30 Polk U. affinis 10 6 28 58 U. quadrifasciata 2 8 4 14 Washington U. affinis 72 74 84 68 68 U. quadrifasciata 10 38 20 22 48 NE Antelope U. affinis -0­ 4 -0­ U. quadrifasciata 84 34 58 Holt U. affinis 6 -0­ 6 U. quadrifasciata 30 60 66 Knox U. affinis 2 U. quadrifasciata 32 Pierce U. affinis -0­ -0­ U. quadrifasciata 52 46 Rock U. affinis 2 -0­ U. quadrifasciata 2 26 NV White Pine U. af{inis 26 U. quadrifasciata -0­ ND Kidder U. affinis 6 -0­ U. quadrifasciata -0­ 2 SD Pennington U. affinis 6 34 66 U. quadrifasciata 10 50 10 Shannon U. affinis -0­ -0­ U. quadrifasciata 54 34 Todd U. af{inis -0­ -0­ 4 U. quadrifasciata 22 32 78 Tripp U. affinis -0­ -0­ U. quadrifasciata 34 46 UT Wasatch U. affinis -0­ -0­ 6 8 U. quadrifasciata -0­ 4 60 78 Weber U. af{inis 12 6 4 28 U. quadrifasciata 30 18 44 16 WI St. Croix U. af{inis -0­ U. quadrifasciata 2 112 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

Table 2. Continued Percent infestation by year State and insect 1991 1992 1993 1994 1995

U. affinis -0­ 26 4 2 -0­ U. quadrifasciata 4 4 18 30 16 Waukesha U. affinis 2 10 60 U. quadrifasciata -0­ 30 26 WY Albany U. affinis -0­ U. quadrifasciata 10 Bighorn U. affinis -0­ U. quadrifasciata 6 Carbon U. affinis -0­ U. quadrifasciata 2 Crook U. affinis -0­ U. quadrifasciata 58 Johnson U. af{inis -0­ 6 U. quadrifasciata 52 50 Laramie U. af{inis -0­ U. quadrifasciata 12 Lincoln U. af{inis 90 U. quadrifasciata 12 Natrona U. affinis 6 30 U. quadrifasciata 32 8 Sheridan U. affinis -0­ U. quadrifasciata 78 Teton U. affinis 20 U. quadrifasciata 2 Uinta U. affinis -0­ U. quadrifasciata 18

adults, and quickly migrate into new knapweed-infested areas (Harris 1980, Roitberg 1988), This is supported by the detection of U, quadrifasciata in prerelease samples from Indiana, Michigan, Minnesota, and Wyoming as mentioned above. These recoveries of U. affinis and U, quadrifasciata help to complete the known distribution of these two Urophora flies in the United States. The APHIS distribution program has extended the documented range of U. affi­ nis and U. quadrifasciata into knapweed-infested areas of the western and midwestern US, Urophora quadrifasciata will probably continue to spread on 1997 THE GREAT LAKES ENTOMOLOGIST 113 its own throughout these regions. Urophora affinis does not disperse as read­ ily, and will require collection and distribution from established populations to facilitate spread.

