h e monitoring cucumber beetles in sweetpotato and

HORTICULTURAL ENTOMOLOGY

Monitoring Cucumber Beetles in Sweetpotato and Cucurbits withKairomone-Baited Traps

D. MICHAEL JACKSON,1 K. A. SORENSEN,2 C. E. SORENSON,2 AND R. N. STORY3

J. Econ. Entomol. 98(1): 159Ð170 (2005)

ABSTRACT Seven kairomone formulations (Trece, Inc., Salinas, CA) were evaluated for theireffectiveness as attractants for luring three species of cucumber beetles into Pherocon CRW traps(Trece, Inc.) in cucurbit and sweetpotato Þelds. The spotted cucumber beetle, Diabrotica undeci-mpunctata howardiBarber; the banded cucumber beetle,Diabrotica balteataLeConte; and the stripedcucumber beetle,Acalymmavittatum(F.), were captured in this study. TRE8276 (TIC mixture: 500 mgof 1,2,4-trimethoxybenzene, 500 mg of indole, and 500 mg of trans-cinnamaldeyde) and TRE8336 (500mg of 1,2,4-trimethoxybenzene, 500 mg of trans-cinnamaldeyde, 500 mg of 4-methoxyphenethanol)were the most effective lures for spotted and striped cucumber beetles. None of the kairomone lureswas very effective for attracting banded cucumber beetles. Three population peaks of spottedcucumber beetles were observed in cucurbit and sweetpotato Þelds at the U.S. Vegetable Laboratory(Charleston, SC). The efÞcacy of TRE8276 declined rapidly after 2 wk in the Þeld. An improved designof the Pherocon CRW trap, with a yellow bottom and more-tapered top section, was more effectivefor capturing cucumber beetles than the original trap design made entirely of clear plastic. Bandedcucumber beetles were not captured in sweetpotato Þelds at inland locations in North Carolina orSouth Carolina.

KEY WORDS Diabrotica undecimpunctata howardi, Diabrotica balteata, Acalymma vittatum, Ipo-moea batatas, Cucurbitaceae

VEGETABLE CROPS IN THE United States are damaged byseveral species of diabroticite beetles (Coleoptera:Chrysomelidae: Galerucinae: Luperini: Diabroticina),including the cucumber beetles (Krysan 1986, Arnettet al. 2002). The spotted cucumber beetle (or south-ern corn rootworm), Diabrotica undecimpunctatahowardi Barber; the banded cucumber beetle, Dia-brotica balteata LeConte; and the striped cucumberbeetle, Acalymma vittatum (F.), are the most impor-tant diabroticite pests of vegetable crops in the south-eastern United States (McKinlay 1992, Capinera2001). The spotted and banded cucumber beetleseach have a wide host range (Arant 1929, Isley 1929,Saba 1970), including cucurbits (Cucurbitaceae) andsweetpotatoes, Ipomoea batatas Lam (L.) (Convolvu-laceae). However, larvae of the striped cucumber bee-tle are found primarily on cucurbits (Houser andBalduf 1925).

The chemical ecology of diabroticite beetles hasreceived considerable research attention, and it isbelieved that these species coevolved with the cucur-bits (Metcalf and Lampman 1989b, 1991; Metcalf 1986,1994). The importance of cucurbitacins as powerful

phagostimulants and arrestants has been well docu-mented (Chambliss and Jones 1966, Metcalf et al.1980). Shaw et al. (1984) reported that a mixture ofcucurbitacin and carbaryl killed corn rootworms intraps so that they could be easily counted. However,cucurbitacins are not active as long-range olfactoryattractants (Rhodes et al. 1980, Branson and Guss1983), so volatile components are needed to lure cu-cumber beetles toward traps or toxic, cucurbitacin-containing baits.

Adult cucumber beetles are attracted to the blos-soms of many cucurbit species, where they feed onnectar, pollen, ßowers, and developing fruits (Arant1929, Fisher et al. 1984, Hesler 1998). More than 40volatile chemicals have been isolated from the ßowersof Cucurbita maxima (Andersen and Metcalf 1986,1987, Andersen 1987). Among them are chemicallyrelated volatile phenylpropanoids that are producedin large amounts by ßowers during pollen maturationand are attractive to adult cucumber beetles (Metcalfand Lampman 1991). A major attractant for thespotted cucumber beetle from C. maxima blossoms is(E)-cinnamaldehyde (trans-cinnamaldehyde) (Met-calf and Lampman 1989a, 1991). Morgan and Crumb(1928) Þrst reported that spotted cucumber beetleswere attracted to cinnamic aldehyde and cinnamicalcohol, but they did not associate these chemicalswith any host plants. Lampman et al. (1987) also foundthat several structurally related benzenoid com-

1 USDAÐARS, U.S. Vegetable Laboratory, 2700 Savannah Highway,Charleston, SC 29414.

2 Department of Entomology, North Carolina State University,Raleigh, NC 27695.

3 Department of Entomology, Louisiana State University, BatonRouge, LA 70803.

pounds, including chavicol, phenylacetaldehyde, ben-zyl acetone, phenethyl alcohol, phenyl acetate, vera-trole, methyl eugenol, methyl isoeugenol, eugenol,and isoeugenol, acted as attractants for spotted cu-cumber beetles. Spotted cucumber beetles were moreattracted to phenylacetaldehyde than other species(Lampman et al. 1987). Andersen and Metcalf (1986)found that indole, isolated from the blossoms ofC. maxima cultivars, was highly attractive to stripedcucumber beetles.

