Evaluation of hexaflumuron for controlling the subterranean termite Coptotermes curvignathus (Isoptera: Rhinotermitidae) in Malaysia

Cellulosic bait matrices containing 0.5% hexaflumuron were tested against field colonies of the subterranean termite Coptotermes curvignathus Holmgren in Malaysia. Foraging activities of the termites were first monitored for several months by using survey stakes and wood bundles placed in underground monitoring traps. Infested stakes were replaced with bait devices. Termite workers to act as recruits were collected from infested stakes and placed in the bait devices. Subsequent activities of the termites were monitored by examining and weighing the wood in the monitoring traps. Data show that the four colonies had populations of 166,288-709,052 foragers and mean wood consumption rates of 305.9-508.2 g/mo per trap. These colonies ceased their activities at all monitoring devices 25-44 d after baiting had commenced. Colonies consumed approximately 27-79 g of bait matrix or 137.5-395 mg of hexaflumuron. Thus, C. curvignathus colonies could be monitored, characterized, baited, and foraging eliminated by using a bait matrix containing hexaflumuron.     

Elimination of subterranean termite (Isoptera: Rhinotermitidae) colonies using a refined cellulose bait matrix containing noviflumuron when monitored and replenished quarterly

Using a quarterly (3-mo) monitoring and bait-replenishment interval, 122 subterranean termite colonies throughout the United States were baited with a refined cellulose bait matrix containing 0.5% noviflumuron. All colonies were eliminated in less than 1 yr after initiation of baiting as determined by long-term monitoring and genetic markers. Sixty-three percent of the colonies were eliminated during the first quarter after the initiation of baiting and 77% of colonies were eliminated after consuming two bait tubes or less. This suggests that a single baiting cycle and bait installed in response to a single active monitoring device were sufficient to eliminate the majority of colonies. Although termites temporarily abandoned stations after depleting bait, workers resumed feeding when baits were replenished. Colonies that consumed large amounts of bait before elimination foraged into multiple stations, thus allowing adequate amounts of bait to sustain feeding. The time to eliminate termite colonies with bait replenished quarterly was similar to that previously reported for laminated cellulose bait replenished monthly. Our data support the conclusion that extending the bait replenishment interval from monthly to quarterly for bait tubes with refined cellulose containing 0.5% noviflumuron did not adversely impact colony elimination.     

Attraction of subterranean termites (Isoptera) to carbon dioxide

Subterranean termites, Reticulitermes spp., were attracted to carbon dioxide (CO2) in laboratory and field tests. In behavioral bioassays, Reticulitermes flavipes (Kollar), Reticulitermes tibialis Banks, and Reticulitermes virginicus Banks were attracted to CO2 concentrations between 5 and 50 mmol/mol. In further bioassays, R. tibialis and R. virginicus were attracted to the headspace from polyisocyanurate construction foam that contained 10-12 mmol/mol CO2. In soil bioassays in the laboratory, more termites foraged in chambers containing CO2-generating formulations than in unbaited control chambers. In field tests, stations containing CO2-generating baits attracted R. tibialis away from wooden fence posts at rangeland sites in Colorado. For all of the CO2 formulations tested, termites foraged in significantly more bait stations at treatment fenceposts than in bait stations at the control fenceposts. By the end of the 8-wk study, the number of bait stations located by termites at treatment fenceposts ranged from 40 to 90%. At control fenceposts, termites foraged in only a single station and the one positive station was not located by termites until week 5 of the study. At treatment fenceposts, termites foraged equally in active stations (containing a CO2-generating bait) and passive stations (with no CO2-generating bait), indicating that bait stations may benefit passively from a proximal CO2 source in the soil. CO2 used as an attractant in current baiting systems could improve their effectiveness by allowing earlier exposure of termites to an insecticide.     

