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HOUSEHOLD AND STRUCTURAL Efficacy of Simulated Barrier Treatments Against Laboratory Colonies of Pharaoh

GRZEGORZ BUCZKOWSKI,1, 2 MICHAEL E. SCHARF,3 CATINA R. RATLIFF,1 1 AND GARY W. BENNETT

J. Econ. Entomol. 98(2): 485Ð492 (2005) ABSTRACT Five selected were applied to four substrates and evaluated in laboratory studies for repellency and toxicity against the Pharaoh ant, pharaonis (L.). We tested both repellent and nonrepellent formulations on outdoor (concrete and mulch) and indoor (ceramic and vinyl) substrates. Repellency was evaluated using a behavioral bioassay in which colonies were given a choice to leave the treated zone and move into empty nests provided in the untreated zone. We used a novel experimental design whereby walked on a Slinky coil suspended from a metal support frame, thus permitting a long foraging distance with a minimum use of space and resources. Cypermethrin, a repellent pyrethroid , resulted in colony budding, although the response was delayed. Toxicity of insecticides was evaluated as worker, queen, and brood mortality. The most effective treatment was Þpronil, which provided 100% reduction in pretreatment activity by 2 d posttreatment on both concrete and mulch. Chlorfenapyr was highly effective on both outdoor and indoor substrates. SigniÞcant substrate effects were observed with insecticides applied to nonabsor- bent substrates (ceramic tile), which performed better than insecticides applied to absorbent sub- strates (vinyl tile). Other highly absorbent materials (mulch and concrete), however, did not reduce insecticide efÞcacy. This is because ants relocated nests into and/or under these attractive nesting materials, thus increasing their exposure to toxic insecticide residues. Our results demonstrate efÞcacy of nonrepellent liquid insecticides as indoor treatments for the control of Pharaoh ants and possibly as exterior perimeter treatments.

KEY WORDS Monomorium pharaonis, Pharaoh ant, repellency, residual insecticide

THE PHARAOH ANT, Monomorium pharaonis (L.), is an may be initiated at any time. Traditionally, part of the introduced species that exhibits several tramp ant success and persistence of the Pharaoh ant had been characteristics such as generalist diet, polygyny, poly- attributed to its frequent budding habits (Edwards domy, large colony size, reproduction by budding, and 1986). Colony fragmentation may occur due to a wide close association with humans (Passera 1994). These range of biotic and abiotic factors; however, only lim- traits make colonies of Pharaoh ants successful invad- ited information exists on the factors responsible for ers of human built structures (Edwards and Baker inducing the budding behavior. Possible factors in- 1981, Edwards 1986) and also make them extremely clude overcrowding, response to weather (seasonal difÞcult to eradicate. Of the habits listed above, re- changes in a structureÕs central heating and cooling production by budding (sociotomy) is perhaps the system), physical disturbance, dietary change (deple- most critical to the success of Pharaoh ants. Although tion of or discovery of new resource), or chemical the majority of ant species disperse and start new disturbance (application of a repellent pesticide). colonies by individual females after a mating ßight, Traditionally, applications of repellent spray insecti- Pharaoh ants mate in the nest and colony reproduc- cides have been alleged to be the major factor pro- tion occurs by fragmentation of mature colonies (Ed- moting fragmentation of colonies (Green et al. 1954, wards 1986). During budding, a reproductively com- Lee et al. 1999). However, no detailed experimental petent colony fragment migrates to a new location to data exist to either support or refute this claim. In establish a new nest (Peacock et al. 1955, Vail and addition, residual insecticides are usually ineffective Williams 1994), which may or may not remain in for complete colony elimination because they only association with the parent colony. Thus, new colonies affect foraging workers and fail to reach reproductive are created independently of gyne production and individuals (Lee et al. 1999). Recently, new classes of nonrepellent pesticide 1 Department of Entomology, Purdue University, West Lafayette, chemistries have become available for urban con- IN 47907Ð2089. trol. The phenyl pyrazoles (Þpronil) and the struc- 2 Corresponding author, e-mail: [email protected]. 3 Entomology and Nematology Department, P.O. Box 110620, Uni- turally similar pyrroles (chlorfenapyr) are both non- versity of Florida, Gainesville, FL 32611Ð0620. repellent residual insecticides that have proven

