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HOUSEHOLD ANDSTRUCTURALPESTS

Evaluation of Attractants for Monitoring Populations of the GermanCockroach (Dictyoptera: Blattellidae)

GODFREY NALYANYA AND COBY SCHAL

Department of Entomology, North Carolina State University, Raleigh, NC 276957613

J. Econ. Entomol. 94(1): 208214 (2001)

ABSTRACT Lures that are used to attract German cockroaches,Blattella germanica(L.), to trapswere compared in olfactometer assays in the laboratory and in trapping experiments in cockroach-infested homes anda swine farm. In olfactometerassays,AgriSense GP-2 was themost attractive lure,followed by peanut butter, and distillers grain. Other lures, including Trapper tablet; Victorpheromone, a crude fecal extract that ostensibly containsB. germanica aggregation pheromone; andVictor food lure elicited upwind orientation from50% of the test insects. Peanut butter anddistillers grain were equally attractive in trapping experiments in swine production barns and theycaptured signicantly more cockroaches than the GP-2 tablet or the Victor pheromone lure; thecommercial lures failed to attract signicantly more cockroaches than the unbaited control traps.When tested against blank controls, cockroaches preferred to rest in shelters that contained theaggregation pheromone-based lure (Victor), but this lure was the least attractive to cockroaches inolfactometer assays. These results do not support claims that commercial crude fecal extracts attractcockroaches to traps, and they highlight a need for developing more attractive lures for detectionof cockroaches and for monitoring populations.

KEY WORDS Blattella germanica,attractants, baits, traps

MANY REDUCED-RISK APPROACHES for controlling cock-roach populations are based on attracting cockroachesto traps, insecticide baits, and onto surfaces treatedwith residual insecticides or biological control agents.Pest monitoring, a central tenet of integrated pestmanagement (IPM), also requires the deployment ofeffective attractants. Yet, commercially available trapsand baits have a limited range over which they attractcockroaches and probably fail to lure cockroaches outof deep refuges. Trap catch therefore may be highlydependent upon location effects within an infestedstructure, and traps may fail to accurately estimate theprevailing cockroach population (Wright and Dupree

1983, Ross and Brett 1989, Schal and Hamilton 1990,Nalyanya 1995).

Extensive research has been conducted on food-based lures and synthetic odorants with the aim toincorporate them into cockroach control tactics.Among them, fatty acids and esters (Tsuji 1966,Wileyto and Boush 1983), cyclohexylalkanoates andn-alkylcyclohexaneacetates (Sugawara et al. 1975,Wileyto and Boush 1983), and tetrahydropyran esters(Pandey et al. 1994, 1995) have been evaluated. Inaddition, various household food materials (Tsuji1965, Reierson and Rust 1977, Rust and Reierson 1981,

Ballard andGold 1982) and insecticide baits (Rust andReierson 1981, Scharf et al. 1994; G.N. unpublisheddata) have been evaluated for attractiveness to cock-roaches. Most of these lures, however, are mediocreattractants, and results from these studies have beeninconsistent.

The lack of effective attractants for deployment intraps is probably the single most signicant factorcontributing to the abandonment of population mon-itoring efforts and a reliance on scheduled applica-tions of insecticides in indoor environments. Effectiveattractants and efcient traps should increase the re-active distance of cockroaches to traps and thus en-hance the sensitivity and accuracy of the decision-making component of IPM programs. We conductedlaboratory behavioral assays and trapping experimentsin the eld to evaluate the relative attractiveness ofseveral lures that are used in cockroach traps.

Materials and Methods

Insects. An insecticide-susceptible strain ofBlattellagermanica (L.), originally obtained from AmericanCyanamid (Princeton, NJ), was used in these exper-iments. The colony was maintained at 27 1C, am-bient relative humidity, and a photoperiod of 12:12(L:D) h, and was provided with water and Purina RatChow No. 5012 (Purina Mills, St. Louis, MO) ad libi-tum. In the laboratory tests, 10- to 20-d-old adult malecockroaches were used.