LITERATURE CITED Gardner, D. E. 1990. Role of biological control as a management tool in national parks and other natural areas. U.S. Dept. of Interior, Nat!. Park Service Tech. Rep. NPS&'"'Rm:u::-.."'RTR 90. 42 p. Gillespie, R. L. 1983. Bionomics of Urophora affinis Frfld. and U. quadrifasciata Meigen (Diptera: Tephritidae) in northern Idaho. MS Thesis, University of Idaho, Moscow, ID. 90 p. Harris, P. 1980. Establishment of Urophora affinis Frfld. and U. quadrifasciata (Meig.) (Diptera: Tephritidae) in Canada for the biological control of diffuse and spotted knapweed. Z. ang. Entomo!. 89::504-515. Harris, P. & R. Cranston. 1979. An economic evaluation of control methods for diffuse and spotted knapweed in western Canada. Can. J. Plant Sci. 59:375-382. Hirsch, S. A. and J. A. Leitch. 1996. The impact of knapweed on Montana's economy. Xorth Dakota State Vniv., Agric. Exp. Sta., Fargo, Agric. Econ. Rept. 355. 43 pp. Hoebeke, E. R. 1993. Establishment of Urophora quadrifasciata (Diptera: Tephritidae) and Chrysolina quadtigemina (Coleoptera: Chrysomelidae) in portions of eastern Lnited States. Entomo!. News 104:143-152. Maddox, D. :lo1. 1979. The Knapweeds: their economics and biological control in the western states, L.S.A. Rangelands 1:139-14l. :'>1ass, F. H. 1989. How to contain knapweed in far western Montana. Pp. 113-117 In: P. K. Fay, and J. R. Lacey, eds. Proc., Knapweed Symposium. Montana State Univ., Bozeman. Muller-Scharer, H. 1993. The biological control of Centaurea spp. in North America: do insects solve the problem? Pestic. Sci. 37:343-353. Rees, X. E. & J. :lot Story. 1991. Host plant testing of Urophora quadlifasciata (Diptera: Tephritidae) against Carthamus tinctorius and two North American species ofCentuurea. Entomophaga 36:115-119. Roitberg, B. D. 1988. Comparative flight dynamics of knapweed gall flies Urophora quadrifasciata and [I. uffinis (Diptera: Tephritidae). J. Entomo!. Soc. B. C. 85:58-64. Story, J . .:'>1. 1985. First report of the dispersal into Montana of Urophora quadrifasci­ ata (Diptera: Tephritidae), a fly released in Canada for biological control of spotted and diffuse knapweed. Can. Entomo!. 117:1061-1062. United States Department of Agriculture. 1971. Common weeds of the United States. Dover Publications Inc., Xew York. 383 p. Wallander, R. T., B. E. Olson, & J. Lacey. 1995. Spotted knapweed seed viability after passing through sheep and mule deer. J. Range Manage. 48:145-149. Watson, A. K. & A J. Renney. 1974. The Biology of Canadian weeds. 6. Centaurea dif­ fuse and C maculosa. Can. J. Plant Sci. 54:687-701. Wheeler, A. G. 1995. Urophora quadrifasciata (Diptera: Tephritidae introduced seed­ head fly new to Midwestern North America. Great Lakes Entomo!. 28:235-236. Wheeler, A. G. Jr., & C. A. Stoops. 1996. Establishment of Urophora affinis on spotted knapweed in Pennsylvania within new Eastern U. S. records of U. quadrifasciata (Diptera: 'Thphritidae). Proc. Entomo!. Soc. Wash. 98:93-99. ZwOlfer, H. 1970. Investigations on the host-specificity of Urophora affinis Frfld. (Dipt., Trypetidae). Commw. Inst. of Bio!. Contr. Delmont, Switzerland. Progress Re­ port No. 25. 28 p. 1997 THE GREAT LAKES ENTOMOLOGIST 115

INTRODUCED PURPLE LOOSESTRIFE AS HOST OF NATIVE SATURNIIDAE (LEPIDOPTERA)

James G. Barbourl and Erik Kiviat2

ABSTRACT Purple loosestrife (Lythrum salicaria,Lythraceae) arrived in North America nearly 200 years ago. In 1969 we first found larvae of the native Ce­ cropia (Hyalophora cecropia) and Polyphemus (Antheraea polyphemus) moths (Lepidoptera: Saturniidae) on loosestrife in the Hudson River Valley, New York, and we have since found 10 (Automeris io) on this plant. A census of 4th and 5th instar saturniids in four 0.25 ha plots in purple loosestrife­ gray dogwood (Comus racemosa) wet meadows near Saugerties in 1984 indi­ cated that Polyphemus and Cecropia larvae occurred much more frequently on loosestrife than on dogwood, a native host. The switch from native woody hosts to an introduced herb may have been facilitated by the dense shrub­ like habit, high productivity, and high tannin content ofloosestrife.

The ecological relationships between native species and introduced species are of theoretical and practical interest (Mooney and Drake 1986). Complex interactions of insects with native and introduced hosts may affect population and range dynamics of biota, pollination, herbivore impacts on plant community composition, decomposition and nutrient cycling, agricul­ ture, and the potential for biological control of weeds. Purple loosestrife, Lythrum salicaria L. (Lythraceae), was introduced from Europe to the northeastern United States in the early 1800s (Thompson et al. 1987). Loosestrife is a broad-leaved, cespitose, perennial herb 1-3 m tall with a woody root system and herbaceous aerial stems that die but per­ sist erect through winter. In older plants the root crown may become an ele­ vated pedestal greater than 30 cm in diameter and 30 cm high that supports 25-50+ stout stems. Loosestrife leaves are sessile, 3-12 cm long and 1-2 cm wide. Loosestrife is abundant in ditches, shores, wet meadows, nontidal marshes, low-salinity tidal marshes, and disturbed upland soils in many re­ gions of the northern United States and southern Canada. Many animals eat loosestrife leaves, but rarely are more than a few plants defoliated at a site (Hight 1990; Barbour and Kiviat, personal observations). Many Saturniidae (silk moths) are known for the large size and bold color patterns of the adults and larvae. Antheraea polyphemus (Cram.) (Polyphemus), Hyalophora cecropia (L.) (Cecropia), and Automeris io (F.) (10), have broad host and habitat niches, and large geographic ranges, in eastern

15 Fish Creek Road, Saugerties, NY 12477. 2Corresponding author; Hudsonia Ltd., Bard College Field Station, Annandale, i\'Y 12504-0217. 116 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

North America (Tuskes et al. 1996). Here we report purple loosestrife as a larval host for these native Saturniidae (Lepidoptera) in New York.