0022-0493/05/0485Ð0492$04.00/0 ᭧ 2005 Entomological Society of America 486 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 98, no. 2

Fig. 1. Experimental design. Three plastic trays were connected by steel coils suspended from a metal support frame. The length of the coil between any two trays was 100 feet. Nest containing the colony was placed in the center tray and surrounded by treated surfaces. Alternative nesting trays were provided on either side. effective against a variety of pests of public and med- red acetate sheets to create the darkened conditions ical importance (Scott and Wen 1997, Ameen et al., preferred by the ants. Plastic spacers were inserted 2000). between the nest ßoor and the covering to restrict the The goals of this study were two-fold. Our main ants to a monolayer to facilitate counting of the various objective was to compare the efÞcacy of barrier treat- castes and developmental stages. When the nests were ments of reportedly nonrepellent insecticides (Þpro- observed to contain Ϸ20 queens, 80Ð90 1-cm2 grid nil and chlorfenapyr) to ones that are thought to be squares of brood (counted by overlaying a grid over repellent (synthetic pyrethroids). We hypothesized the nest and marking the outline of the brood), and that nonrepellent insecticides will be associated with Ϸ4,000 workers, single plates were transferred to the greater mortality via direct and indirect (horizontal center tray of the experimental conÞguration shown transfer) exposure to the active ingredient (e.g., Buc- in Fig. 1. zkowski and Schal 2001). In contrast, repellent treat- The experimental setup consisted of three 38 by ments will cause lower mortality due to colony relo- 50-cm Fluon-coated plastic trays connected by steel cation (budding) once the insecticide is detected. Our (Slinky, James Industries, Inc., Hollidaysburg, PA) second objective was to evaluate the efÞcacy of re- coils suspended from a metal support frame (Fig. 1). sidual insecticides on a variety of outdoor and indoor The length of the coil between any two trays was 100 substrates. Previous studies emphasized the impor- feet, and previous work in our laboratory has shown tance of surface type on insecticide efÞcacy (Chad- that Pharaoh ants will routinely traverse this distance wick 1985, Knight and Rust 1990, Osbrink and Lax without difÞculty (Ratliff and Bennett 2003). Empty 2002) with nonporous substrates generally providing nests were placed in trays adjacent to the center tray. better, longer lasting control. We examined four sub- To promote initial exploration of the center tray and strates likely encountered by foraging Pharaoh ants: the coils, food was provided mid-way (25 feet) along two outdoor substrates (concrete and mulch) and two the coils immediately adjacent to the center plate. indoor substrates (nonabsorptive ceramic and absorp- Food placements consisted of boiled egg yolk, peanut tive vinyl tile). oil, 10% sucrose, and whole cricket provided three times a week. After a 7-d acclimation period, treated substrates Materials and Methods were placed on three sides of the nest in the center Laboratory Pharaoh ant colonies were maintained tray, 3 cm from the nest. We tested two types of at constant temperature and humidity in an environ- outdoor substrates and two types of indoor substrates. mentally controlled rearing and testing room (25 Ϯ Outdoor substrates were concrete pavers (20 by 10 by 1ЊC, 65% Ϯ 10% RH). Colonies were reared in 38 by 4 cm in height) or hardwood mulch (Western White 50-cm Fluon-coated trays (DuPont Polymers, Wilm- Wood Mulch, Menards Inc., Eau Claire, WI; arranged ingtom, DE) on a regular diet of 10% sucrose, whole in aluminum foil trays 20 by 10 by 4 cm in height). cricket, peanut oil, and boiled egg yolk. Nests were Indoor substrates were ceramic (i.e., nonabsorptive plastic dishes Þlled with moist plaster. Experimental surface) and vinyl (i.e., absorptive surface) tile, both colony units were obtained by placing empty nests (9 10 by 10 cm by 0.5 cm in height. Five different liquid cm in diameter) in colony rearing trays for voluntary spray insecticides were examined (Table 1), although colonization by the ants. The nests were covered with not all insecticides were tested on all four substrates. April 2005 BUCZKOWSKI ET AL:COLONY BUDDING IN PHARAOH ANT 487