A mixture of Cucurbita blossom components, 1,2,4-trimethoxybenzene, indole, and trans-cinnamalde-hyde (TIC mixture), is very attractive to several spe-cies of diabroticite beetles, including spotted andstriped cucumber beetles (Metcalf and Lampman1991). Lampman and Metcalf (1987) reported thatspotted cucumber beetles also were attracted to amixture of veratrole, indole, and phenylacetaldehyde(VIP mixture). Relatively minor changes in themakeup of these semiochemical mixtures can radicallychange the speciÞcity and attractiveness to cucumberbeetles (Lampman et al. 1987; Lampman and Metcalf1987, 1988). Lewis et al. (1990) reported that trapsbaited with the three-component TIC mixture showeda synergistic effect and captured twice as many stripedcucumber beetles as would be expected with the in-dividual components. Banded cucumber beetles havenot been evaluated for response to volatile compo-nents from cucurbits, even though they aggregate andfeed heavily on them (Hesler 1998).

Volatile ßoral attractants from cucurbits and maizeshow promise as semiochemicals for monitoring and

managing diabroticite beetles (Metcalf and Metcalf1992, Metcalf 1994). TIC-baited traps have been usedto monitor cucumber beetle populations in peanuts,Arachis hypogaea L. (Herbert et al. 1996), cucurbits(Cucurbitaceae) (Lewis et al. 1990, Hoffmann et al.1996), soybean,Glycinemax(L.) Merr. (Whitworth etal. 2002), and maize, Zea mays L. (Lampman andMetcalf 1987). Volatile semiochemicals also might beused to attract and concentrate preovipositional fe-males into Þelds to complement pest control strategies(Hammack 2003). Volatile attractants also have beentested as an orientation disruption tool for adult west-ern corn rootworm, Diabrotica virgifera virgifera Le-Conte (Wennemann et al. 2001).

Unbaited Pherocon AM sticky traps (Trece, Inc.,Salinas, CA) were suggested as a scouting tool formonitoring adult Diabrotica spp. populations and forpredicting damage by corn rootworm larvae (Heinand Tollefson 1984, 1985). However, Whitworth et al.(2002) tested a version of the Pherocon CRW kairo-mone trap (Trece, Inc.) and found it to be moreeffective than Pherocon AM or Multigard traps formonitoring corn rootworm populations. Nowatzki etal. (2002) also used Pherocon CRW kairomone trapsand found them to be much more efÞcient than emer-gence traps for predicting the Þrst day of corn root-worm emergence.

Trece, Inc., is currently marketing Pherocon CRWkairomone traps and Þve kairomone lures for moni-toring cucumber beetles: Pherocon TR-CRW-M(TRE8275) for Mexican corn rootworm, Diabroticavirgifera zeaeKrysan & Smith; Pherocon TR-CRW-W

Table 1. Details of cucumber beetle trapping experiments in South Carolina, North Carolina, and Louisiana, 1999–2003

Experimentdesignation(state-yr)

Field locationCounty/parrish

stateGeographic coordinates Date exp setup (DOYa)

SC-1999-A USVL Charleston Co., SC (32.81 N, 80.07 W) 22 Sept. (DOY 265)SC-1999-B USVL Charleston Co., SC (32.81 N, 80.07 W) 22 Sept. (DOY 265)SC-2000-A USVL Charleston Co., SC (32.81 N, 80.07 W) 7 June (DOY 159)SC-2000-B USVL Charleston Co., SC (32.81 N, 80.07 W) 12 June (DOY 164)SC-2000-C Duford Horry Co., SC (34.20 N, 79.05 W) 15 June (DOY 167)SC-2000-D Blackville Barnwell Co., SC (33.35 N, 81.30 W) 30 June (DOY 182)SC-2001-A USVL Charleston Co., SC (32.81 N, 80.07 W) 15 June (DOY 166)SC-2001-B USVL Charleston Co., SC (32.81 N, 80.07 W) 15 June (DOY 166)SC-2001-C Ehrhardt Bamberg Co., SC (33.21 N, 81.07 W) 21 June (DOY 172)SC-2002-A USVL Charleston Co., SC (32.81 N, 80.07 W) 12 June (DOY 163)SC-2002-B USVL Charleston Co., SC (32.81 N, 80.07 W) 19 June (DOY 170)SC-2002-C Hollywood Charleston Co., SC (32.72 N, 80.28 W) 5 Aug. (DOY 217)SC-2002-D Ehrhardt Bamberg Co., SC (33.15 N, 81.10 W) 8 Aug. (DOY 220)SC-2003-A USVL Charleston Co., SC (32.81 N, 80.07 W) 9 July (DOY 190)SC-2003-B USVL Charleston Co., SC (32.81 N, 80.07 W) 9 July (DOY 190)NC-2000-A Fremont Wayne Co., NC (35.56 N, 78.02 W) 25 July (DOY 207)NC-2001-A McGee Crossroads and

ClintonJohnston Co., NCSampson Co., NC

(35.52 N, 78.58 W)(35.02 N, 78.28 W)

4 June (DOY 155)4 June (DOY 155)

NC-2001-B Newton Grove, McGeeCrossroads, and Clinton

Sampson Co., NCJohnston Co., NCSampson Co., NC

(35.27 N, 78.32 W)(35.52 N, 78.58 W)(35.02 N, 78.28 W)

9 July (DOY 190)9 July (DOY 190)9 July (DOY 190)

NC-2001-C AtchinsonÕs Mill, HaresCrossroads, and Narron

Johnston Co., NCJohnston Co., NCJohnston Co., NC

(35.66 N, 78.26 W)(35.66 N, 78.25 W)(35.71 N, 78.24 W)

19 July (DOY 200)19 July (DOY 200)19 July (DOY 200)

NC-2002-A Heßin, Stancils Chapel, andKenly

Johnston Co., NCJohnston Co, NCJohnston Co, NC

(35.72 N, 78.20 W)(35.70 N, 78.20 W)(35.63 N, 78.15 W)

12 Aug. (DOY 224)12 Aug. (DOY 224)12 Aug. (DOY 224)

LA-2001-A Baton Rouge E. Baton Rouge, LA (30.41 N, 91.10 W) 9 July (DOY 190)

a Day of the year (Julian date).

