PROC. ENTOMOL. SOC. WASH. 102(1). 2000, pp. 15U161 ISCHNOPTERAPION (CHLORAPION) VIRENS (HERBST) (COLEOPTERA: CURCULIONOIDEA: BRENTIDAE: APIONINAE), A PALEARCTIC CLOVER PEST NEW TO NORTH AMERICA: RECOGNITION FEATURES, DISTRIBUTION, AND BIONOMICS E. Richard Hoebeke, Robert A. Byers, Miguel A. Alonso-Zarazaga, and James Stimmel F. (ERH) Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY 14853, U.S.A. (e-mail: [email protected]); (RAB) Pasture and Watershed Management Research Unit, ARS, U.S. Department of Agriculture, U.S. Regional Pasture Research Laboratory, University Park, PA 16802, U.S.A.; (MAAZ) Depto. de Biodiversidad y Biol- ogia Evolutiva, Museo Nacional de Ciencias Naturales, Jose Gutierrez Abascal, 2. 28006 Madrid, Spain; (JFS) Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, PA 17110, U.S.A. — Abstract. Ischnopterapion (Chlorapion) virens (Herbst) is reported for the first time in North America based on collections in 22 counties in Pennsylvania, 5 counties in New York, 3 counties each in Maryland and New Jersey, and 1 county each in western Con- necticut, northern Delaware, and northern Virginia. This immigrant weevil is a pest of clover {Trifolium spp.), with adults injuring the foliage, and larvae mining in the petioles, stems (stolons), root-crowns, and roots. A detailed redescription and a thorough diagnosis of the adult are given to allow its identification and separation from similar Nearctic species, and its biology and seasonal history in the Palearctic region are summarized from the European literature. Key Words: Ischnopterapion (Chlorapion) virens, immigrant weevil, cloverpest, system- atics, bionomics The first North American collections of subfamily Apioninae' [currently placed in the Palearctic apionine weevil Ischnopter- the Brentidae, see Kuschel (1995) and apion virens (Herbst), a pest ofclover, were Lawrence and Newton (1995)] historically made during an inventory to: (1) determine have been assigned to the exceptionally invertebrate species richness and abundance large and difficult genus Apion, which in- in dairy farm paddocks under different eludes nearly 1,600 species. In America grazing regimes in the northeastern United north of Mexico, more than 150 species of ^P^on have been recorded (O'Brien and States; and, (2) to identify invertebrate spe- cies that may potentially affect sustainabil- Wibmer 1982). In this paper, we recognize ity of pasture systems. Soil sampling for ^he supraspecific taxa proposed by Alonso- this invertebrate survey was conducted by Zarazaga (1990) in which many of the for- RAB and Gary M. Barker (Agresearch, r7ij\ XHTamil-twon, TNVTew Zealand) on '2^1irP*ennsyl1va- ' The classification presented in Kuschel (1995) and -^ L,awrence andj Nxtewton (,,1,9^9,^5^). is f^o.l.lowed here, with nia farms irom iyy4—Ivvo. jj^g Apioninae recognized as a subfamily ofthe Bren- Most of the world's described taxa in the tidae (ERH). 152 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mer subgenera ofApion are elevated to ge- cality, 3-X-1997. Salem Co., Woodstown, NEW neric rank. 8-X-1997. Warren Co., 6-X-1997. Here, we list and map the known distri- YORK: Chemung Co., Lowman, 16-VII- butional records of /. virens in the eastern 1997. Chenango Co., Earlville, 15-VII- United States; provide characters to identify 1997. Orange Co., Westtown, 29-VII-1998. this newly detected immigrant and to allow Tompkins Co., Ithaca, 28-VII-1998. it to be distinguished from similar native Westchester Co., Yorktown Heights, 23- apionine weevils found in eastern North XII-1996. PENNSYLVANIA: Berks Co., America; and summarize information on Robesonia, V-1994, 1996 and VII, IX-1996, the biology, habits, food plants, and feeding 6-VI-1997. Bucks Co., Quakertown, 1-VI- damage of /. virens in its native range. 