Table 1. Insecticides and active ingredients used in the study

Insecticide trade Active Application rate Substrate Insecticide class Manufacturer name ingredient (␮g/cm2) tested on Cinnamite Cinnamaldehyde Aldehyde Mycotech, Butte, MT 12,973.0 Ceramic, vinyl Demon EC Cypermethrin Type II pyrethroid Syngenta Crop Protection 3.0 Ceramic, vinyl Inc., Greensboro, NC Phantom Chlorfenapyr Pyrrole BASF Corp, Research 34.5 Concrete, mulch, Triangle Park, NC ceramic, vinyl Talstar F Bifenthrin Type I pyrethroid FMC Corp., Philadelphia, PA 2.6 Concrete, mulch Termidor SC Fipronil Phenylpyrazole Aventis Environmental 3.7 Concrete, mulch Science, Montvale, NJ

The insecticides were applied to the top surface of all Results and Discussion substrates by using a plant mister and allowed to dry Colony Movement. Workers frequently traversed for 4 h. The treatments remained in place for the entire the distance between nesting and alternative trays, length of the study. All insecticides were applied at and small groups of workers (50Ð100 individuals) oc- label rates speciÞed by the manufacturers and in equal casionally explored and/or colonized side nests. In volumes. Water-treated substrates were used as con- fact, all replicates in all treatments had small groups of trols. Three replicates of each treatment were per- workers nesting in alternative trays at some point formed, and assays were run for 6 wk (42 d). Obser- during the study. However, no queens and/or brood vations were made on day 2, 4, and 7 and subsequently were observed in side trays, with the exception of once a week until day 42. Collected data consisted of treatments involving Demon (cypermethrin; a repel- 1) the presence or absence of colony relocation be- lent type II pyrethroid). Reproductively competent haviors (deÞned as movement of reproductively com- colony fragments colonized alternative trays in three petent colonies into side trays), 2) worker foraging of three Demon replicates involving vinyl tile, and one activity, 3) worker mortality, 4) queen mortality, and of three Demon replicates involving ceramic tile. 5) brood mortality. We deÞne reproductively compe- However, colonies did not migrate to side trays until tent colonies as a group of 50 or more workers ac- day 35. It remains unknown why colonies showed this companied by at least one queen and/or brood (Pea- delayed budding behavior, long after the initial expo- cock et al. 1955). In the Pharaoh ant, queens are not sure to insecticide. Other researchers, too, reported essential for successful foundation of new colonies delayed budding with pyrethroid insecticides (Knight because workers can rear new reproductives from and Rust 1990). Perhaps budding occurs only after a existing brood (Vail and Williams 1994). Worker for- certain threshold level of colony mortality is reached. aging activity was monitored by counting foragers Worker numbers steadily declined from day 1 to day crossing a marked line on all four 50-foot sections of 35, and excessive mortality by day 35 may have sig- the Slinky coil. The marks were 20 feet from the pans. naled a need to abandon the original nesting site in The number of workers that crossed the mark in 5 min favor of one that offered greater protection from toxic was recorded, and all counts were done before the insecticides. Nest and/or colony relocation seems to ants were given food. Worker and queen mortality was be a common behavior in ant colonies exposed to toxic assessed by aspirating dead individuals from the pans chemicals. Imported Þre ant Solenopsis invicta (Bu- and counting them at the end of the 6-wk study. For ren) colonies relocated their nests after insecticide workers, results are normalized to an assumed starting applications (Collins and Callcott 1995) and Argen- number of 4,000 workers. Brood mortality was esti- tine ants, Linepithema humile (Mayr) avoided nesting mated by counting the number of 1-cm2 squares of in areas containing toxic aromatic cedar mulch brood at the beginning and end of the study. For (Meissner and Silverman 2003). Nest relocation may queens and brood results are normalized to starting help colonies survive insecticide treatments and may quantities. be partly responsible for the difÞculty in eradicating Statistical analyses were performed in SAS 8.1 (SAS established colonies. Other pyrethroid insecticides, Institute 2002) and consisted of analysis of variance including cypermethrin and permethrin, also have (ANOVA) (PROC ANOVA) on mean ant numbers by been shown to be repellent to ants (Knight and Rust substrate category. A separate ANOVA test was per- 1990). In temperate climates, Pharaoh ants nest mostly formed for each of the four parameters measured indoors; however, they also may forage outside in (foraging activity, queen mortality, worker mortality, warmer climates (Oi et al. 1994). Thus, repellent py- and brood mortality) and for each substrate category rethroid insecticides should not be used for eradica- (outdoor versus indoor). Thus, eight separate ANOVA tion of indoor colonies of Pharaoh ants. Such insecti- tests were performed: four for outdoor substrates and cides may, however, be useful in creating a repellent four for indoor substrates. Each ANOVA was followed and potentially effective barrier to exclude Pharaoh by the least signiÞcant difference (LSD) t-test to test ants from entering buildings in warmer climates. for signiÞcant differences between means. The level of Despite the relatively low incidence of budding into signiÞcance was set at ␣ ϭ 0.05. side trays, entire colonies were observed to readily 488 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 98, no. 2