MATERIALS AND METHODS Our general study area in northeastern Ulster and western Dutchess counties has elevations of 0-150 m adjoining the Hudson River midway be­ tween New York City and Albany, New York. Annual precipitation is ca. 1000 mm. Woody vegetation, mostly post-agricultural, covers about half the land­ scape, and wetlands cover perhaps 5-10%. Barbour conducted censuses 18-21 August 1984 in shrubby wet fields 1.5 and 4 km north of Saugerties (Ulster County), elevation 45 m, where he had found the highest local densities of saturniid larvae the previous two years. These sites are 1.~2.0 km from the tidal Hudson River on deep, nearly level, somewhat poorly drained to poorly drained soils on glaciolacustrine silty clay and glacial outwash (Tornes 1979). At each of the two sites, two 50 x 50 m (0.25 ha) plots, at least 50 m from forest edges, were selected as representa­ tive. Each plot bordered a drainage ditch or mowed right-of-way, and was di­ vided into 25 subplots (10 x 10 m). Barbour searched each subplot once, recording 4th and 5th instar saturniids, and visually estimating the cover of purple loosestrife and gray dogwood (Comus racemosa Lam.). We believe the census was complete for 5th instar and nearly complete for 4th instar sat­ urniids. There were no trees in the plots, and few other plants except aster (Aster) and goldenrod (Solidago). The plots rarely flood and did not flood in 1983-84. We computed Spearman's rank correlations (rho), Kruskal-Wallis one­ way analysis of variance by ranks, Wilcoxon matched-pairs tests (T), and two-tailed Fisher exact tests with Statistica version 5.1 (StatSoft, Tulsa, OK).

RESULTS In 1969 Barbour found 2 Cecropia and 1 Polyphemus (all 4th instar) on purple loosestrife near Kingston, New York. In 1976, Kiviat found a copulat­ ing pair, eggs, and larvae (instars 1-3) on purple loosestrife in the towns of Clinton and Wappinger, Dutchess County. In 1979 Barbour found several Ce­ cropia cocoons in loosestrife in a highway intersection "island" where there were no potential woody hosts. In September 1980 he found 45 5th instar Ce­ cropia feeding on purple loosestrife in a highway intersection island at Kingston, and 63 on loosestrife near Stone Ridge (Ulster County), In the early 1980s he found more Polyphemus larvae on loosestrife in Ulster County. In 1991, Barbour found 3 Cecropia larvae on loosestrife at West Point Military Academy (Orange County, New York), and in 1992 he found Cecropia and Polyphemus larvae on loosestrife in the Bethlehem train yards (Albany County, New York). We have seen 1st instars of both moths feeding on loosestrife. Since 1984 there has been no obvious change in saturniid use of loosestrife in the Hudson Valley. In the 1980s, Robert Dirig (Cornell Uni­ versity, pers. communication 1997) found a Cecropia cocoon on loosestrife in Ithaca (Tompkins County, central New York), suggesting use of this plant for food elsewhere in the state. On the 4 census plots combined (total 1 hal there was a total of 79 4th and 5th instar saturniids, comprising 50 Polyphemus, 27 Cecropia, and 2 10. All larvae were on loosestrife except 4 Polyphemus on gray dogwood. Polyphemus was significantly more abundant than Cecropia (Wilcoxon 1997 THE GREAT lAKES ENTOMOLOGIST 117

Table 1. Probabilities ofSpearman rank correlations (rho) for saturniid larvae and veg­ etation cover for n 100 subplots (each 10 x 10 m) at four sites; '" indicates a negative correlation. The number of plots (50 x 50 m, n = 25) with rho significant (p < 0.05) fol­ lows the probability. Polyphemus Cecropia 'Ibtal saturniids Loosestrife Cecropia 0.00011 2 Loosestrife cover 0.03 2 0.26 o 0.06 2 Dogwood cover * 0.038 1 * 0.024 2 * 0.0045 1 * 0.0017 4 Ls. + dw. cover 0.56 '" 0.69 * 0.87