1998; Nockamixon State Park, 1-VI-1998. Carbon Co., Normal Square, 20-V-1998; Initial Detection, Additional U.S. Lehighton, 20-V-1998; Jim Thorpe, 20-V- Records, and Specimen Deposition 1998. Chester Co., Honey Brook, 30-VII- The first specimens of/. virens were tak- 1997. Columbia Co., Centralia, 8-VI-1998; en in pitfall traps on a farm near Robesonia Numidia, 8-VI-1998. Cumberland Co., Car- (Heidelberg Twp.), Pennsylvania (Berks lisle, 14-VI-1999; Summerdale, 10-VII- Co.) in May 1994. During 1994-1997, nu- 1998. Dauphin Co., Harrisburg, 18-V-1998. merous additional specimens of /. virens Delaware Co., Swarthmore, 18-VI-1998. were collected in pitfall traps set in pad- Lancaster Co., Kirkwood, 28-V1I-1997. docks of grazed pastures of 5 other Penn- Lebanon Co., Myerstown, 6-VIII-1997. Le- sylvania counties. Other locality records, high Co., Coopersburg, 1-VI-1998; Fogles- many from 1998 and 1999, have become ville, 7-VIII-1997. Luzerne Co., Freeland, available for Connecticut, Delaware, Mary- 20-VII-1998; Hazelton, 20-VII-1998; land, New York, New Jersey, Pennsylvania, White Haven, 20-VIM998. Lycoming Co., and Virginia. The collection of specimens Montgomery, 25-VI-1998. Monroe Co., of /. virens from Connecticut, Maryland, Appenzell, 7-VII-1998; Brodheadsville, 7- and Virginia was the direct result of those VII-1998; Saylorsburg, 7-VII-1998; states' Cooperative Agricultural Pest Sur- WStroudsburg, 7-VII-1998. Montgomery Co., vey-Eastern Region (CAPS) activities for of Boyertown, IX-1997. Montour Co., 1999. All known eastern U.S. collections of Danville, 26-V-1998. Northampton Co., /. virens (mostly by pitfall trapping, and Bethlehem, 1-VI-1998. Northumberland sweep-net or vacuum sampling) are listed Co., Dalmatia, 22-V-1998; Malta, 8-VI- below and mapped in Fig. 1. 1998. Perry Co., Marysville, 27-VII-1998; CONNECTICUT: Litchfield Co., East Roseglen, 7-VI-1999. Philadelphia Co., Canaan, 15-VI-1999; Norfolk, 15-VI-1999. Philadelphia, Fairmont Park, 14-VII-1998. DELAWARE: New Castle Co., Newark, 21 Schuylkill Co., Lavelle, 8-VI-1998; New- & 22-IV-1998; Smyrna, 25-VII-1999. town, 18-V-1998; Pitman, 8-VI-1998; MARYLAND: Cecil Co., Calvert, 22-IV- Schuylkill Haven, 18-V-1998. York Co., 1999; Chesapeake City, 22-IV-1999; Con- Longlevel, 28-V-1998. VIRGINIA: Lou- owingo, 22-IV-1999; Cowentown, Fair Hill doun Co., Rte. 15, nr. Gilberts Corner, 30- NRMA, 22-IV-1999; Elkton, 22-IV-1999; VI-1999. Rising Sun, 22-IV-1999; Warwick, 22-IV- The first specimens of /. virens, repre- 1999. Harford Co., Dublin, 26-IV-1999. senting the initial detection of this Palearc- Kent Co., Chestertown, 3-V-1999; Golts, tic species in North America, were initially 22-IV-1999; Kennedyville, 3-V-1999; Mill- identified by ERH and later confirmed by ington, 22-IV-1999. NEW JERSEY: Hun- MAAZ. Voucher specimens are deposited terdon Co., Snyder research farm, 30-VI- in collections of the following individual 1992 and 14 «fe 21-VII-1992; no precise lo- and institutions: Cornell University (Ithaca, VOLUME 102, NUMBER 1 153 154 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2—6. Ischnopterapion virens. 2, Adult, male, dorsal aspect. 3, Head andpronotum, male, lateralaspect. 4, Head and pronotum, female, lateral aspect. 5, Head and pronotum, male, dorsal aspect, showing punctation in part. 6, Elytra, male, dorsal aspect. Scale line for Fig. 2 = 0.5 mm. (Figs. 3-4 after Morris 1990; Figs. 5-6 after Gonget 1997.) way, Sweden) (Gonget 1997). Occurring black, with aeneous or metallic greenish re- commonly and abundantly throughout Eng- flection, elytron distinctly metallic greenish land, Wales, Ireland, and locally in Scotland or greenish-blue, shining, vestiture rather (Fowler 1891, Morris 1990), it is recorded sparse. Male rostrum (Fig. 3) shorter, slight- also from Siberia, Syria, and northern Africa ly curved, about IVa times as long as pro- (Algeria) (Hoffmann 1958, Brito-Castro and notum, pubescent at apex. Female rostrum Oromi-Masoliver 1986), and the Canary Is- (Fig. 4) longer, curved, about P/i times as lands (Tenerife and La Palma) (Brito-Castro long as pronotum, pubescent only at base. and Oromi-Masoliver 1986). Frons slightly convex, finely and sparsely punctured. Vertex finely and sparsely punc- Adult Redescription tured apically and transversely striate and The following redescription is adapted shining basally. Eye prominent, medium- from Gonget (1997). Length 1.8-2.6 mm, sized to large and almost round. Pronotum oblong, slightly convex (Fig. 2). Body (Fig. 5) broader than long, broadest behind VOLUME 102. NUMBER 1 155 middle, slightly convex longitudinally, dif- 7 with 3+4 and 8 with 5+6 (joining 7+8 fusely, finely and shallowly punctured. Bas- in Kissingeria). It differs from Apion pan- al fovea ofpronotum (Fig. 5) rather distinct amense Sharp (still placed in Apion sensu but short. Elytron (Fig. 6) oblong to elon- lato) by the shorter rostrum in both sexes, gate with sides rounded, broadest just be- ca. 1.25 X length of pronotum in the male hind middle, slightly convex longitudinally and ca. 1.50 in the female (ca. 1.42 in the and