Lures and Traps.The following lures that are com-

mercially used in cockroach traps were evaluated:Trapper (Bell Laboratories, Madison, WI), Victorpheromone and food lures (Woodstream Industries,Lilitz, PA), and the GP-2 tablet (AgriSense, Fresno,CA). Several other foods were evaluated for attrac-tiveness, including peanut butter (Jif, Proctor and

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Gamble, Cincinnati, OH), distillers grain (JackDaniels, Lynchburg, TN), and feces-contaminated l-ter paper, containingB. germanica aggregation pher-omone (Ishii 1970).

The trapping efcacy of the Trapper and Victortraps was compared. The Trapper trap is tent-shaped(18.5 cm long, 4 cm wide, 4 cm high) with 120 cm2

of adhesive surface area. In our eld experiments, thistrap was divided into its three component traps (40cm2 each) and used with the supplied (170 mg at-tractant tablet, Trapper lure). The Victor pheromoneroach trap is box shaped (14 cm long, 10 cm wide, 1.7cm high) with 77 cm2 of adhesive surface area anda laminated plastic pheromone dispenser glued to theceiling of the trap. According to the manufacturer, thelure contains the crude extract of cockroach feces,containing B. germanica aggregation pheromone(Gerhet and Chang 1997).

Laboratory Experiments. Choice of Shelter Assays.

Two different assays were designed to investigate theshort-range attraction of cockroaches to the phero-mone dispenser in the Victor trap and its arrestantactivity. In the arrestant assay,we predicted that cock-roaches would preferentially rest in shelters contain-ing the aggregation pheromone. Shelters were madeof inverted paper Dixie cups (Norwalk, CT, 9.5 cmdiameter, 5 cm high, Steltenkamp et al.1992) with fourholes cut in the rim to facilitate entry. Victor phero-mone lures were taped to the inside bottom of the cupsand control cups received blank dispensers (gift fromWoodstream). The control and pheromone cups were

inverted symmetrically at opposite ends of a plasticcage (55 cm long, 30 cm wide, 21 cm high) the innerwalls of which were treated with a thin layer of pe-troleum jelly to prevent cockroaches from escaping.Twenty adult male German cockroaches were re-leased in the cage andprovided with water and a pelletof rat chow. Their choice of shelter was recorded 24 hlater, in the photophase, and the experiment was rep-licated ve times.

A second shelter preference assay investigated theattraction of cockroaches to a Victor pheromone dis-penser that they could not contact. The pheromoneand a blank control dispenser were placed in petri

dishes (60 mm diameter, 15 mm high) with ne-meshscreened lids to prevent cockroaches from contactingthe dispenser. The petri dishes were placed underDixie cups at opposite ends of a cage and the choiceof shelter was recorded after 24 h.

Olfactometer Assays. Two-choice olfactometerswere used for the behavioral assays. Each olfactom-eter consisted of a Plexiglas tube (54.5 cm long, 3.2 cmi.d.) with a 15 cm long divider sealed vertically in theupwind end. A tube cage (15 cm long, 3.2 cm i.d.) witha swivel metal screen gate was used to introduce cock-roaches into the downwind end of the olfactometer.

Sixteen such olfactometers were connected to a vac-uum pump that provided a linear air velocity of 25cm/s through each tube; the tubes were exhaustedoutside the building. Fluorescent lights covered withred photographic safety lters placed 60 cm belowandabove the olfactometers facilitated observation in the

dark. Lures were aired in the fume hood for 1520 minbefore dispensing 350 mg of each lure into a 0.5-mlmicrocentrifuge tube (cut to a height of 1.5 cm) thatwas used to dispense the odorants in the olfactom-eters. A strip of the Victor pheromone dispenser (1 by3 cm) was cut and t into each tube. A single malecockroach was placed in each olfactometer for 30min to acclimate to the airow and thereafter onlyquiescent insects were used in the assays. Thegate wasopened carefully and the lure introduced upwind. Aresponse was considered positive when the test insectran out of the cage to the end of the olfactometer tube,contacting the lure within 3 min. Each dispenser wasused in no more than seven assays to preserve theconcentration of odorants. A fresh nave male wasused in each assay and each kind of lure was testedseparately on 2530 cockroaches. The olfactometertubes were subjected to the 25 cm/s airow betweentests and cleaned with detergent solution every 4 5 d.