T 117, P = 0.01). ANOVAs indicated that numbers of total saturniids and Polyphemus differed significantly among the 4 plots (p =0.0039, 0.012, re­ spectively), but Cecropia did not differ (p > 0.05). Loosestrife cover and dog­ wood cover were negatively correlated (Table 1). Loosestrife cover was 0-1.00 (mean 0.317, median 0.25) and dogwood cover 0-0.70 (mean 0.128, median 0.10). Loosestrife cover was significantly greater than dogwood cover (Wilcoxon T =783.5, P =0.00001). Polyphemus was correlated with Cecropia and with loosestrife cover, and Polyphemus and Cecropia were each negatively correlated with dogwood cover (Table 1). Total saturniids were negatively correlated with dogwood cover but were not correlated with loosestrife cover (Table 1, Fig. 1). Saturni­ ids separately or combined, however, were not correlated with total ''brushy'' cover (Le. dogwood + loosestrife). The August censuses were in the early part of the 5th instar period for Polyphemus, and between the two peaks for late instar Cecropia (adult Ce­ cropia exhibit bimodal emergence and oviposition [Waldbauer and Sternburg 1973]). On 5 September 1984, Barbour re-censused 1 plot. There were 3 Polyphemus, 9 Cecropia, and 3 10 (compared to 5 Polyphemus, 2 Cecropia, and 0 10 on 21 August). The numbers of Polyphemus and Cecropia were not significantly different between the two dates (Fisher p = 0.074). On 2 Sep­ tember 1984, Barbour censused 4th and 5th instar saturniids along 835 m of wet drainage ditches with loosestrife and other plants (habitat width 1-2.5 m, habitat area ca. 0.15 hal in a hay field near one plot. There were 3 Polyphemus, 27 Cecropia, and 2 10, all on loosestrife. Saturniid density was equivalent to 213 ha-1 (Polyphemus 20 ha-1, and Cecropia 180 ha-1).

DISCUSSION Cecropia and Polyphemus are associated with savanna-like habitats (scattered trees and shrubs), forest ecotones such as riparian and lacustrine margins, open shrubby wetlands (Stratton-Porter 1910), old fields and burned forests (Waldbauer 1996:90, 257), and barrier beach shrublands (John Cryan, New York, NY, pers. communication 1988). Polyphemus also oc­ curs in deciduous forests. Both moths readily colonize disturbed habitats and may abound in suburbs and cities (Scarbrough 1970, Waldbauer 1996), post­ industrial shrublands, and railroad rights-of-way. We have found saturniids feeding on loosestrife in wet meadows, pond shores, ditches, and wetland fill. Kiviat has found Cecropia cocoons on loosestrife distant from woody plants in a freshwater-tidal marsh of the Hudson River. Loosestrife stands supporting 118 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

5

4

~~ ~~ Figure 1. Occurrence of 4th and 5th instar saturniids (Antheraea polyphemus, Hyalophora cecropia, and Automeris io combined) in 10 x 10 m subplots (n = 100) in relation to cover of purple loosestrife and gray dogwood.

saturniid larvae comprise denser, larger plants more often than sparser, smaller plants. Figure 2 shows a Cecropia larva on loosestrife. Purple loosestrife is probably as abundant in the Hudson Valley as any­ where in North America. Other regions with extensive loosestrife include central and western New York, the St. Lawrence River corridor in Quebec, the Lake Erie shore of Ohio, areas of Michigan and Wisconsin, and north­ eastern Massachusetts (Thompson et al. 1987). Cecropia larvae were found on purple loosestrife in Orange County, New York in 1986-87 (Hight 1990). A population of the Hemileuca maia (Drury) complex in Wisconsin (Tuskes et al. 1996:121) and Saturnia pavonia (Linnaeus) in Europe (Stone 1991) are the only other saturniids reported to feed on loosestrife to our knowledge. Loosestrife thrives in disturbed, moist or wet soils, and often forms thick­ ets adjoining or intermingled with native hosts of Polyphemus and Cecropia such as shrubby dogwoods (Comus spp.) and willows (Salix spp.). On our study plots, Polyphemus and Cecropia occurred in areas with low dogwood cover and moderate to high loosestrife cover (Fig. 1). Larvae occurred almost exclusively on loosestrife, in close proximity to dogwood. Our qualitative ob­ servations also indicate that Polyphemus and Cecropia are at least as com­ mon on loosestrife as on their native woody hosts. 1997 THE GREAT LAKES ENTOMOLOGIST 119

Figure 2. Hyalophora cecropia larva on purple loosestrife, Ulster County, New York. Photo by Anita F. Barbour.