Table 2. Summary of observations indicating migration from as counting the number of foragers crossing an imag- original nests into or under treated substrates inary line along a foraging trail (Knight and Rust 1990). Ants that reached the line had crossed the Insecticide Brick Mulch Ceramic Vinyl barrier and were thus not repelled. Although this Control (3/3) (3/3) (3/3) (3/3) methodology offers a direct assessment of repellency, Termidor SC (Þpronil) (0/0)* (0/0)* N/A N/A Talstar F (bifenthrin) (1/3) (0/3) N/A N/A its potential drawback is the underestimation of re- Phantom (chlorfenapyr) (2/3) (3/3) (2/3) (1/3) pellency due to rapid knockdown and/or mortality Cinnamite (cinnamaldehyde) N/A N/A (3/3) (3/3) caused by some insecticides. Knight and Rust (1990) Demon EC (cypermethrin) N/A N/A (1/3) (2/3) avoided this potential problem by normalizing counts to the number of ants surviving the treatments at each Number of replicates where relocation occurred is provided in parentheses (n ϭ 3). N/A, not applicable. count. This method requires that all surviving ants be * Rapid onset of mortality prevented assessment of colony move- counted at each sampling period. Therefore, it is labor- ment. intensive and may not be always practical, such as when colonies nest in mulch. An advantage of our methodology is that there is no need for exact worker relocate into mulch and under pavers, ceramic, and counts at each sampling period. A potential disadvan- vinyl tile (Table 2). Other tramp ant species also have tage is the assumption that lack of movement indicates been shown to relocate colonies into alternative sub- repellency. Lack of movement may only indicate pref- strates. Meissner and Silverman (2001) demonstrated erence to occupy the original nest versus the alterna- that Argentine ants and odorous house ants, Tapinoma tive substrate provided. sessile (Say), abandoned plaster nests and colonized Foraging Activity. The toxicity of various insecti- cypress mulch in choice assays. It remains unknown cides was determined by observing changes in colony whether nesting in the vicinity of the treated sub- foraging behavior, i.e., the number of workers crossing strates promoted or impeded mortality. On the one a line on the Slinky coil. Foraging activity decreased hand, colonies that relocated were in proximity to signiÞcantly for all treatments relative to controls, toxic insecticides. However, the undersides of pavers except for Cinnamite, where worker counts were of- and tiles, as well as the bottom layer of mulch were ten higher than controls (Fig. 2). Of all insecticides untreated, which may have allowed some protection. tested, Termidor was the most effective at reducing Alternatively, ants could have become contaminated ant foraging activity. Pretreatment activity declined with insecticide residues in the process of exploring by 100% by 2 d posttreatment on both concrete and alternative nesting locations and moving brood over mulch. Phantom was also highly effective on both the treated substrates. Furthermore, after colonizing outdoor and indoor substrates. On outdoor substrates, the underside of pavers, mulch, and tile, workers were Phantom showed Ͼ80% reduction of pretreatment still readily observed exploring the top surface of all foraging activity by 2 d posttreatment. On indoor sub- substrates. strates, it provided reduction in the range of 70Ð100%. The differential relocation behaviors outlined in Phantom showed differences between indoor sub- Table 2 can serve to provide an indirect measure of strates, performing slightly better on ceramic tile, relative repellency. Both control and Cinnamite treat- where almost 100% control was maintained through- ments were associated with colony relocation in all out the 6-wk period. Foraging activity also decreased replicates, demonstrating lack of repellency by these in the controls (Fig. 2). We have consistently ob- treatments. Termidor, a nonrepellent material, acted served this effect with Pharaoh ants in various exper- so rapidly that no assessments on colony relocation imental setups, and other researchers also reported could be made. However, our observations indicated declines in foraging activity in controls (Knight and that workers readily explored Termidor-treated Rust 1990). The decline in foraging activity may have mulch and pavers, thus conÞrming Þpronil to be a several underlying causes. First, initial foraging activ- nonrepellent insecticide. Chlorfenapyr also resulted ity may have been high due to the ants exploring the in ants relocating from their original nests and into the newly discovered foraging space and marking the ter- treated substrates, also demonstrating a lack of repel- ritory with species-speciÞc home range . lency by this insecticide. Both Talstar, a type I (non- Such behavior is common in other ant species (Aron ␣-cyano) pyrethroid and Demon, a type II (␣-cyano) et al. 1990, Mayade et al. 1993). Second, ants may have pyrethroid, prevented colony relocation in most as- become satiated with the food source. To minimize says. The absence of colony relocation with Talstar this effect, we performed forager counts at least 24 h and Demon suggests moderate repellency by these after colonies were provided with food. materials. It is possible, however, that other pyrethroid In treatments involving outdoor substrates all treat- insecticides will be less repellent to Pharaoh ants. ments had signiÞcant, but similar impacts on average In the current study, we used the nest relocation cumulative foraging activity over the entire 6-wk pe- behavior as an indirect method of estimating insecti- riod (Fig. 3A). Also, no signiÞcant differences oc- cide repellency. This method provides information at curred between concrete and mulch substrates in cu- the colony level, i.e., it suggests that a collective de- mulative foraging activity. In treatments involving cision is made by numerous individuals of different indoor substrates (Fig. 3B), Phantom and Demon had castes. Other studies have used more direct methods signiÞcant impacts on cumulative foraging activity, that provide information on the individual level, such with Phantom performing signiÞcantly better than April 2005 BUCZKOWSKI ET AL:COLONY BUDDING IN PHARAOH ANT 489