Field Experiments. Two trapping studies were con-ducted, one in apartments managed by the RaleighHousing Authority and the second in the farrowingbarns of a cockroach-infested swine farm in SampsonCounty, NC.

Trapping in Apartments. To select apartments, seventraps (six in the kitchen and one in the bathroom)were placed in each apartment for 48 h. Apartmentswith a minimum of 20 cockroaches were selected forthe study and only those locations within the apart-ments where cockroaches were trapped were used inlater studies. Traps and lures were compared using a

paired trapping procedure.A pairof traps (control andbaited traps) were placed in each of the sites, 0.5 mapart. After 24 h the traps were retrieved and replacedwith fresh traps, which were also left in place for 24 h.To minimize the effect of trap location, the positionsof the new traps were reversed relative to the previousday.

The rst experiment was conducted to determinethe attractiveness of the Trapper lure, so Trapper trapwith and without its lure were compared (six apart-ments). The second experiment was designed to eval-uate the attractiveness of the Victor pheromone lure.Because Victor traps without dispensers could not be

obtained, the pheromone dispenser from the Victorpheromone trap was carefully removed from the trap,stapled to the inner surface of a Trapper trap andcompared with an unbaited Trapper trap (12 apart-ments). The third experiment compared the attrac-tiveness of GP-2 tablet and Victor pheromone lure (12apartments) using a similar procedure as in experi-ment 1. Because GP-2 is formulated as a 750-mgtablet, the tablet was carefully divided into pieces of170 mg, comparable to the weight of Trapper tablets.Finally, the Victor roach trap with its pheromone lurewas compared with the Trapper trap with its tablet (14

apartments). Each site within each apartment wasconsidered a replicate, and a minimum of 30 replica-tions were made per comparison.

Trapping in the Swine Barns. To investigate the at-tractiveness of the AgriSense GP-2 tablet, the Victorpheromone lure, and several gel baits, trapping was

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conducted in rooms (15 m long, 13 m wide) in far-rowing barns of a cockroach infested swine farm. Ma-son jars (0.95 liter) wrapped in a 12-cm cotton sleeveto facilitate cockroach entry, were used as traps. Pe-troleum jelly was spread on the inner lip to preventcockroaches from escaping. Traps were baited with asingle GP-2 tablet (750 mg), Victor pheromone (1dispenser), 750 mg of peanut butter, or 750 mg ofdistillers grain. Lures were placed in small petri dishes(60 mm diameter, 15 mm) with screened lids whichpermitted odorant emission while preventing trappedcockroaches from contacting the lures. The experi-mental design was a randomized complete block.Baited traps were deployed for 24 h 1 m apart on theoor along the cockroach-infested walls of farrowingrooms. Trap location was randomized along each wall(block) and 10 walls were used concurrently.

Data Analysis.Olfactometer assays were analyzedby Ryans test. Assays for choice of shelter in two-

choice preference assays were analyzed by the chi-square test at 0.05. Students paired t-test was usedto analyze trap catches of paired traps in apartments.Because few adult males and females were captured inapartments, the sum of all cockroaches (males, fe-males, and nymphs) per trap was used in the analysesand only trapping sites with a mean trap catch of twoor more cockroaches per day were included in theanalysis. Mean trap catch per location per day for eachtrap type or trap and lure combination was computed.Trap counts from the pig farm were square-root trans-formed and analyzed by PROC GLM (SAS Institute

1996). LSMEANS/PDIFF was used to compute themean trap catch and to separate differences among thelures. The data analyzed from the swine farm is asubset of a larger data set from a study in which otherlures were evaluated. To use this subset of data, cor-relation coefcients of trap catch between the controland lures, and between pairs of lures were computed.The correlation coefcients were highly variable andmostly insignicant suggesting that there was poorassociation among the various treatments.

Results

Shelter Preference Assays.In the contact arrestantassays, all the cockroaches preferred the shelter con-taining the Victor pheromone lure over the controlshelter (2 10.0, df 1, P 0.001) (Fig. 1A).Similarly, in short-range attractant assays, where thedispenser was screened from the cockroaches, signif-icantly more males preferred the shelter with the lurethan the control shelter (2 6.74, df 1,P 0.001)(Fig. 1B).