Polyphemus and Cecropia share many woody hosts, especially shrubby dogwoods, willows, maples (Acer spp.), and cherries (Prunus spp.) (Tuskes et al. 1996; Barbour, personal observations). Besides purple loosestrife, only 3 herbs have been reported as Cecropia hosts: Decodon verticillatus (L.) Ell. (Lythraceae), a native species similar to and in the same family as purple loosestrife, native hops (Humulus lupulus L., Moraceae), and introduced gar­ den peony (Paeonia officinalis L., Ranunculaceae) (Eliot and Soule 1902, Stone 1991, Waldbauer 1996:89). The Decodon record was based on cocoons over water (Eliot and Soule 1902:251), suggesting oviposition on this plant. Only 1 herb has been reported as a Polyphemus host, wild-indigo (Baptisia tinctoria [L.] Vent., Fabaceae) (Stone 1991). The host niche breadth of Saturniidae is indicated by lists of larval food plants in Stone (1991) which we analyzed for nominate subspecies of moths only. These lists are a useful index although Stone (1991) presumably in­ 120 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

eluded laboratory as well as wild host records, and some wild host records are probably based on cocoon locations (e.g. mature Cecropia larvae may move up to 10 m from host plant to a spinning site on another species [Wald­ bauer and Sternburg 1967aD. Larval food plants listed by Stone (1991) for Polyphemus, Cecropia, and 10 comprise 40, 49, and 56 genera, respectively (arbitrarily combining Pyrus, Malus, and Sorbus). For 9 native species of Saturniidae now found in the Hudson Valley but that we have not found on purple loosestrife, Stone (1991) listed 4-31 (median == 6) genera oflarval food plants. For Polyphemus, Cecropia, and 10, 1, 2, and ca. 9 genera of herbs are among the food plants listed, whereas only 0-1 (median 0) herb genera were listed for the other 9 saturniids. (For two additional saturniids, the in­ troduced ailanthus silk moth [Samia cynthia Drury] and the historically pre­ sent imperial moth [Eacles imperialis Drury), Stone (1991) listed 41 and 42 genera of food plants, respectively.) These figures suggest that, of the sat­ urniids in long contact with loosestrife in the Hudson Valley, the species with the broadest host niches and the most herbaceous food plants are the moths that have accepted loosestrife as host. It is also significant that the saturniid species using loosestrife pupate aboveground, as the soil is often wet under loosestrife. In the case of Cecropia, several factors may explain the successful switch to purple loosestrife: 1. Cecropia is a generalist that accepts many hosts; 2. Loosestrife is taller than most herbs and the same height as gray dogwood, and thus may intercept the flight path of ovipositing moths; 3. Loosestrife is very productive, and the dense, rapidly re-growing foliage of mature plants provides abundant food and concealment for large larvae (see Waldbauer 1996:90); 4. There is little competition for loosestrife leaves from other herbi­ vores; 5. Loosestrife leaves appear to have a higher moisture content than leaves of common woody hosts e.g. Comus, possibly increasing the availabil­ ity of nutrients relative to woody plants (see Scriber 1975); 6. Cecropia co­ coons among low dense shrnb stems or basal tree shoots are more likely to escape bird predation (Waldbauer and Sternburg 1967b), and the stiff, crowded, winter-persistent loosestrife stems offer sturdy attachment and bet­ ter concealment for the large cocoons than most trees and shrubs; 7. Sat­ urniid larvae in general prefer tannin-rich leaves (Bernays and ,Janzen 1988), and although the leaves of most woody dicots contain tannins and herbaceous dicots do not (Bate-Smith and Metcalfe 1957, Swain 1979, Rhoades 1983), purple loosestrife leaves have high tannin levels (Vincent and Segonzac 1954, Gibbs 1974, Shishkin and Bobrov 1974). Except for factor 6, these factors may also apply to Polyphemus and 10. We hypothesize that: 1. The switching of Polyphemus and Cecropia from native woody hosts to loosestrife was initially possible because of the wide host range of the moths, the abundance of dense mature loosestrife in associ­ ation with traditional hosts in habitats that rarely flood, and the attraction of the moths to tannin-rich leaves; and 2. The switch was successful because abundant, moisture-rich foliage allows higher growth rates; also larvae feed­ ing in dense loosestrife foliage and cocoons spun among crowded loosestrife stems are more likely to escape predators than larvae and cocoons on woody plants. Recent, large-scale switches from native to introduced leguminous herbs have been documented in the butterflies Erynnis baptisiae Forbes (Shapiro 1979) and Glaucopsyche lygdamus (Doubleday), ssp. couperi Grote (Dirig and Cryan 1992). (Vicia cracca L., the new host of G. l. couperi, may be native in New York, nonetheless this butterfly has two alien legume hosts in Canada [Robert Dirig, pers. communication 1997]). Purple loosestrife is considered a pest in North America because it alters the marsh and wet meadow habitats of stenotopic species (Thompson et al. 1997 THE GREAT lAKES ENTOMOLOGIST 121