somewhat depressed on disc, separately male and ca. 1.70 in the female in Apion rounded at apex. Elytral striae moderately panamense) and by the latter having only strongly punctured, first stria deeper on disc one specialized seta on interstria 9. From than others. Striae joined at apex as fol- the group ofspecies assigned to Ceratapion lows: 1+2+9, 3+4+7, and 5+8+6. Fine by Kissinger (but not belonging to this ge- vestiture arranged in single row on each in- nus, cf. Alonso-Zarazaga 1990) and which, terstice, with one specialized seta at apical being absent from his key, should key also third of 7th interstice. Legs black with fem- to the same couplet, /. virens is distinguish- ora having greenish or bluish metallic re- able by the union of striae at the apex (as flection. Flight wings fully developed or ru- above), and the lack of a spine on the 1st dimentary. Male genitalia illustrated by mesotarsomere in the male (present in the Alonso-Zarazaga (1990: 120). so-called Ceratapion) and genital features. Following the criteria explained in Alon- Diagnosis so-Zarazaga (1990), Kissingeria and the so- Ischnopterapion virens can be separated called Ceratapion sensu Kissinger belong from the majority of North American to the tribe Oxystomatini subtribe Trichap- apionine weevils by the following combi- iina. Ischnopterapion virens belongs to the nation ofcharacters: rostrum curved in both subtribe Synapiina of the same tribe, which sexes, ventral surface ofhead with low sub- includes three other Palearctic genera, not ocular ridges, dorsal margin of scrobe not yet known to be in the Nearctic. The two dentiform, prothorax moderately constrict- subtribes formerly present in the Nearctic ed behind middle, lacking basal flange, el- Region were: Trichapiina (including the ytra apically neither expanded nor with large and heterogeneous genus Trichapion deep pits, male metastemum lacks median Wagner, Kissingeria Alonso-Zarazaga, and tubercle near posterior margin, male tibiae several species groups still placed in Apion without hooks (non-mucronate), legs dark, sensu lato, which will probably need genera tarsal segments 1 longer than wide, tarsal of their own after a thorough revision and segments 3 moderately large, strongly bi- study of their relationships) and Oxysto- lobed, tarsal claws with acute basal tooth. matina (represented in the Nearctic Region Using these characters, in Kissinger's by two genera: Mesotrichapion Gyorffy (1968) key to North and Central American (with the single species Mesotrichapion apionid weevils, this newly detected species {Loborhynchapion) cyanitinctum (Fall)) keys to couplet 27. It differs from the mem- and Eutrichapion Reitter, embracing three bers of Pseudapion sensu Kissinger {Apion species in two different subgenera, Eutri- disparatum and A. varicorne species chapion s. str. and Leconteapion Alonso- groups, most of the species transferred to Zarazaga, namely: Eutrichapion {Eutri- genus Kissingeria by Alonso-Zarazaga chapion) viciae (Paykull) (= Apion alas- (1990)] by the black legs and antennae (legs kanum Fall), E. {Leconteapion) cavifrons in part and/or antennae yellow in both (LeConte) and E. (L.) huron (Fall). All the groups of species in Kissingeria), only ely- Nearctic members of Oxystomatina have a tral interstria 7 with one specialized seta median tubercle on the male metastemum (one specialized seta on elytral interstria 7 and key separately at couplet 13 in Kissin- and 9 in Kissingeria), striae at apexjoining ger's key. Even if the median tubercle in 156 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON these species is overlooked, members of widely in mixture with grasses for animal Eutrichapion have blackelytra and one spe- feed, it is one ofthe most importantherbage cialized seta on interstriae 7 and 9, and legumes in Britain (Wiech and Clements members of Mesotrichapion subgenus Lo- 1992). Chief among the invertebrates dam- borhynchapion, although having metallic aging white clover in southeastern England elytra, have simple claws, the frons is strio- are apionine weevils, especially /. virens late and the male mesorostrum is strongly (Wiech and Clements 1992). In Bavaria in lobed in dorsal view. In addition, /. virens 1930 and 1931, a serious local and heavy is different from all these species by its first infestation by /. virens occurred in red clo- elytral stria being more deepened than the ver {T. pratense); larvae, by