(A) concrete (B) mulch

100 100

80 80

60 60

40 40

20 20

0 0

on control

i control Termidor (fipronil) Termidor (fipronil) Talstar (bifenthrin) -20 -20 Talstar (bifenthrin) uct Phantom (chlorfenapyr) Phantom (chlorfenapyr) d

-40 -40 0 5 10 15 20 25 30 35 40 45 0 5 10 15 20 25 30 35 40 45

(C) vinyl tile (D) ceramic tile

100 100 ng percent re i

80 80 orag

F 60 60

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0 control 0 control Cinnamite (cinnamaldehyde) Cinnamite (cinnamaldehyde) Phantom (chlorfenapyr) Phantom (chlorfenapyr) -20 Demon (cypermethrin) -20 Demon (cypermethrin)

-40 -40 0 5 10 15 20 25 30 35 40 45 0 5 10 15 20 25 30 35 40 45

Day Fig. 2. Average (ϮSEM) percentage of reduction in worker foraging activity in response to various treatments at 2, 4, 7, 14, 21, 28, 35, and 42 d posttreatment (n ϭ 3). Shown in A and B are results for outdoor substrates, and in C and D, indoor substrates. Data shown were normalized to pretreatment counts.

Demon on both vinyl and ceramic tile. The results for formed similarly to Termidor. On indoor substrates, Cinnamite were not different from the controls. In Phantom was the most effective insecticide. It re- fact, cumulative foraging activity for Cinnamite was duced the number of workers by 83% on ceramic tile higher than the controls. Based on comparisons of and 20% on vinyl tile. Demon was signiÞcantly less percentage of reductions in foraging activity over time effective than Phantom on ceramic tile and caused (Fig. 2) versus cumulative foraging activity (Figs. 3A only 33% reduction in worker numbers. Demon was and 4A), it seems that monitoring of foraging activity ineffective at reducing colony size on vinyl tile and reductions would sufÞce for comparisons of cumula- colonies grew by 27%. Similarly, Cinnamite was inef- tive foraging activity by insecticide treatments. fective on both substrates. Worker Mortality. Based on previous research Queen Mortality. Impacts of treatments on queen (Ratliff 2003), starting worker numbers were normal- mortality are shown in Figs. 3C and 4C. SigniÞcant ized to 4,000. By 6 wk, worker numbers increased in substrate effects were observed in queen reduction in controls, with mulch showing the greatest increase. In controls. This was likely caused by the difÞculty in treatments involving outdoor substrates, Termidor assessing weekly queen numbers in paver and mulch had the most impact on worker numbers. It caused assays, due to the fact that colonies moved under 99% mortality on concrete and 100% mortality in pavers and into mulch in control and nonrepellent mulch by 6 wk. Talstar mortality ranged from 40 to insecticide assays. Termidor was highly effective at 50%, whereas Phantom mortality was different be- killing queens and resulted in 100% mortality on both tween substrates. Phantom effects on pavers were substrates. Talstar and Phantom provided signiÞcantly similar to Talstar, whereas Phantom in mulch per- lower levels of control relative to Termidor. Talstar 490 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 98, no. 2

(A) foraging activity (C) queen mortality aa 350 b a 100 300 bc a 80 250 c c 200 60 c b 150 b b b b 40 100 b d 20 50

0 0 Average number foragingweeks over6 Average percent reductionweeks at6

(B) worker mortality (D) brood mortality