Olfactometer Assays. In the behavioral assays, up-wind orientation responses were highly dependentupon the presence of an attractantlure (Fig.2). All the

lures that were tested except Victor food and Victorpheromone were signicantly more attractive thanthe control (P 0.05).Of the commercial lures tested,the GP-2 tablet elicited the highest and signicantlymore responses (P 0.05) than any other commerciallure or cockroach contaminated lter papers. Male

response to peanut butter and distillers grain was notsignicantly different (P 0.05), but peanut butterattracted more cockroaches than all the commerciallures, except GP-2 (Fig. 2).

Trapping in Apartments. Paired assays in apart-

ments partly conrmedourolfactometer results. Trap-per traps with GP-2 lure captured signicantly morecockroaches than Trapper traps with Victor phero-mone (t 3.62, df 38,P 0.001) (Fig. 3C). How-ever, Trapper traps with or without the Trapper luretrapped equal numbers of cockroaches (t 0.77, df

Fig. 1. Arrestant and short-range attractant assays of theVictor pheromone dispenser. Columns represent mean SEnumber of cockroaches in each shelter. Chi-square test wasused to compare theresults to theexpected 50:50 distribution(n 5 replicates with 20 cockroaches each).

Fig. 2. Upwind orientation responses of male Germancockroaches to various lures in olfactometer assays. Columnswith different letters are signicantly different (Ryans test,P 0.05; n 2530 cockroaches per lure).

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32,P 0.05) (Fig. 3A), suggesting that the lure wasless effective in the eld. Similarly, there was no sig-nicant difference in trap catch between Trappertraps with or without the Victor pheromone lure (t1.72 df 79,P 0.05) (Fig. 3B), indicating that the

pheromone-based lure failed to enhance trap catch inthe eld. And nally, no signicant difference wasfound in the mean trap catch per day of Trapper trapscontaining the Trapper lure and the Victor trap con-taining the Victor pheromone lure (t 1.976, df 38,P 0.05) (Fig. 3D). However, apartments containedrelatively low cockroach populations and the hetero-geneity of kitchens delimited the paired traps to0.5m apart. We therefore expanded these assays to ahighly infested swine farm.

Trapping in Swine Barns. There were signicantdifferences in the number of cockroaches trapped by

different lures (F 30.16; df 4, 146;P 0.001) (Fig.4). The block by bait interaction was signicant (F4.67; df 32, 146; P 0.001), suggesting that cock-roach distribution was uneven among the barn wallsand along each of the walls. There were also signicantdifferences in the numbers of trapped males, females

and nymphs (F 24.69; df 8, 146; P 0.001), but thestage by bait interaction was not signicant (F 1.03;df8, 146; P 0.05). According to LSMEANS/PDIFFanalysis, only peanut butter and distillers grain con-sistently lured signicantly more males, females, and

nymphs than the blank control and other lures (Fig.4). The GP-2 lure also attracted and trapped morenymphs than the blankcontrol.In contrast, therewereno signicant differences between Victor pheromonelure and the blank control in their attractiveness tonymphs and adults.

Discussion

Several important inferences may be derived fromour data. First, the olfactometer results and the dif-ferential attractiveness of the baits in the eld indicate

that all stages of B. germanica perceive and orienttoward the source of food odors and other olfactorycues over some distance.These ndings are in contrastto the generally held view thatB. germanicadoes notrespond in a consistent fashion to any attractant andthat no lure attracts cockroaches from a distance of

Fig. 3. Trap catches of German cockroaches in apart-ments using two types of traps and three lures. Columnsindicate mean ( SE) (ns, not signicantly different byStudents pairedt-test,P 0.05).

Fig. 4. Comparativetrapcatches of German cockroachesin swine barns by using several lures. Columns indicatemean SE and those with different letters are signicantlydifferent from each other (LSMEANS, P 0.05; n 10).