tunnelling in rest on the disc. Mesotrichapion (L.) cyan- the taproot and root-crown, reduced plant itinctum is recorded only from Manitoba growth so that fodder and seed yields were and Quebec, and its host plant appears to much decreased (Andersen 1932). In April be Astragalus (Fabaceae). and May 1926 in upper Austria, fields of red clover were heavily damaged by /. vi- Host Plants and Damage rens, with as much as 80% of this forage Plants of the family Fabaceae (= Legu- crop damaged (Wemeck 1930). minosae) are the principal hosts of adult The following species of Trifolium have and larval /. virens, with a decided prefer- been recorded as host plants of/. virens: T. ence shown for species of clover (Trifoli- campestre Schreb. (Brito-Castro and wn). These weevils are responsible for two Oromi-Masoliver 1986); T. pratense L. types of feeding injury to clover: adults (Fowler 1891, Wemeck 1930, Markkula feed on the foliage (leaflets), whereas lar- and Myllymaki 1957, Stein 1965, 1968); T. vae feed or mine in the petioles, stems (sto- alexandrimnn L. (Frauenfeld 1866, Delas- lons), root-crown, and possibly taproots. sus 1936, Dieckmann 1977); T. arvense L. Ischnopterapion virens larvae mine the (Hoffmann 1958); T. incarnatum L. and T. stems of Trifolium pratense and generally fragiferum L. (Dieckmann 1977); and T. re- bore downward towards the root-crown and pens L. (Dieckmann 1977, Stein 1968, roots during development (Stein 1965). In Wiech and Clements 1992). one instance, root damage was observed in Biology, Seasonal History, and red clover, T. pratense var. spontaneiim Habitat (Stein 1965). Adult feeding creates small (ca. 0.5 The following account ofthe biology and mm-), elongate punctures in the leaf tissue seasonal history of /. virens is a summary (Wiech and Clements 1992: 438). Feeding drawn from field and laboratory observa- sites vary depending on the age ofthe wee- tions of various investigators: Werneck vil. For example, newly emerged (first sea- (1930), Andersen (1932), Bovien and Jor- son) adults (in July) exhibit apreference for gensen (1934, 1936), Markkula and Myl- feeding on the apical portion ofclover leaf- lymaki (1957, 1958), Kokorin (1964), lets, while "older" adults (14-21 days min- Scherf (1964), and Stein (1965, 1968, imum since emergence) use the basal por- 1972a, b). Detailed studies on the mor- tion of leaflets (see Stein 1965: 393, fig. 3). phology, biology, seasonal history, and be- Adult feeding damage to white clover fo- havior of /. virens are provided by Stein liage is illustrated in Fig. 7. (1965, 1968). Although /. virens is generally consid- Adult weevils overwinter in the soil near ered to be a minor pest of clover crops in host plants or among plant debris at the Europe (Balachowsky 1963), it can be a base of their hosts {Trifolium spp.) and re- major pest causing significant damage. Be- appear in the spring, generally as early as cause white clover {T. repens) is grown late February-early March. After weevils VOLUME 102. NUMBER 1 157 Figs. 7-10. Ischnopterapion virens, adult and larval feeding damage to white clover and immature stages. 7. Adult feeding damage to white clover leaflet. 8, Larva tunneling in white clover stem. 9, Pupa exposed in white clover stem. 10. Pupa in white clover stem, enlarged. emerge from their overwintering sites, in- emerge, harden and become fully colored tensive feeding begins, followed by a spring in about 2-3 days. Preimaginal (egg, larval mating period, generally from April to and pupal) development takes place from June. Oviposition generally begins in mid- May through much of August. Adults April to early May and continues into late emerge throughout the summer, but gener- May-early June. The female uses her man- ally in July and August. Newly emerged dibles to chew a cavity in the petiole or weevils feed intensively on the leaves for stem, lays a single egg, and seals off the about four weeks, followed again by a non- egg cavity with a mixture ofexcrement and feeding period of about 40 days, and then abdominal secretions, probably to protect a dispersal, or migratory phase. Adults fly the egg and to prevent its desiccation. After mainly during late summer and autumn about 7-8 days, first-instar larvae hatch and (August-November), with a lower temper- chew long mines in the stems, mostly ature threshold of 10°C (Stein 1972a), after downward. Larvae move downward into which they feed until the beginning