160 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 98, no. 1

(TRE8275) for western corn rootworm; Pherocon TR-CRW-S (TRE8276) for southern corn rootworm(spotted cucumber beetle); Pherocon TR-CRW-N(TRE8391) for northern corn rootworms, Diabroticabarberi Smith & Lawrence; and Pherocon TR-CRW-W/N (TRE8391) for mixed populations of westernand northern corn rootworms for use in maize pestmanagement (Trece, Inc. 2003). The purpose of thisstudy was to evaluate the Trece kairomone lures andPherocon CRW traps in cucurbit and sweetpotatoÞelds in the southeastern United States.

Materials and Methods

Experiments at the U.S. Vegetable Laboratory(USVL), Charleston, SC. Trapping experiments forcucumber beetles were conducted at the USVL from1999 to2003(Table1).Eachyear, the samethreeÞelds(0.68, 0.70, and 0.83 ha, separated by at least 200 m)were planted to alternating, adjacent rows of ÔEarlySummer CrookneckÕ squash (Seeds by Design, Inc.,Maxwell, CA), ÔSmall Sugar/New EnglandÕ pumpkin(Hollar Seeds, Rocky Ford, CO), ÔHales Best JumboÕcantaloupe (Western Hybrid Seed, Inc., HamiltonCity, CA), and ÔPoinsett 76Õ cucumber (Dorsing Seeds,Inc., Nyssa, OR). Only one-third of each Þeld wasplanted on a given planting date. New plantingswere made every 3Ð4 wk to ensure that healthy, ßow-ering plants were present throughout the season. TheÞrst planting was made in early April, and the Þnalplanting was made in September of each year. Fertil-

izer (455 kg/ha of 10Ð10-10) was incorporated intothe plant beds. Before planting, metolachlor (Dual IIMagnum, Syngenta Crop Protection, Inc., Greens-boro, NC) and oryzalin (Surßan A.S., Dow Agro-Sciences, Indianapolis, IN) were incorporated into thesides, but not the top, of the bedded rows to helpcontrol weeds. No insecticides were used. Rows were�1 m apart, and seeds were planted on �30-cm cen-ters. Fields were irrigated and cultivated as needed.

Each year, traps also were placed in two sweet-potato Þelds that were separated by at least 300 m atthe USVL. Over the 5 yr of this study, seven differentsweetpotato Þelds were used. They were 0.60, 0.61,0.75, 0.82, 0.88, 0.92, and 1.10 ha. Fields consisted ofmixed plantings of several sweetpotato clones fromthe sweetpotato breeding program (Dukes et al.1992). Fertilizer (545Ð682 kg of 10Ð10-10) was incor-porated into the rows, and Þelds were planted in Juneof each year. Soon after planting, metolachlor (Dual IIMagnum) was applied for weed control. Sethoxydim(Poast, BASF, Research Triangle Park, NC) was usedas needed for control of grasses. During these exper-iments each year, one of the sweetpotato Þelds at theUSVL was sprayed two to three times with carbaryl(Sevin XLR Plus; Bayer Corporation, Kansas City,MO) as part of the protocols for management of thesweetpotato weevil, Cylas formicarius elegantulus(Summers). No insecticides were used on the otherÞelds.

The seven kairomone lures obtained from Trece,Inc., were TRE8274 (low dose of TIC mixture: 250 mg

Table 1. Continued

Crop Replicates Trap design Times traps checked Lures evaluated (TRE-)

Cucurbit 2 Old 10 8274, 8275, 8276, 8279, 8282, 8336Sweetpotato 2 Old 10 8274, 8275, 8276, 8279, 8282, 8336Sweetpotato 2 Old 39 8274, 8275, 8276, 8279, 8282, 8336, 8391Cucurbit 2 Old 39 8274, 8275, 8276, 8279, 8282, 8336, 8391Sweetpotato 2 Old 4 8274, 8275, 8276, 8279, 8282, 8336, 8391Sweetpotato 2 Old 4 8274, 8275, 8276, 8279, 8282, 8336, 8391Cucurbit 3 Old & New 56 8276, 8279, 8282, 8336Sweetpotato 2 Old & New 56 8276, 8279, 8282, 8336Sweetpotato 3 New 9 8276, 8279, 8282, 8336Cucurbit 3 Old & New 74 8276, 8279, 8282, 8336, 8391Sweetpotato 2 Old & New 74 8276, 8279, 8282, 8336, 8391Sweetpotato 3 New 8 8276, 8279, 8282, 8336, 8391Sweetpotato 3 New 6 8276, 8279, 8282, 8336, 8391Cucurbit 3 New 36 8276, 8279, 8282, 8336, 8391Sweetpotato 2 New 36 8276, 8279, 8282, 8336, 8391Sweetpotato 2 Old 4 8274, 8276, 8279, 8336, 8391Sweetpotato 1 New 12 8274, 8275, 8276, 8279, 8282, 8336, 8391sweetpotato 1 New 12 8274, 8275, 8276, 8279, 8282, 8336, 8391Sweetpotato 1 New 7 8276, 8336Sweetpotato 1 New 7 8276, 8336Sweetpotato 1 New 7 8276, 8336Sweetpotato 2 New 5 8276, 8336Sweetpotato 2 New 5 8276, 8336Sweetpotato 2 New 5 8276, 8336Sweetpotato 3 New 4 8276, 8336Sweetpotato 3 New 4 8276, 8336Sweetpotato 3 New 4 8276, 8336Sweetpotato 4 New 3 8274, 8275, 8276, 8279, 8282, 8336, 8391