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more than a few centimeters (Meisch and Howell1967, Rust and Reierson 1981, Owens 1995, Reierson1995). Second, not all lures used in cockroach controlare equally attractive, and indeed the most attractivebaits that we found were off-the-shelf foods (e.g.,peanut butter) and industrial by-products (distillersgrain). And third, claims of attractiveness attributed tosome commercial lures cannot be supported by eitherlaboratory or eld results.

Some of the lures that we tested were attractive tocockroaches in both olfactometer assays in the labo-ratory and in trapping assays in the eld. The stage bybait interaction was not signicant in the eld, indi-cating that any given lure was equally attractive toimmatures and adults. In laboratory olfactometer as-says AgriSenses GP-2 tablet, peanut butter, and dis-tillers grain elicited responses from 50% of adultmales. Trapper lure outperformed the blank controlbut was not signicantly more attractive than Victor

food or pheromone lures, which in turn failed to at-tract more cockroaches than the blank control (Fig.2). Peanut butter and distillers grain were highlyattractive to cockroaches in swine barns (Fig. 4),whereas GP-2 and Victor pheromone were not signif-icantly more attractive than the blank control. How-ever, GP-2 trapped signicantly more nymphs thanVictor pheromone lures and the control. Interestingly,the GP-2 lure was highly attractive in olfactometerassays and in apartments, but it failed to trap signi-cantly more adults than the blank control traps in theswine barns. We suspect that the use of mason jars as

traps probably favored trapping adults, as indicated bythe relatively fewer nymphs that were captured (Fig.4). In adhesive traps, which are more commonly de-ployed in pest control programs, more nymphs thanadults are generally captured, and we would thereforeexpect better efcacy from the GP-2 lure under theseconditions. Indeed this was the case as shown bytrapping results fromapartments where adhesive trapswere used. The difference in attractiveness of this lurecould also be due to strain differences in olfactoryresponse (G.N., unpublished data).

The data we obtained in apartments failed to dis-criminate between most of the paired treatments. The

Trapper trap was no better with the Trapper lure thanwithout it, and it was no better with the Victor pher-omone lure than without it (Fig. 3). Likewise, theTrapper trap plus its lure was no different from theVictor trap with its pheromone lure. In contrast, Trap-per trap with GP-2 was signicantly more attractivethan Trapper trap with Victor pheromone lure. Theseapartments contained relatively few cockroaches andthe relatively poor lures that were tested failed tocontribute signicantly to the traps. Nevertheless, re-sults with GP-2 suggest that a more potent lure canenhance trap catch.

Scharfet al.(1994) reported that GP-2 wasrepellentto cockroaches, in marked contrast to our nding thatit elicited responses from 90% of the males in olfac-tometer assays. Peanut butter elicited upwind re-sponses from 84%of the cockroaches in our assays, butRust and Reierson (1981) did not nd it to be as

attractive. Such discrepancies may be due to differ-ences in cockroach strains, lure formulations, and inmethodology for evaluating attractants. Although be-haviorally discriminating bioassays are required to dis-tinguish the component behaviors that result in a pos-itive response (Kennedy 1977, Sakuma and Fukami1985),most eld assays of lures, including ourown, arebased on trapping assays that fail to discriminate be-tween orientation behavior and trap efciency.

The Victor pheromone lure contains an extract ofGerman cockroach feces, which includes the aggre-gation pheromone. However, this lure was not signif-icantly more attractive to cockroaches than a blankcontrol in either olfactometer (Fig. 2) or in eld ex-periments (Fig. 4). It is important to note that toevaluate its attractiveness, we tested the lure inde-pendently of the Victor trap in which it is normallydeployed. The trap may be highly effective, but fromour data we must draw the conclusion, in contrast to

previous studies (Kaakeh and Bennett 1997a, 1997b),that high trap catch with Victor traps cannot be at-tributed to its aggregation pheromone lure. Also, rel-atively low responses to the Victor food lure (Fig. 2)would suggest that neither lure contributes substan-tially to the capture efciency of this trap.