Field evaluation of potential control strategies against the invasive eastern subterranean termite (Isoptera: Rhinotermitidae) in Chile

Four different termite control strategies consisting of two soil treatments with cypermethrin and fipronil and two bait treatments with hexaflumuron and sulfluramid were evaluated for their efficacy and potential for controlling the invasive eastern subterranean termite, Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae), in Quillota and Valparaiso, Chile. Monitoring stations were installed at all sites to measure the overall termite activity next to and at distances up to 30 m away from infested structures, most being 3-5 m. Foraging activity was extremely seasonal with the maximum consumption rates occurring from January to April. Termite foraging activity was low from July to September. After treatments in January 2003, there was a significant decrease in wood consumption, percentage of active stations, and numbers of termites in traps for the first 6 mo with hexaflumuron baits. No significant change in R. flavipes foraging activity was observed on wood consumption and percentage of active stations for sulfluramid bait or cypermethrin and fipronil soils treatments. The seasonal pattern of foraging activity remained unchanged for all treatments in 2004 and 2005, except structures baited with hexaflumuron. Feeding activity at the monitors in the area surrounding the hexaflumuron baits declined to 0 after January 2004. Alate flights occurred in approximately 38% of the structures treated with cypermethrin and sulfluramid bait in 2003 and in approximately 40% of the structures treated with cypermethrin, fipronil, and sulfluramid bait in 2004. Sulfluramid baits and cypermethrin soil treatments did not have any impact within a 15-m radius of the structures. Soils treatments of fipronil and cypermethrin prevented termites access in 75% of the homes. The number of active monitoring stations only declined within 2 m of the fipronil-treated zone. Hexaflumuron exhibited a reduction in activity for at least a 15-m radius.     

Hermetically sealed baits for subterranean termites (Isoptera: Rhinotermitidae)

Cellulose baits containing 0.5% hexaflumuron were hermetically sealed in a closed cell polyethylene sheet envelope and placed in soil to test their durability and efficacy against field colonies of the Formosan subterranean termite, Coptotermes formosanus Shiraki, or the eastern subterranean termite, Reticulitermes flavipes (Kollar). The closed cell polyethylene sheet was readily tunneled through by termites, yet it was impervious to water and protected the cellulose baits and hexaflumuron from the environment. Only in a few incidents did the polyethylene envelope become infiltrated by plant roots, resulting in water intrusion and apparent degradation of cellulose baits. After consuming one to three sealed baits, three colonies each of both termite species were eliminated. The sealed baits may be placed in soil for months or years without the need of monitoring, and they are readily penetrated and fed upon by termites when they are present. Application of baits hermetically sealed in a protective sheet may save labor costs by bypassing the monitoring phase, circumvent the station avoidance by some termite species, and enable the use of baiting technologies in large areas such as agricultural fields in which the manual monitoring is impractical.     

Control of Coptotermes havilandi (Isoptera: Rhinotermitidae) with hexaflumuron baits and a sensor incorporated into a monitoring and baiting program

A sensor consisting of a wooden monitor painted with a conductive circuit of silver particle emulsion was placed in a monitoring station to detect feeding activity of the subterranean termite Coptotermes havilandi Holmgren. Sensor accuracy was 100% 1 mo after installation, but 9 mo after sensor placement, the rate declined to 73%. After the detection of C. havilandi in the stations, baits containing the chitin synthesis inhibitor hexaflumuron were applied in five colonies, and four colonies were eliminated within 3-5 mo. Baiting could not be completed for the remaining one colony because the site became inaccessible.     

Targeted versus standard bait station placement affects subterranean termite (Isoptera: Rhinotermitidae) infestation rates

A major challenge to termite baiting in soil habitats is the prolonged time that it may take for subterranean termites (Isoptera: Rhinotermitidae) to infest stations. The objective of this research study was to determine whether the location of food sources (Sentricon in-ground monitoring stations and wooden monitors) influences the likelihood of infestation by termites. In field trials conducted at 15 structures in central Ohio, standard placement of stations at 3-4.5 m intervals was compared with targeted placements based on evidence of termite activity indoors and outdoors as well as conducive moisture conditions. Termites infested significantly more targeted placements (70/374) than standard placements (35/372) around structures. At the targeted placement sites, termites infested more wooden monitors than Sentricon stations, but this was not statistically significant. This implies that placement, rather than cellulose composition, was the more important factor. Termites first infested stations/monitors an average of 38 d sooner at targeted sites than standard placement sites. This research indicates that evidence of termite activity indoors and outdoors should be a prime consideration when placing in-ground stations.     