February 2005 JACKSON ET AL.: MONITORING CUCUMBER BEETLES 161

of 1,2,4-trimethoxybenzene, 250 mg of indole, and250 mg of trans-cinnamaldeyde), TRE8275 (500 mg ofeugenol and 750 mg of 4-methoxycinnamaldehyde;marketed as Pherocon TR-CRW-W or PheroconTR-CRW-M), TRE8276 (high dose of TIC mixture:500 mg of 1,2,4-trimethoxybenzene, 500 mg of indole,and 500 mg of trans-cinnamaldeyde; marketed asPherocon TR-CRW-S), TRE8279 (1,500 mg of euge-nol), TRE8282 (750 mg of 4-methoxyphenethanol and750 mg of 4-methoxycinnamaldehyde), TRE8336(500 mg of 1,2,4-trimethoxybenzene, 500 mg of trans-cinnamaldeyde, 500 mg of 4-methoxyphenethanol),and TRE8391 (750 mg of eugenol and 750 mg of 4-methoxycinnamaldehyde, marketed as PheroconTR-CRW-N or Pherocon TR-CRW-W/N) (Trece,Inc. 2003).

Two versions of the Pherocon CRW kairomone trapfrom Trece, Inc., were evaluated. The Þrst style ofCRW trap was constructed entirely of rigid, clearplastic. It had a cylindrical bottom cup (5.5 in depth,8 cm in diameter) that snapped onto a top section thatwas shaped like a truncated cone (8 cm in depth, 13 cmin diameter at bottom and 7 cm in diameter at the top)(illustrated in Whitworth et al. 2002). The bottom cupportion of the trap projected 4 cm into the top sectionuntil it engaged three tabs that held it in place. Be-cause of the difference in diameter of the trap sectionsand because the tabs held them apart, there was anopen space around the two portions of the trap thatvaried from 3 cm at the bottom to 0.5 cm where thesections met. This space allowed insects access intothe trap. The specially designed cylindrical kairomonelure dispensers (1.0 by 3.5 cm in diameter) (Trece,Inc. 2003) were attached by a plastic tab passedthrough a slit on a ßange at the bottom edge of the topportion of the trap so that they hung outside and belowthe middle of the trap. After the foil lining was re-moved, the kairomone-impregnated dispenser facedinward toward the open space between the trap sec-tions. At the top of the upper section of the trap therewas an indentation that held a cylindrical, toxic-baitpellet (1.2 by 1.3 cm in diameter) containing cornmeal, 3.9% carbaryl, and pulverized roots of buffalogourd, Cucurbita foetidissima H.B.K. (Trece, Inc.2003). This bait pellet was used to attract and killcucumber beetles that entered the trap. In addition tocucurbitacins, buffalogourd rootpowderalsocontainsvolatile attractants that further enhance feeding byDiabrotica larvae (Cosse and Baker 1999). Cucurbit-acins themselves are not volatile attractants of cucum-ber beetles (Rhodes et al. 1980, Branson and Guss1983). The bottom portion of the trap had a cylindricalindentation (2 cm in depth, 2.3 cm in diameter) for theattachment of a piece of PVC rigid conduit (2.3 cm indiameter). These support poles were 60 cm in lengthand sharpened so they could be pushed into theground. An improved version of the Pherocon CRWtrap, now being marketed (Trece, Inc. 2003), wasevaluated in 2001 and 2002. This improved trap, whichwe will refer to as the new trap design, had a cylin-drical bottom portion identical to the old design ex-cept that it was colored bright yellow. The upper

portion of the new design was similar to the originaldesign except that the top or roof of the trap wastapered instead of ßat, which was intended to directthe trapped beetles toward the cucurbitacin-contain-ing toxic bait at the peak. Each type of trap was po-sitioned, baited, and checked in the same way in theÞeld.

Traps were placed within rows of cucurbits and inopen spaces between plots of sweetpotatoes at theUSVL. In 2001 and 2002, old and new trap designs werecompared for each kairomone lure. In 1999 and 2000,only the old trap design was used, and in 2003 only thenew trap design was tested. Each Þeld was designatedas a replication in a randomized complete block de-sign. Traps were checked and beetles removed two tothree times per week (Table 1). Lures were changedevery 2 wk, and the lures were assigned randomly tonew trap positions at that time. Data were analyzed byanalysis of variance (ANOVA) using PROC GLM, andtreatment means were separated by Fisher least-sig-niÞcant difference (LSD) test (P � 0.05) (SAS Insti-tute 1989).Experiments at Other Locations. Trapping experi-

ments were conducted at Þve other Þeld locations inSouth Carolina during 2000Ð2003. Experiments wereset up as randomized complete block designs with twoto three replications, and traps were checked four tonine times on an irregular basis (Table 1). Lures thathad been in the Þeld for at least 2 wk were changedat the time the traps were checked. Lures were as-signed randomly to new trap positions each time theywere changed. In 2000, Pherocon CRW traps (olddesign) with all seven Trece lures were set up in agrowerÕs sweetpotato Þeld near Duford, and in asweetpotato Þeld at the Clemson UniversityÕs EdistoResearch and Education Center, Blackville, SC. In2001, traps (new design) with four Trece lures(TRE8276, TRE8279, TRE8282, and TRE8336) wereset up in a cooperating growerÕs sweetpotato Þeld nearEhrhardt, SC. In 2002, traps (new design) with ÞveTrece lures (TRE8276, TRE8279, TRE8282, TRE8336,and TRE8391) were placed in a sweetpotato Þeld at adifferent location near Ehrhardt, and in a cooperatinggrowerÕs sweetpotato Þeld in Hollywood, SC. Exceptfor the Duford site in 2000, no insecticides were ap-plied to these Þelds. At Duford, cultural practices wereat the growerÕs discretion; a preplant insecticide ap-plication of chlorpyrifos (Lorsban 15G, Dow Agro-Sciences) was made, but no foliar insecticides wereapplied.