aa ab a a a 100 100 bc b 80 b b 80 60 cd cd 40 60 d 20 c d 40 d 0 -20 20 -40 -60 0 -80 Average percent reductionweeks at6 Average percent reductionweeks at6 control Termidor Talstar Phantom control Termidor Talstar Phantom Fig. 3. Impacts of various treatments on colony attributes on outdoor substrates. Solid bars represent concrete pavers; open bars represent hardwood mulch. (A) Cumulative foraging activity over the course of the entire 6-wk study. (B) Cumulative worker mortality over the course of the study. (C) Queen numbers at the end of 6 wk. (D) Brood quantity at the end of 6 wk. Means with the same letter are not signiÞcantly different by the LSD t-test (P Յ 0.05). queen mortality on concrete was similar to concrete dor and Talstar, performed equally well in mulch and controls, suggesting no efÞcacy of Talstar against on concrete. The impact of Phantom on worker mor- queens. On indoor substrates, Phantom and Demon tality, queen number, and brood quantity was slightly, provided moderate reduction in queen numbers although not always signiÞcantly, higher in mulch. The (Ϸ40%). Cinnamite was ineffective on either vinyl effect of substrate was more pronounced with vinyl or ceramic tile and resulted in negligible queen mor- and ceramic tile, with all three insecticides tested tality. generally performing better on ceramic tile. This was Brood Mortality. No signiÞcant substrate effects most likely due to the nonabsorptive nature of this were observed in association with brood reduction on material. Phantom was the only insecticide tested on outdoor substrates. Both Termidor and Phantom were all four substrates, thus allowing a comparison be- highly effective at reducing the number of brood tween outdoor and indoor materials. Phantom gener- present. Talstar had no signiÞcant effect on brood ally performed better on outdoor substrates (Fig. 3). reduction relative to controls. On indoor substrates, This result is rather unexpected given that concrete Phantom was the most effective insecticide, providing and mulch are highly absorptive. However, nest relo- 45% brood reduction on vinyl tile and 83% reduction cation behaviors might provide an explanation for this on ceramic tile. The efÞcacy of Cinnamite and Demon was not different from the controls. result. Pharaoh ants moved into the mulch in three of Effect of Substrate on Insecticide Performance. Nu- three replicates, two-thirds of concrete replicates, merous biotic and abiotic factors affect insecticide one-third of vinyl tile replicates, and zero of three performance, with the application substrate being one ceramic tile replicates. Higher incidence of nesting in of the most important factors (Wagner and Strawn treated outdoor substrates most likely promoted mor- 1980, Chadwick 1985, Su and Scheffrahn 1990). Our tality by increasing proximity to the insecticides. Ants results revealed signiÞcant substrate effects, with the could have become contaminated with insecticide res- nonporous ceramic tile causing faster and greater mor- idues in the process of exploring alternative nesting tality relative to the porous substrates (mulch, con- locations and colonizing treated substrates. Further- crete, and vinyl tile). We did not, however, observe a more, after colonizing the treated substrates, workers signiÞcant difference in insecticide performance be- were readily observed exploring the top surface of tween concrete and mulch. Two insecticides, Termi- mulch and concrete. Such behaviors may have con- April 2005 BUCZKOWSKI ET AL:COLONY BUDDING IN PHARAOH ANT 491