1987), including plants, graminoid-feeding insects, graminoid-nesting marsh birds, and muskrat. European beetles have been released in the northern U.S. during the 1990s to control loosestrife (Malecki et al. 1993). Polyphe­ mus, Cecropia, and 10 moths are generalized herbivores that may switch from native woody hosts to purple loosestrife in regions other than the Hud­ son Valley as loosestrife becomes more abundant, and perhaps switch back to woody hosts where biological control causes loosestrife populations to de­ cline. Other saturniids with broad food plant niches, such as ailanthus silk moth and imperial moth, might also switch to loosestrife. Monitoring these interactions would add to knowledge ofloosestrife ecology in North America, and the ecology of invasions in general. The switching of native insects to an introduced host plant could indicate incipient "natural" control of purple loosestrife (perhaps akin to the control of Eurasian watermilfoil [Myriophyl­ lum spicatum L.J by native insects, see Creed and Sheldon [1995]). We think it unlikely, however, that saturniid larvae can reach abundances sufficient to reduce Hudson Valley loosestrife populations.

ACKNOWLEDGMENTS Anita F. Barbour participated in field work. John Bleiler and Stephen Hight provided literature citations. Robert Dirig and Gil Waldbauer com­ mented on drafts. This is Bard College Field Station-Hudsonia Contribu­ tion 70.

LITERATURE CITED Bate-Smith, E. C. and C. R. Metcalfe. 1957. Leuco-anthocyanins. 3. The nature and systematic distribution of tannins in dicotyledonous plants. Jour. of the Linn. Soc. of London; Botany 55:669-705. Bernays, E. A and D. H. Janzen. 1988. Saturniid and sphingid caterpillars: two ways to eat leaves. Ecology 69(4):1153-1160. Creed, R. P., Jr. and S. P. Sheldon. 1995. Weevils and watermilfoil: did a North Ameri­ can herbivore cause the decline of an exotic plant? Ecol. Appl. 5(4):1113-1121. Dirig, R. and J. F. Cryan. 1992. The status of silvery blue subspecies (Glaucopsyche lygdamus lygdamus and G. l. couperi) in New York. Jour. of the Lepidopt. Soc. 45:272-290. Eliot, 1. M. and C. G. Soule. 1902. Caterpillars and their moths. Century Co., New York. Gibbs, R. D. 1974. Chemotaxonomy of flowering plants. McGill-Queen's UniVersity Press, Montreal. 4 volumes. Hight, S. D. 1990. Available feeding niches in populations of Lythrum salicaria L. (pur­ ple loosestrife) in the northeastern United States. Proceedings of the International Symposium on Biological Control of Weeds 7:269-278. Malecki, R. A, B. Blossey, S. D. Hight, D. Schroeder, L. T. Kok, and J. R. Coulson. 1993. Biological control of purple loosestrife. BioScience 43(10):680-686. Mooney, H. A and J. A Drake, eds. 1986. Ecology of biological invasions of North America and Hawaii. Springer-Verlag, New York. Rhoades, D. F. 1983. Herbivore population dynamics and plant chemistry. pp. 155-222. In: R. F. De=o and M. S. McClure, eds. Variable Plants and Herbivores in Natural and Managed Systems. Academic Press, New York. Scarbrough, A G. 1970. The occurrence of Hyalophora cecropia (L.) as related to ur­ banization. Ph.D. Thesis, University of Illinois, Urbana-Champaign. Scriber, J. M. 1975. Comparative nutritional ecology of herbivorous insects: general­ 122 THE GREAT LAKES ENTOMOLOGIST Vol. 30, No.3