Trapping experiments were conducted at 11 Þeldlocations in North Carolina during 2000Ð2002 (Table1). The old design of the Pherocon CRW kairomonetrap was used in 2000, whereas the new design wasused for 2001 and 2002 tests. Experiments were set upin randomized complete block designs, and traps werechecked four to 12 times each year (Table 1). Luresthat had been in the Þeld for at least 2 wk werechanged at the time the traps were checked. Lureswere assigned randomly to trap positions each timethey were changed. Cultural practices, including pre-plant insecticide applications, were at the growersÕ


discretions. In 2000, traps were set up with two rep-lications of Þve Trece lures (TRE8274, TRE8276,TRE8279, TRE8336, and TRE8391) in a growerÕssweetpotato Þeld near Fremont, NC. In 2001, threetrapping experiments were conducted in North Caro-lina. For the Þrst 2001 experiment (NC-2001-A,Table 1), traps were set up in sweetpotato Þelds at theNorth Carolina State UniversityÕs Clinton ResearchStation and near McGee Crossroads, NC. For the sec-ond 2001 experiment (NC-2001-B), traps were placedin sweetpotato Þelds near McGee Crossroad, at theClinton Research Station, and near Newton Grove,NC. For the third 2001 experiment (NC-2001-C), trapswere set up in sweetpotato Þelds near AtchinsonÕsMill, Hares Crossroads, and Narron, NC. In 2002, trapswere placed in sweetpotato Þelds near Heßin, StancilsChapel, and Kenly, NC.

In 2001, an experiment with Pherocon CRW kairo-mone traps (new design) was set up in randomizedcomplete block design with all seven Trece lures in asweetpotato Þeld at the Louisiana State UniversityAgCenter, Burden Research Plantation in BatonRouge, LA (Table 1). Traps were checked three times.Lures were changed and assigned randomly to newÞeld positions on 29 July. Typical cultural practices forsweetpotatoes were used during this trapping period,but no insecticides were applied during this period.Lure Aging Experiment, 2001. To be useful for

integrated pest management (IPM) programs, semio-chemical lures must be active in the Þeld for an ex-

tended period of time, and the controlled release ofactive components is critical to the lureÕs functionality(Kydonieus and Beroza 1982). Eventually, most of theattractant chemicals are released from the lure and itbecomes ineffective. Therefore, in 2001, we con-ducted an experiment to test the effective Þeld lifeof one of the Trece kairomone lures (TRE8276). On11 June, lures were opened (foil seal removed) andclipped hanging downward (in the same position asthey would be if attached to a trap) to an unshelteredrack in a Þeld at the USVL. Lures were aged in this wayfor 0, 1, 2, 3, 4, 5, 6, 7, or 8 wk. Weather conditions weretypical for late June and July in Charleston, with hot,humid, sunny, or partly cloudy days, and occasionalbrief afternoon rain showers. Minimum temperatures-maximum temperatures and rainfall for the lure-agingperiod were week 1, 21Ð34�C, 28 mm; week 2, 17Ð32�C,104 mm; week 3, 20Ð32�C, 20 mm; week 4, 21Ð33�C,66 mm; week 5, 18Ð36�C, 5 mm; week 6, 18Ð34�C,20mm;week7, 21Ð32�C,107mm;andweek8, 19Ð31�C,8mm.After lureshadbeenaged, theywereplaced intozip-lock plastic bags and put into a freezer (�17.8�C)until needed for the Þeld experiment. On 6 August,Pherocon CRW kairomone traps (new design) wereplaced into the three USVL cucurbit Þelds describedabove. Lures of each age classiÞcation were set up ina randomized complete block design with each Þeldrepresenting one replication. Traps were checked andbeetles were removed nine times (every 2Ð3 d) overa 3-wk period. After the Þrst week and after the second

Fig. 1. Average number of spotted, striped, and banded cucumber beetles captured in Pherocon CRW traps baited withkairomone lures (Trece, Inc.) for 4 yr (2000Ð2003) in cucurbit Þelds at the U.S. Vegetable Laboratory, Charleston, SC.


week, a new 0-age lure was added to the experiment,and the age designations of the lures in the Þeld wereadvanced 1 wk. Therefore, by the end of the experi-ment, there were 10 lure-age treatments in each ÞeldclassiÞed as 0Ð1, 1Ð2, 2Ð3, 3Ð4, 4Ð5, 5Ð6, 6Ð7, 7Ð8, 8Ð9,and 9Ð10 wk old. Data were analyzed by ANOVA, anddifferencesamongmeans(adult cucumberbeetlespertrap per day) for the lure-age treatments were sepa-rated by FisherÕs LSD test (P � 0.05) (SAS Institute1989).