A computerized system for remote monitoring of subterranean termites near structures.

Sensors comprising wooden stakes painted with conductive circuits of silver particle emulsion were inserted in Sentricon stations in soil near structures. Sensors were wired to a datalogger that was programmed to test for circuit breakage every 2 h and store the data in its memory. A host computer was programmed to access the datalogger through telephone communication lines for data download every 4 d. The computerized monitoring system was tested in three remote sites, and site visits were conducted monthly for 6 mo to examine system accuracy in detecting termite activity. The mean monthly accuracy for the system to correctly report the presence (true positive) or absence of termites (true negative) in the stations was 85%, but the accuracy at 6 mo after system installation ranged from 41 to 79%. Mean sensor longevity, defined as the time for a sensor circuit to break in the absence of termites, was approximately 4.4 mo.     

Effect of fipronil on subterranean termite colonies (Isoptera: Rhinotermitidae) in the field

To assess possible colony-level effects of fipronil, a commonly used nonrepellent termiticide, we conducted a field study of eight houses in the Raleigh, NC, area with infestations of the eastern subterranean termite Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae). We installed an extensive grid of in-ground monitoring stations on each property (mean = 68.6 monitors per property) and collected samples from these stations as well as from mud tubes in the structure and wood debris in the yard for approximately 6 mo. We genotyped all samples by using microsatellite markers to identify the number and locations of colonies present on each property. Houses were treated with either a full treatment (n = 5) or exterior/localized interior treatment (n = 3). After treatment, the monitors were checked monthly for 3 mo and then quarterly for 3 yr to track the fate of colonies. Wood debris in natural areas was checked semiannually for 3 yr. All 11 of the treated colonies (those attacking structures) disappeared within 90 d of treatment and were not found again. These colonies were presumed to be eliminated. In contrast, 60% of untreated colonies (those located > 6 m from the foundation wall at the time of treatment) continued to persist throughout the study, as did 25% of the likely treated colonies (those occupying monitors 0.5 m from the foundation wall where the treatment was applied). Our results provide strong evidence for potent colony wide effects of fipronil on subterranean termites leading to colony suppression and likely colony elimination under field conditions.     

Population density, species abundance, and breeding structure of subterranean termite colonies in and around infested houses in central North Carolina

Pressure from subterranean termites is known to vary geographically across the United States, but there are few quantitative studies concerning the threat of structural infestation for any geographic region. We assessed the number and locations of termite colonies present on 20 infested residential properties in central North Carolina, where subterranean termite pressure is considered to be heavy. This was achieved by using microsatellite markers to determine colony identity of termites collected over 6-14 mo from mud tubes in structures, below-ground monitors, and wood debris in the yard. In total, we identified 188 distinct colonies and determined their breeding structures. Reticulitermes flavipes (Kollar) was by far the most common species, accounting for nearly 90% of all colonies; the remaining colonies belonged to Reticulitermes hageni Banks and Reticulitermes virginicus (Banks). In four cases, there were two colonies infesting a structure simultaneously; in all other cases only a single colony was detected in the structure. Colony densities were high, averaging 62 colonies per ha (25 per acre) with a maximum of 185 colonies per ha (75 colonies per acre). Foraging ranges of R. flavipes and R. hageni colonies were generally small (<30 linear m), and most colonies were headed by a single pair of monogamous reproductives with nearly all the remaining colonies headed by relatively few inbreeding descendants of the original monogamous pair. These results provide the most detailed picture to date of the number, distribution, and colony characteristics of subterranean termite colonies located in and around residential structures.     