ized and specialized feeding strategies in the Papilionidae and Saturniidae (Lepi­ doptera). Ph.D. Thesis, Cornell University, Ithaca, New York. Shapiro, A. M. 1979. Erynnis baptisiae (Hesperiidae) on crown vetch (Leguminosae). Jour. of the Lepidopt. Soc. 19:227-230. Shishkin, B. K. and E. G. Bobrov, eds. 1974. Flora of the U.S.S.R. Volume 15. Malvales, Parietales, Myrtiflorae. Israel Program for Scientific Translations, Keter Publishing House, Jerusalem. (Originally published in Russian 1949, Akademiya Nauk SSSR, Moskva-Leningrad.) Stone, S. E. 1991. Foodplants of world Saturniidae. Lepidopt. Soc. Mem. 4. Stratton-Porter, G. 1910. Music of the wild, with reproductions of the performers, their instruments and festival halls. Jennings and Graham, Cincinnati, Ohio. Swain, T. 1979. Tannins and lignins. pp. 657-682. In: G. A. Rosenthal and D. H. Janzen, eds. Herbivores; Their Interactions with Secondary Plant Metabolites. Acad­ emic Press, New York. Thompson, D. Q., R. L. Stuckey, and E. B. Thompson. 1987. Spread, impact, and con­ trol of purple loosestrife (Lythrum salicaria) in North American wetlands. U.S. Fish and Wildlife Service, Fish and Wildlife Research 2. Tornes, L. A. 1979. Soil survey of Ulster County, New York. U.S. Department of Agri­ culture, Soil Conservation Service. Tuskes, P. M., J. P. Tuttle, and M. M. Collins. 1996. The wild silk moths of North Amer­ ica. Cornell University Press, Ithaca. Vincent, D. and G. Segonzac. 1954. Quelques donnees nouvelles sur la salicaire Lythrum salicaria L. et son action pharmacologique. Acta Phytotherapeutica 1(6):1-13. Waldbauer, G. P. 1996. Insects through the seasons. Harvard University Press, Cam­ bridge. Waldbauer, G. P. and J. G. Sternburg. 1967a. Host plants and the locations of the baggy and compact cocoons of Hyalophora cecropia (Lepidoptera: Saturniidae). Ann. Entomol. Soc. America 60(1):97-10l. Waldbauer, G. P. and J. G. Sternburg. 1967b. Differential predation on cocoons of Hyalophora cecropia (Lepidoptera: Saturniidae) spun on shrubs and trees. Ecology 48(2):312-315. Waldbauer, G. P. and J. G. Sternburg. 1973. Polymorphic termination of diapause by Cecropia: genetic and geographical aspects. Biological Bull. 145:627-641. r--_

1997 THE GREAT LAKES ENTOMOLOGIST 123

THE ASSASSIN BUG ZELUS LURIDUS (HETEROPTERA: REDUVIIDAE) IN MICHIGAN'S UPPER PENINSULA

Philip A. Cochran, James R. Hodgson and Adam A. Leisten 1

On 17 July 1992, an assassin bug (Zelus luridus Stal) was flushed from the stomach of a smallmouth bass (Micropterus dolomieu) collected in West Long Lake of the University of Notre Dame Environmental Research Center, Gogebic County, Michigan. The bass measured 237 mm in total length and weighed 187 g. Although Z. luridus was reported by McPherson (1992) from 26 of 67 counties in Michigan's Lower Peninsula, he did not list it from the Upper Peninsula. Including Z. luridus, only 5 of 28 reduviids known from Michigan have been collected in the Upper Peninsula, a region that appar­ ently has been under-collected (McPherson 1992). The Z. luridus reported herein probably was eaten by the smallmouth bass after falling from a shoreline bush or tree to the water's surface. Al­ though smallmouth bass typically eat fish and crayfish primarily, their diet in unproductive lakes in northern Michigan may be dominated by insects (Clady 1974). The stomach contents ofthe bass that had eaten the Z. luridus specimen contained, in addition, a mixture of both aquatic and terrestrial insects. Chemical defense systems are characteristic of the Heteroptera, and as­ sassin bugs possess not only scent glands in the metathorax, but also toxic saliva (McGavin 1993). Indeed, Cochran (1990) observed a treefrog reject as­ sassin bugs after taking them into its mouth. However, McGavin (1993) sug­ gested that particular groups of bugs may have different sets of primary predators, with chemical defenses evolved to target the most appropriate en­ emies in each case. It seems likely that the chemical defenses of assassin bugs are more effective against whatever terrestrial predators they typically encounter than against aquatic predators such as bass. In addition, it is our experience that some individual bass appear to specialize on noxious prey (Cochran et aI., unpublished data). Thus, we are not surprised to find a pre­ sumably toxic or distasteful assassin bug among the gut contents of a pre­ sumably occasional predator.

ACKNOWLEDGMENTS We thank J.E. McPherson for reading a preliminary draft of this manu­ script and for identification of the specimen reported herein. It has been placed in the entomology collection at Southern Illinois University. We are grateful to the personnel ofthe University of Notre Dame Environmental Re­ search Center for logistical support. Our fish dietary analyses were funded by the National Science Foundation.