Results

Experiments at the USVL, Charleston, SC. Overthe 5-yr period (1999Ð2003) of experiments at theUSVL, 2,661 spotted cucumber beetles were capturedin cucurbits and 1,780 were captured in sweetpota-toes. The kairomone trapping system evaluated in thisstudy clearly delineated population peaks of spottedcucumber beetles in cucurbit and sweetpotato Þelds.In 2001, 2002, and 2003, there were three distinctpopulation peaks of spotted cucumber beetles in cu-curbits (Fig. 1) and sweetpotatoes (Fig. 2) at theUSVL. In 2000, the midsummer peak of spotted cu-cumber beetles was indistinct (Figs. 1 and 2). Thisobservation of three population peaks for spotted cu-cumber beetles is in agreement with studies fromother locations with similar warm climatic conditions

as coastal South Carolina (Arant 1929, Michelbacheret al. 1955).

TRE8276 and TRE8336 were the best attractantlures for spotted cucumber beetles in this study(Tables 2 and 3). TRE8274, a dilution of TRE8276, wasalso effective. In cucurbits, TRE8276-baited traps cap-tured signiÞcantly more spotted cucumber beetlesthan did TRE8336-baited traps in 2002, but these twolures worked equally well the other years (Table 2).In sweetpotato, TRE8336-baited traps captured sig-niÞcantly more spotted cucumber beetles than didTRE8276-baited traps in 2000, 2001, and 2003, butthese lures worked equally well in 1999 and 2002(Table 3). None of the other Trece lures were veryeffective for trapping spotted cucumber beetles(Tables 2 and 3).

Over the 5-yr period, 1,047 striped cucumber bee-tles were captured in cucurbit Þelds at the USVL. Nodistinct population peaks of striped cucumber beetleswere seen in 2000. However, in 2001, there was a singlelarge population peak in August (Fig. 1). In 2002 and2003, there were distinct population peaks of stripedcucumber beetles in cucurbits in July and in August(Fig. 1). Each year, TRE8276-baited traps capturedsigniÞcantly more striped cucumber beetles than didtraps baited with the other lures (Table 2). As ex-pected, few (58) striped cucumber beetles were cap-tured in sweetpotato Þelds at the USVL. Most of these

Fig. 2. Average number of spotted and banded cucumber beetles captured in Pherocon CRW traps baited with kairomonelures (Trece, Inc.) for 4 yr (2000Ð2003) in sweetpotato Þelds at the U.S. Vegetable Laboratory, Charleston, SC.


Table 2. Season-long captures of three species of cucumber beetles by seven Trécé kairomone lures in Pherocon CRW kairomonetraps in fields of mixed cucurbits, Charleston, SC, 1999–2003

LureBeetles/trap/da

1999 2000 2001 2002 2003 Avg

Spotted cucumber beetle

TRE8274 0.245ab 0.247bc 0.246TRE8275 0.078b 0.045d 0.062TRE8276 0.377a 0.498a 0.462a 0.267a 0.833a 0.487TRE8279 0.042c 0.044d 0.106b 0.052c 0.059b 0.051TRE8282 0.250ab 0.065d 0.154b 0.088c 0.178b 0.147TRE8336 0.375a 0.372ab 0.536a 0.188b 0.631a 0.420TRE8391 0.108cd 0.051c 0.142b 0.100

Banded cucumber beetle

TRE8274 0.108 nsb 0.027abc 0.068TRE8275 0.147 0.017bc 0.082TRE8276 0.236 0.059a 0.034a 0.022 ns 0.002 ns 0.071TRE8279 0.158 0.042ab 0.009b 0.017 0.012 0.048TRE8282 0.108 0.003c 0.011b 0.011 0.000 0.027TRE8336 0.092 0.021bc 0.002b 0.012 0.000 0.025TRE8391 0.037abc 0.018 0.002 0.019

Striped cucumber beetle

TRE8274 0.146ab 0.166b 0.156TRE8275 0.000b 0.007c 0.004TRE8276 0.237a 0.377a 0.569a 0.131a 0.184a 0.300TRE8279 0.000b 0.003c 0.047b 0.010b 0.022b 0.016TRE8282 0.000b 0.012c 0.031b 0.012b 0.059b 0.023TRE8336 0.017b 0.030c 0.087b 0.019b 0.054b 0.041TRE8391 0.013c 0.010b 0.021b 0.015

aFor each beetle species, means within columns followed by a common letter are not signiÞcantly different (P� 0.05, FisherÕs least signiÞcantdifference).b NonsigniÞcant F value in AVOVA.

Table 3. Season-long captures of three species of cucumber beetles by seven Trécé kairomone lures in Pherocon CRW kairomonetraps in sweetpotato fields, Charleston, SC, 1999–2003

LureBeetles/trap/da

1999 2000 2001 2002 2003 Avg

Spotted cucumber beetle

TRE8274 0.033 nsb 0.151b 0.092TRE8275 0.000 0.029c 0.015TRE8276 0.059 0.159b 0.394b 0.204a 0.374b 0.238TRE8279 0.025 0.020c 0.117c 0.046b 0.048c 0.051TRE8282 0.067 0.057c 0.148c 0.060b 0.106bc 0.088TRE8336 0.125 0.276a 0.592a 0.154a 0.764a 0.382TRE8391 0.040c 0.023b 0.097bc 0.053

Banded cucumber beetle

TRE8274 0.133 ns 0.022 ns 0.078TRE8275 0.150 0.003 0.077TRE8276 0.000 0.019 0.012 ns 0.008 ns 0.000 ns 0.008TRE8279 0.275 0.014 0.016 0.007 0.003 0.063TRE8282 0.292 0.003 0.004 0.002 0.000 0.060TRE8336 0.067 0.037 0.015 0.001 0.000 0.024TRE8391 0.029 0.004 0.000 0.011

Striped cucumber beetle

TRE8274 0.000 ns 0.003 ns 0.002TRE8275 0.000 0.000 0.000TRE8276 0.050 0.003 0.057a 0.000 ns 0.000 ns 0.022TRE8279 0.000 0.000 0.011b 0.000 0.003 0.003TRE8282 0.000 0.000 0.010b 0.000 0.000 0.002TRE8336 0.000 0.000 0.014b 0.002 0.000 0.003TRE8391 0.000 0.002 0.000 0.001

aFor each beetle species, means within columns followed by a common letter are not signiÞcantly different (P� 0.05, FisherÕs least signiÞcantdifference).b NonsigniÞcant F value in AVOVA.


were captured in 2001, probably due to the proximityof this sweetpotato Þeld to a cucurbit Þeld (Table 3).