(A) foraging activity (C) queen mortality a a 350 a a 100 300 80 250 b a 200 a 60 bc a 15 0 ab c 40 bc 10 0 bc c 20 c 50 c

0 0 Average percent reductionweeks at6 Average number foragingweeks over6

(B) worker mortality (D) brood mortality a 100 a 80 100 b 60 bc 40 80 20 b 60 0 c -20 bcd 40 c c c -40 cd c c cd d -60 20 d -80 0 Average percent reductionweeks at6 Average percent reductionweeks at6 control Phantom Cinnamite Demon Control Phantom Cinnamite Demon Fig. 4. Impacts of various treatments on colony attributes on indoor substrates. Solid bars represent vinyl tile, open bars represent ceramic tile. (A) Cumulative foraging activity over the course of the entire 6-wk study. (B) Cumulative worker mortality over the course of the study. (C) Queen numbers at the end of 6 wk. (D) Brood quantity at the end of 6 wk. Means with the same letter are not signiÞcantly different by the LSD t-test (P Յ 0.05). tributed to the higher mortality associated with out- used, with Þpronil being a good candidate. Fipronil is door substrates. highly effective against Pharaoh ants in bait (Lee In conclusion, residual insecticide barriers are com- 2000) and spray (current study) formulations. Com- monly used to control various species of house invad- bining both approaches might provide effective Pha- ing ants (Vail and Bailey 2002, Scharf et al. 2004), raoh ant control under Þeld conditions where numer- including Pharaoh ants (Edwards 1986, Oi et al. 1996). ous colonies may exist. They are generally applied outdoors as barrier sprays designed to kill and/or repel foraging ants. Our results demonstrate that nonrepellent residual insecticides, Acknowledgments most notably Þpronil, can be an effective tool for controlling Pharaoh ants indoors. The Pharaoh ant is Chris Paulsen faithfully counted dead workers and main- primarily an indoor pest, although it also may nest tained ant colonies. Jessie Hoteling and Dean Brad provided and/or forage outside in warmer climates (Kohn and valuable suggestions. We thank BASF Corp. (Research Tri- angle Park, NC) for providing partial funding for the project. Vlcˇek 1986, Oi et al. 1994, Vail and Williams 1994). As a result, treatments for Pharaoh ants are usually per- formed indoors, although outdoor treatments with baits are also effective (Oi et al. 1994, Vail et al. 1996). References Cited In the past, residual insecticide sprays have been re- Ameen, A., W. Kaakeh, and G. Bennett. 2000. Integration of garded as inferior to baits and ineffective at eradicat- chlorfenapyr into a management program for the German ing infestations. Here, we show that nonrepellent re- cockroach (Dictyoptera: Blattellidae) J. Agric. Urban En- sidual insecticides, which are active at low doses (i.e., tomol. 17: 135Ð142. Þpronil, and to a lesser degree chlorfenapyr) can re- Aron, S. J., M. Pasteels, S. Goss, and J. L. Deneubourg. 1990. sult in a complete eradication of colonies. Further- Self-organizing spatial patterns in the Iri- domyrmex humilis (Mayr), pp. 438Ð451. In R. K. Vander more, our results indicate that residual sprays should Meer, K. Jaffe, and A. Cedeno [eds.], Applied myrme- be applied to nonporous materials to maximize efÞ- cology, a world perspective. Westview Press, Boulder, cacy. To maximize control efforts in complex Þeld CO. situations (numerous nesting sites in large buildings), Buczkowski, G., and C. Schal. 2001. Emetophagy: Þpronil- a combination of baits and residual sprays might be induced regurgitation of bait and its dissemination from 492 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 98, no. 2

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