ofwin- the stem where actual feeding will begin ter. Mating takes place in early to mid-au- (Fig. 8). After about four weeks, a larva tumn. Thus, there are two periods of mat- completes its development and pupates in a ing, one in autumn and one in spring. Dur- cavity in the lower part of the clover stem ing a weevil's lifespan, at least three feed- (Figs. 9, 10), just under the epidermis, di- ing periods occur: one directly after adult rectly above the root. The pupal stage lasts emergence in summer, a second from Sep- about 7 days, and first generation adults tember to the beginning of the overwinter- 158 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ing period, and a third after the overwin- and meadows, and in grassy fields and cul- tering period in spring (Stein 1965). There tivated clover fields (Gonget 1997). In Brit- is one generation annually. ain, this weevil is found ''in a wide variety Egg development requires approximately ofopen and grassy biotopes, including sand 8 days, with a range of 6-12 days (Stein dunes and agricultural land" (Morris 1990). 1968). Bovien and Jorgensen (1936) re- Sampling and Natural History ported a range of 14-18 days for egg de- velopment. Three larval instars occur in /. Observations in Pennsylvania virens. Larval development takes about 27- To gain a better understanding of popu- 28 days (Stein 1968), whereas Bovien and lation numbers of /. virens at the original Jorgensen (1936) noted that 28-30 days collection site in Robesonia (Heidelberg were necessary for larval development. The Twp.), PA, one of us (JFS) conducted a average duration from egg to the pupal sampling regime of white and red clover. stage is approximately 34 days, with little Weekly, 1 m-, vacuum samples, taken or no difference between males and fe- from April 1998 through mid-July 1999, males, and a range of 30-39 days. The av- have yielded adult weevils nearly constant- erage length of the pupal stage is 7 days. ly. Even during the winter months (Decem- Adults can survive as long as 360 days. Un- ber 1998 through March 1999), adults were der laboratory conditions, each mated fe- taken in vacuum samples; the 1998-99 win- male is capable oflaying a maximum of90 ter was unusually mild, with temperatures to 166 eggs; however, for the majority of seldom falling below freezing and with lit- beetles in the field the number is much less, tle to no snow cover for the majority ofthis approximately 40-45 eggs per female period. Numbers of adults taken in the (Stein 1968). weekly samples actually increased during The seasonality and habits of/. virens in the winter months, as compared to those the Old World vary according to the inves- collected during the dry, warm months of tigator. In upper Austria, for example, eggs summer and early autumn of 1998. From are laid on the root-crown or stems of red the surprisingly high numbers taken weekly clover, with first instar larvae found in Sep- during the winter, the population climbed tember; early instars overwinter in the tap- somewhat during the spring, until new gen- roots, make their way into the heart of the eration adults began to emerge. At this plant in spring, and pupate in July (Wemeck point, sample sizes increased dramatically 1930). In Germany, larvae of /. virens live to a high of 180 adults from a 1 m- area on in the tap root or root collar and overwinter June 15, 1999. Winter and early spring as adults in the soil (Andersen 1932). Based sample numbers averaged ca. 32 adults/m-^. on laboratory rearings in Denmark, Bovien From the June 15 peak, the number of and Jorgensen (1934) found that field-col- adults present in the weekly samples de- lected adults laid eggs in May in the mid- clined drastically to a low of in the July ribs of leaves, that eggs hatched in early 13, 1999 sample. The extreme drought of June, and that larvae later mined in the leaf the 1999 summerhas all but eliminated suc- petioles, eventually penetrating into the culent host plant material in the sampling roots of smaller plants, but continuing to area. The July 13 sample is the first such feed in the stems of larger plants. Markkula sample to be devoid of adults since sam- and Myllymaki (1957) found that /. virens pling began in April 1998. females oviposited in the stems of red clo- On numerous occasions in 1998 and ver from late May to early August. 