Only 218 banded cucumber beetles were capturedin cucurbits and 124 were captured in sweetpotatoesat the USVL, 1999Ð2003. No distinct population peaksof banded cucumber beetles could be determined forany year at the USVL. Adult banded cucumber beetleswere sometimes seen on sweetpotato and cucurbitfoliage, but many more spotted cucumber beetleswere observed. For the most part, there were no sig-niÞcant differences among lure treatments for season-long captures of banded cucumber beetles (Tables 2and 3). However, in 2000 and 2001 in cucurbits,TRE8276-baited traps captured signiÞcantly morebanded cucumber beetles than did traps baited withthe other lures (Table 2). Nevertheless, none of thekairomone lures distinguished itself as being espe-cially attractive to banded cucumber beetles. Giventhis speciesÕ habit of frequenting cucurbit ßowers,additional ßoral volatiles should be evaluated for usein Pherocon CRW kairomone traps.Experiments at Other Locations. The spotted cu-

cumber beetle was the dominant species captured(6,190 beetles) by using kairomone-baited PheroconCRW traps in sweetpotato Þelds at all locations in2000Ð2002 (Table 4). Only 19 banded cucumber bee-tles were captured at the Louisiana location, and asingle banded cucumber beetle was collected at theEhrhardt, SC, site in 2002. Only 11 striped cucumberbeetles were captured at two sites in North Carolinain 2000 and 2001 (Table 4).

For these locations, the effectiveness of the sevenTrece lures for capturing spotted cucumber beetleswas similar to the results from sweetpotato at theUSVL (Tables 3 and 4). Overall at locations other thanthe USVL, TRE8336-baited traps captured more spot-ted cucumber beetles than did TRE8276-baited trapsin three of 11 experiments, but these lures wereequally effective in the other eight experiments insweetpotatoes (Table 4).TrapDesign.Overall, the new version of the Phero-

con CRW kairomone trap was an improvement overthe old design (Fig. 3). Increased trap captures withthe new trap design were most evident for stripedcucumber beetles in cucurbits, where treatment dif-ferences were signiÞcant both years (2001, P� 0.005;2002, P � 0.013). For spotted cucumber beetles in2001, the new trap design captured signiÞcantly morebeetles than the old design in cucurbits (P � 0.004)and sweetpotatoes (P� 0.030). However, there wereno signiÞcant differences in trap design in 2002 forspotted cucumber beetles (cucurbits, P � 0.095;sweetpotatoes, P � 0.349) (Fig. 3). The new trapdesign caught signiÞcantly more banded cucumberbeetles in cucurbits (P � 0.017), but not in sweet-potato (P � 0.937) in 2002. Trap design was not sig-niÞcant for banded cucumber beetles in either cucur-bits (P � 0.854) or sweetpotato (P � 0.263) in 2001(Fig. 3).Lure Aging Experiment, 2001. The age of TRE8276

lures signiÞcantly affected trap captures of spotted(F11,219 � 7.8; P � 0.0001) and striped (F11,219 � 8.2;

Tab

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P� 0.0001) cucumber beetles in mixed cucurbit plant-ings (Table 5). However, there were no signiÞcantdifferences (F11,219 � 1.6; P � 0.095) in captures ofbanded cucumber beetles, which were trapped in lownumbers (22 beetles) during the 3 wk of this exper-iment. TRE8276 lost considerable activity after 2 wk of

aging in the Þeld, but after 10 wk, some beetles werestill captured in the Pherocon CRW kairomone traps(Table 5). Although more striped cucumber beetles(321 beetles) were captured during this experiment,the effects of lure age affected spotted (76 beetles) ina similar manner. This experiment clearly shows thatTRE8276 lures should be changed at least every 2 wkin the Þeld.

Discussion

Pherocon CRW traps baited with Trece kairomonelures provide an effective trapping system for spottedand striped cucumber beetles in cucurbits and sweet-potatoes. Because TRE8274, TRE8276, and TRE8336have 1,2,4-trimethoxybenzene and trans-cinnama-ldeyde in common, their attractiveness to spotted andstriped cucumber beetles is most likely due to thesecomponents. This result is consistent with previousreports on the attraction of spotted and striped cu-cumber beetles (Metcalf and Lampman 1989a, 1991).However, none of the other four Trece lures hadeither 1,2,4-trimethoxybenzene or trans-cinnama-ldeyde, and they were much less attractive to cucum-ber beetle adults.

The new design of the Pherocon CRW trap was animprovement over the original design. Because morethan one improvement was made to the trap design, itwas not possible to determine exactly why the newtrap worked better than the old one. However, it islikely that the new yellow bottom had more of aneffect on trap captures than did the slight improve-ment in shape of the top portion of the trap. This isbecause yellow attracts cucumber beetles from a dis-tance (Hesler and Sutter 1993), whereas the improvedtop portion acts simply to direct the beetles to theinsecticide bait.