The aggregation pheromone contains both attract-ant and arrestant components and cockroaches re-spond to the attractants by olfaction (Sakuma andFukami 1985) and to the arrestants by contact che-moreception (Sakuma and Fukami 1991). Although itis apparent that the Victor pheromone lure is only

minimally attractive in olfactometer bioassays, it ap-pears to function as a short-range attractant and ar-restant in cage bioassays (Fig. 1). Cockroaches clearlypreferred shelters that contained the pheromone lureover control shelters. Why do Victor pheromone dis-pensers fail to attract cockroaches? It is possible thatthe attractant components of the lure volatilize at ahigh rate or are lost during extraction, handling, orstorage. Indeed, Sakuma et al. (1997) caution of theextremely high volatility of several amines they iso-lated as cockroach attractant aggregation phero-mones. However, recent assays using a locomotioncompensator treadmill conclude that carboxylic acids,

both highly volatile and some less volatile compo-nents, elicit the attractant-aggregation response(Scherkenbeck et al. 1999). Clearly, more bioassay-guided chemical characterization is needed beforethese materials can be used in pest control operations.

Potent lures are needed to meet the challenges oftrapping cockroaches in detection, monitoring, andpest reduction efforts. The attractants and baits incurrent use are thought to be relatively ineffective(Ballard and Gold 1982, Gold 1995), yet some of thelures assayed in the eld signicantly increased trapcatch (Fig. 4) and presumably would enhance the

correlation between trap catch and infestation level.Moreover, similar studies with food-based baits (GN,unpublished data) and with pheromone- and food-baited traps(Liang et al.1998)conrm that some lurescan signicantly enhance trap catch. Clearly, cock-roaches respond to odorant lures from a distance.

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Food stuffs like peanut butter, bread, and distillersgrain, as well as various baits elicited more responsesin the laboratory, and trapped more cockroaches thandid unbaited traps in the eld. These ndings implythat even currently deployed lures can detect andmonitor infestations, and a concerted research effortto discover more potent attractants should yield moreeffective cockroach lures.

Tremendous advances are being made in the dis-covery and implementation of semiochemicals in IPMin eld crops (Jones 1998), forestry (Borden 1993),and stored products (Burkolder 1990, Phillips 1997).Notwithstanding, semiochemicals have not been usedin cockroach control, probably because of difcultiesin their characterization and identication. The sexpheromones of several species havebeen tested undereld conditions (Bell et al. 1984, Schal 1992, Liang etal. 1998), but have not been used in practical pestmanagement. Crude fecal extracts containing aggre-gation pheromones have shown promise in eld stud-ies aimed at reducing repellency of contact insecti-cides (Rust and Reierson 1977a, 1997b), and inlaboratory assays of baits (Miller et al. 1996), but theirdeployment as attractants appears to be less impres-sive (current study). Nevertheless, efforts to integratepheromone and food lures in cockroach populationmanagement are certain to optimize trap catch, aswithSupella longipalpa(Liang et al. 1998).

It appears that insecticide baits are relatively effec-tive cockroach control tactics, even when as few as 12bait stations are used per apartment (Reierson 1995).

It would seem that if traps were baited with attractivelures, if their trapping efciency were signicantlyenhanced, and if they were deployed in a similarmanner as baits, then theoretically, the efcacy of thetwo tactics should be the same. Assuming equallyeffective lures, both traps and baits should be encoun-tered by equal numbers of cockroaches, and the hor-izontal transmission of bait active ingredient (Silver-man et al. 1991; Kopanic and Schal 1997, 1999)notwithstanding, their efcacy should be similar. Thegreatdisparity between the efcacyof these two cock-roach control tactics highlights the need to better

understand the chemical ecology of the cockroach,develop better lures, and design traps that retain cap-tured cockroaches.

Acknowledgments

We thank C. Moore and B. Deasy for technical assistance;E. Vargo, C. Apperson, and M. Waldvogel for critically re-viewing the manuscript; and Woodstream Industries for sup-plying us with German cockroach fecal extracts, Victor ag-gregation pheromone dispensers, and blank dispensers. Thisresearch was supported in part by the Blanton J. Whitmire

Endowment at North Carolina State University, the USDA/PMAP (9804680), and the W. M. Keck Center for BehavioralBiology at North Carolina State University. G.N. also grate-fully acknowledges scholarships from the North CarolinaPest Control Association, the National Conference on UrbanEntomology, and the Rotary Club of N.W. Raleigh.