1999, we have observed the feeding habits Ischnopterapion virens is associated gen- of larvae of /. virens on white and red clo- erally with its leguminous host plants on ver at Robesonia, PA. Mature white clover slopes, along roadsides, in coastal pastures plants spread by creeping, above ground. VOLUME 102. NUMBER 1 159 prostrate stems (stolons) that also root at the surface devoured ranged from 0% to a high nodes (Uva et al. 1997). On May 21, 1998, of 21.25%, and averaged 2.64% of leaf sur- we (ERH, JFS) split open, by means of a face sampled during a 5 week period (sam- razor-blade knife, many stems (stolons) of ples collected weekly from June 15 through white clover plants growing adjacent to July 13, 1999). grazed pastures and found that the majority The pest status of /. virens and its long- were mined by larvae of /. virens (Fig. 8). term impact on white and red clover in the Cut plant stems with larvae were brought eastern United States are equivocal and will into the lab and adults were reared to con- remain so until further field studies are con- firm their identity. In addition to the dis- ducted. covery of many mature larvae, we also ob- served pupae (Figs. 9, 10) inside these Acknowledgments stems (stolons). Random observations oflarval feeding in We are grateful to the following individ- the stems ofwhite cloverrevealedextensive uals for providing us with supplemental consumption of plant tissue. Larvae feed collections of /. virens in the Northeast: W. upon the central, pithy areas of the lower H. Day (USDA, Beneficial Insects Research prostrate stems (stolons). During peak lar- Laboratory, Newark, DE) for Delaware, J. val incidence (late May to early June), Ingerson-Mahar (Rutgers University, New many stolons exhibited evidence offeeding Brunswick, NJ) for New Jersey, and M. C. by larvae over the entire length of the sto- Thomas (Florida State Collection ofArthro- lon. In such instances, larvae were found at pods, Gainesville, FL) for New York. Dis- approximately 2.5 cm intervals over the tributional data acquired through the Co- length of the infested stolon, and those sto- operative Agricultural Pest Survey (CAPS) lons with late-stage larvae and pupae were activities in the following states is grateful- often in a state of semi-collapse, contained ly acknowledged: Connecticut (Donna El- large areas of browned and dessicating tis- lis), Maryland (Dick Bean), and Virginia sue, and often exhibited small holes through (Eric Day). We also thank the following in- the epidermis of the plant. On 26 May 1998, RAB found larvae and pupae in the dividuals for their assistance in the field and for collecting pitfall trap samples in Penn- stems of red clover and noted nearly 100% sylvania, New York, and Vermont during infestation ofthese plants, with many dying the past 4 years: Steve LaMar, John Ever- and browning stems. hart, Ruth Haldeman, Andy Paolini, Kevin In spite of extensive larval feeding dam- Nelson, Jeremy Everhart, Karen Jackson, age to the stems (stolons) of white clover Allison Henry, Chrissa Rose, Sharyn in grazed pastures at Robesonia, PA, these Leach, Andy Davis, Mark Giacomin, Amy plants nonetheless appeared vigorous and Kellerman, Chante Gordon, Cindy Long, healthy. In sharp contrast, the red clover Brian Dougherty, Mike Hutchinson, Jim plants appeared to be more heavily impact- Gavlik, Kanesha Williams, Michelle Lorek, ed by the larval feeding of /. virens, espe- cially the above ground stems (RAB obser- Heather Lagoda, Matthew Hartman, and Claudia Violette. Chad M. Petrovay and vations). On white clover, adults feed mostly in Justin Newell are acknowledged for their interveinal areas of the foliage, producing extensive survey findings of /. virens in oval to elongate-oval holes. In the summer Pennsylvania. We greatly appreciate the of 1999, JFS attempted to quantify the leaf critical review of an earlier draft ofthis pa- area fed upon in relation to the total leaf per by A. G. Wheeler, Jr. (Clemson Uni- surface area, using computer imaging and versity), and also the comments provided measurement packages. Percentage of leaf by two anonymous reviewers. 160 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Literature Cited Biology and phylogeny of Curculionoidea: Pro- ceedings of a symposium convened at the XVIII Alonso-Zarazaga, M. A. 1990. Revision of the su- InternationalCongressofEntomology,Vancouver, praspecific taxa in the Palaearctic Apionidae Canada, July 3-9, 1988. 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