Because larvae are seldom found when sweetpotatoroots are dug, injury often cannot be attributed to aparticular cucumber beetle species. In fact, it is even

Fig. 3. Cumulative numbers of spotted, striped, andbanded cucumber beetles captured by two versions (old andnew) of Pherocon CRW traps (Trece, Inc.) in cucurbit andsweetpotato Þelds at the U.S Vegetable Laboratory, Charles-ton, SC, 2001Ð2002.

Table 5. Captures of three species of cucumber beetles inPherocon CRW kairomone traps baited with TRE8276 kairomonelure that had been aged for 0–10 wk before being evaluated in fieldsof mixed cucurbits, Charleston, SC, 2001

Age oflure (wk)

Beetles/trap/d � SEa

Stripedcucumber

beetle

Spottedcucumber

beetle

Bandedcucumber

beetle

0Ð1 3.09 � 0.83a 0.72 � 0.22a 0.11 � 0.07 nsb

1Ð2 2.94 � 1.04a 0.35 � 0.09b 0.09 � 0.062Ð3 0.60 � 0.14b 0.24 � 0.06bc 0.04 � 0.033Ð4 0.49 � 0.14b 0.08 � 0.04d 0.02 � 0.024Ð5 0.36 � 0.17b 0.10 � 0.05cd 0.01 � 0.015Ð6 0.47 � 0.15b 0.16 � 0.05cd 0.10 � 0.056Ð7 0.22 � 0.07b 0.06 � 0.03d 0.00 � 0.007Ð8 0.20 � 0.07b 0.09 � 0.04d 0.07 � 0.048Ð9 0.15 � 0.06b 0.02 � 0.02d 0.02 � 0.029Ð10 0.06 � 0.04b 0.04 � 0.03d 0.00 � 0.00

a For each beetle species, means within columns followed by acommon letter are not signiÞcantly different (P� 0.05, FisherÕs leastsigniÞcant difference).b NonsigniÞcant F value in AVOVA.


difÞcult to determine whether root injury was causedby wireworms (W), cucumber beetles (Diabrotica sp.[D]), or ßea beetles (Systena sp. [S]), so damage bythese pests is often lumped into a single categorycalled WDS (Schalk et al. 1991). Observed WDS dam-age could be due to one or more of these pests, in-cluding a combination of cucumber beetle species.Clearly, additional tools, such as Pherocon CRW kai-romone traps that could identify what cucumber bee-tles were present, would be useful for making pestmanagement decisions for sweetpotatoes.

Elsey (1988) found that the spotted cucumber bee-tle was numerically dominant over the banded andstriped cucumber beetle in coastal South Carolina.The spotted cucumber beetle was also the most abun-dant species trapped in cucurbits and sweetpotatoes atthe USVL during the current study. This species alsopredominated at all of the upstate South Carolina andNorth Carolina sweetpotato locations we examined.

Our data support the conclusions of Houser andBalduf (1925) that the striped cucumber beetle isprimarily a pest of cucurbits, and not sweetpotatoes.This species is not listed as a pest of sweetpotatoes instate sweetpotato production guides, except for Vir-ginia (Tuckey et al. 2001). We captured few stripedcucumber beetles in sweetpotatoes, although sizablepopulations were trapped nearby in cucurbits. Thestriped cucumber beetle is a more-northerly species,and it is found in fewer numbers in the southeasternUnited States (Houser and Balduf 1925, Isley 1927).

Our data also show that the banded cucumber bee-tle is not present in the major sweetpotato productionareas of North Carolina and South Carolina and thatthe primary cucumber beetle pest on sweetpotato inthese areas is the spotted cucumber beetle. However,the banded cucumber beetle is a major pest of sweet-potatoes in Louisiana (Pitre and Kantack 1962). Since�1900, the banded cucumber beetle has expanded itsrange from southern Texas eastward to the coastalareas of the Gulf of Mexico and Atlantic Ocean up toNorth Carolina (Smith 1966, Krysan and Branson1983). Banded cucumber beetles are generally con-Þned to the coastal areas of South Carolina, usually no�65 km from the coast (Clemson University 2003).However, the distribution map published by Krysan(1986) shows banded cucumber beetles distributedfurther inland in South Carolina and North Carolina.

The northern range of the banded cucumber beetleis limited to areas where hard freezes seldom occur(Saba 1970; Krysan 1986; Elsey 1988, 1989). Saba(1970) reported that banded cucumber beetle adultsdied after two days exposure to �6�C. However, Elsey(1989) showed that although banded cucumber bee-tles are intolerant of severe freezing conditions, adultscan become acclimatized to cold temperatures andsurvive in subfreezing temperatures. Elsey (1988)showed that banded cucumber beetle adults are Þrstcaptured in the Charleston, SC, area in early April,shortly after the last freezing temperatures. This spe-cies would not have had time to produce a new gen-eration of adult beetles by early April, which couldindicate that at least some adults survive in the

Charleston area, even though Charleston averages32Ð33 d with freezing temperatures each year (Anon-ymous 1968, 2004). Therefore, the movement ofbanded cucumber beetles away from coastal areaswhere they survived a mild winter could impactsweetpotato growers some distance inland. A betterkairomone for baiting Pherocon CRW traps couldhelp further delineate the distribution of this pest inthe southeastern United States.

Acknowledgments

We thank Jennifer Cook and Louise Cauthen, U.S. Veg-etable Laboratory, Charleston, SC, for excellent technicalsupport. All kairomone lures and Pherocon CRW traps wereprovided by Trece, Inc.

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Received 16 March 2004; accepted 18 October 2004.


h e monitoring cucumber beetles in sweetpotato and

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