References Cited

Ballard, J. B., and R. E. Gold. 1982. The effect of selectedbaits on the efcacy of a sticky trap in the evaluation ofGerman cockroach populations. J. Kans. Entomol. Soc. 55:8690.

Bell, W., J. Fromm, A. R. Quisumbing, and A. K. Kydonieus.1984. Attraction of American cockroaches (Orthoptera:Blattidae) to traps containing periplanone B and to in-secticide-periplanone mixtures. Environ. Entomol. 13:448450.

Borden, J. H. 1993. Strategies and tactics for the use ofsemiochemicals againstforest insect pests in North Amer-ica, pp. 265279. In B. A. Leonhardt and J. L. Vaughn[eds.], Pest management: biologically-based technolo-gies. American Chemical Society, Washington, DC.

Burkolder, W. E. 1990. Practical use of pheromones andother attractants for stored product insects, pp. 497516.In R. L. Ridgeway, R. M. Silverstein, and M. N. Inscoe[eds.], Behavior-modifying chemicals for insect manage-ment, applications of pheromones and other attractants.

Marcel Dekker, New York.Gerhet, M. J., and F. N. Chang. 1997. Compositions con-

taining cockroach aggregation pheromones, their pro-duction and uses. U.S. Patent no. 5:665,370.

Gold, E. R. 1995. Alternate control strategies, pp. 325344.InM. K. Rust, D. A. Reierson, and J. M. Owens [eds.],Understanding and controlling the German cockroach.Oxford University Press, New York.

Ishii, S. 1970. An aggregation pheromone of the Germancockroach,Blattella germanica(L.) 2. Species specicityof the pheromone. Appl. Entomol. Zool. 5: 3341.

Jones, O. T. 1998. Practical applications of pheromones andother semiochemicals, pp. 261355. In P. Howse, I.Stevens, and O. Jones [eds.], Insect pheromones andtheir use in pest management. Chapman & Hall, London.

Kaakeh, W., and G. W. Bennett. 1997a. Evaluation of trap-ping and vacuuming compared with low-impact insecti-cide tactics for managing German cockroaches in resi-dences. J. Econ. Entomol. 90: 976982.

Kaakeh, W., and G. W. Bennett. 1997b. Evaluation of com-mercial sticky traps used for German cockroach (Dic-tyoptera: Blattellidae). J. Agric. Entomol. 14: 349353.

Kennedy, J. S. 1977. Behaviorally discriminating assays ofattractants and repellents, pp. 215229.In H. H. Shoreyand J. J. McKelvey, Jr. [eds.], Chemical control of insectbehavior: theory and application. Wiley, New York.

Kopanic, R. J., and C. Schal. 1997. Relative signicance of

direct ingestion and adult-mediated translocation of baitto German cockroach (Dictyoptera: Blattellidae)nymphs. J. Econ. Entomol. 90: 10731079.

Kopanic, R. J., and C. Schal. 1999. Coprophagy facilitateshorizontal transmission of bait among cockroaches (Dic-tyoptera: Blattellidae). Environ. Entomol. 28: 431438.

Liang, D., A. Zhang, R. J. Kopanic, Jr., W. L. Roelofs, and C.Schal. 1998. Field and laboratory evaluation of femalesex pheromone for detection, monitoring, and manage-ment of brownbanded cockroaches (Dictyoptera: Blat-tellidae). J. Econ. Entomol. 91: 480485.

Meisch, D. M., and D. E. Howell. 1967. An evaluation ofbaits for cockroaches. Pest Control 6: 16, 18, 20.

Miller, D. N., P. G. Koehler, and R. S. Patterson. 1996. In-uence of German cockroach (Dictyoptera. Blattellidae)fecal extract on food consumption and harborage choice.J. Econ. Entomol. 89: 668672.

Nalyanya, G. 1995. An evaluation of German cockroachmonitoring techniques. M.S. thesis, North Carolina StateUniversity, Raleigh.



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