IDivision ofNatural Sciences, St. Norbert College, DePere, WI 54115. '---

124 THE GREAT lAKES ENTOMOLOGIST Vol. 30, No.3

LITERATURE CITED Clady, M.D. 1974. Food habits of yellow perch, smallmouth bass and largemouth bass in two unproductive lakes in northern Michigan. Amer. Midi. Nat. 91:453-459. Cochran, p.A. 1990. On the anti-predatory capability of assassin bugs. Y.E.S. Quarterly 7(4):5-6. McGavin, G.C. 1993. Bugs of the world. Facts on File, Inc. New York, New York. 192 pp. McPherson. J.E. 1992. The assassin bugs of Michigan (Heteroptera: Reduviidae). Great Lakes Entomol. 25 :25-31 . INSTRUCTIONS FOR AUTHORS

SUBJECTS Papers dealing with almost any aspect of entomology be considered for publication in The Great Lakes Entomologist. Appropriate subjects are those to professional and amateur entomologists in the North Central States and Canada, as general papers and revisions di­ rected to a larger audience while retaining interest to our geographic area. All manuscripts are refereed by two except notes, which are reviewed ex­ ternally at the discretion of the Editor. REQUIREMENTS Manuscripts must be typed or printed, double-spaced, with 1» margins on 8 112 x 11" or equiv­ alent size paper, and submitted in triplicate. Authors submitting only one copy will be asked to send two more as the editor will not make additional copies. Please italicize or underline only those words that are to be italicized. Use subheadings sparingly. Footnotes (except for author's addresses, which must be on the title page, and treated as a footnote), legends, and captions of illustrations should be printed on separate sheets of paper. Titles should be concise, identiJYing the order and family discussed. The author of each species must be given fully at least once in the text, but not in the title or abstract. If a common name is used for a species or group, it should be in accordance with the common names published by the Entomological Society of America. The format for references must follow that used in previous issues of The Great Lakes Entomologist. Literature cited is just that - no unpublished manuscripts or internal memos. FIGURES & TABLES Remember that the printed page area for The Great Lakes Entomologist is 4.5 x 7 inches. Scale your illustrations accordingly. Photographs should be glossy finish, and mounted on stiff white card­ board (transparencies are not acceptable). Drawings, charts, graphs, and maps must be scaled to permit proper reduction without loss of detail. Please reduce illustrations or plates to a size no greater than 9 x 12" to permit easier handling. Ifyou have several photographs, it may be better to group them on a plate, rather than several separate photographs. Attach a figure number to the re­ verse side of each figure and include the authors' names. Unsuitably mounted photographs or poor figures will be returned to authors for revision. Figures printed on dot-matrix printers are highly discouraged. Even if your figures are printed on a laser printer, it is recommended that you affix them to a stiff backing. Do not submit photocopied figures for reproduction. A service bureau or local print shop can provide suitably-sized photostats for the original figures. Tables should be kept as uncluttered as possible, and should fit normally across a page when typeset by the printers. Contributors should follow the Council of Biology Editors Style Manual, 5th ed., and examine recent issues of The Great Lakes Entomologist for proper format of manuscripts. MA..."WSCRIPTS ON DISK Manuscripts must be submitted on computer diskette (Macintosh, MS-DOS, Or Apple II format) along with one printed copy, after they have been accepted for publication. The files may be format­ ted in any popularly used word-processing program (Microsoft Word, WordPerfect, ClarisWorks, MacWrite, AppleWorks, WordStar, or WriteNow), or left as a Rich Text Format (RTF) or ASCII file. ~ficrosoft Word for PC or Macintosh is the preferred format. . Special formatting notes for submitting manuscripts on disk: The organizational format for a manuscript is as seen in the recent issues of the journal: TITLE, Author(s), Abstract, Introduction, Methods & Materials, Results, Discussion, Acknowledgments, Literature Cited, Tables, and List of Figures. Do not use extra spaces between paragraphs or references in the Lit. Cited. The columns oftext in tables should be aligned with TABS, not spaces. Some symbols may not translate properly from one computer system to another, such as ii, ll, 0, ii, ii, and male & female symbols. So long as these symbols are clearly seen in the manuscript, adjustments can be made in the copy sent to the printers. Authors should use italics, rather than underline when submitting manuscripts on disk. PAGE CHARGES Papers published in The Great Lakes Entomologist are subject to a page charge of $35.00 per published page. Members of the Society, who are authors without funds from grants, institutions, or industry, and are unable to pay costs from personal funds, may apply to the Society for financial assistance. Application for subsidy must be made at the time a manuscript is initially sub­ mitted for publication_ Authors will receive page proof, together with an order blank for separates. Extensive changes to the proof by the author will be billed at a rate of $1.00 per line. COVER ARTWORK Cover art or photographs are desired for upcoming issues. They are published free of charge. We only require that they be suitably mounted as described above, and that the subject be identi­ fied as accurately as possible. EDITOR'S ADDRESS All manuscripts for The Great Lakes Entomologist should be sent to the Editor, Mark F. O'Brien, Insect Division, Museum of Zoology, The University of Michigan, Ann Arbor, MI, 48109­ 1079, USA. BUSINESS Other correspondence should to the Secretary, Michigan Entomological Society, do Dept. of Entomology, Michigan State UIllvt'T8JlW. East Lansing, MI 48824-1115.