Laboratory performance and pharmacokinetics of the benzoylphenylurea noviflumuron in eastern subterranean termites (Isoptera: Rhinotermitidae)

A benzoylphenylurea insect growth regulator with the common name noviflumuron was evaluated for use as a baiting toxicant against the eastern subterranean termite, Reticulitermes flavipes (Kollar). Noviflumuron demonstrated significantly greater potency and faster speed of action compared with the commercial standard hexaflumuron. In addition, noviflumuron was not a feeding deterrent on filter paper at concentrations of up to 10,000 ppm. The rates of uptake, clearance, and insect-to-insect transfer of [14C] noviflumuron were measured in R. flavipes in laboratory assays and compared with those previously reported for [14C]hexaflumuron. Under a continuous exposure regime, the uptake profile for noviflumuron was similar to that for hexaflumuron, although the time period of maximal uptake was shorter for noviflumuron. Noviflumuron was cleared from termites in a first order process with a half-life of approximately 29 d, whereas the half-life of hexaflumuron was much shorter (8-9 d). Noviflumuron was efficiently transferred from treated to untreated termites by trophallaxis via kinetics similar to those reported for hexaflumuron; however, the systemic dose of noviflumuron required to result in toxicity of R. flavipes was found to be at least two- to three-fold less than that of hexaflumuron. The faster activity of noviflumuron compared with hexaflumuron in R. flavipes can be at least partially explained by the combination of slower clearance and greater intrinsic activity.     

The Trojan horse approach for managing invasive ants: a study with Asian needle ants, <Emphasis Type="Italic">Pachycondyla chinensis</Emphasis>

Ants are among the most ecologically and economically significant biological invaders and are notoriously difficult to eradicate once established. Invasive ants are typically managed with toxic baits which are often unattractive to the target species, toxic to non-targets, and environmentally persistent. The current study evaluated a novel Trojan horse approach for managing invasive ants in natural habitats based on the use of poisoned prey. Eastern subterranean termites (Reticulitermes flavipes) were topically exposed to fipronil and presented to Asian needle ants (Pachycondyla chinensis) which are a significant invader in natural and disturbed habitats in the eastern United States. In laboratory assays, P. chinensis colonies were offered fipronil-treated termites within experimental arenas. The termites were readily attacked and consumed and results demonstrate that a single termite exposed to 25 ppm fipronil for 1 h is capable of killing 100 P. chinensis workers in 9 h. To evaluate population effects, field studies were conducted in forested areas invaded by P. chinensis. Fipronil-treated termites scattered on the forest floor provided rapid control of P. chinensis and ant densities throughout the treated plots declined by 98 ± 5 % within 28 days. I demonstrate that the poison baiting approach based on fipronil-treated termite prey is highly effective against P. chinensis and may offer an effective alternative to traditional bait treatments against other invasive ants, especially those with predatory and generalist feeding habits. Furthermore, I demonstrate that the poison baiting approach offers environmental benefits by delivering substantially less toxicant to the environment relative to current control methods which rely on commercial bait formulations. In summary, the poison baiting approach evaluated in this study appears highly suitable for controlling invasive ants and should be further tested against other invasive ants.     

Endogenous synchronization of the chemical signature of Reticulitermes (Isoptera: Rhinotermitidae) worker termites

Termites of the genus Reticulitermes are characteristic of temperate regions. Their colonies comprise various castes, the most numerous being that of workers which can develop into soldiers or secondary reproductives (neotenics). Each caste has a mixture of hydrocarbons (HCs) on the cuticle forming a chemical signature. The primary aim of this study was to compare the changes in the chemical signature of a population of worker termites fed on paper with juvenile hormone to differentiate them into soldiers with a control population of termites fed only on paper or wood for one month. Gas chromatography was used to analyze the cuticular profiles of Reticulitermes flavipes termites to determine whether they changed, and, if so, when and how. The data collected over one month showed that the workers fed with JH did not differentiate into soldiers but that there were progressive changes in the hydrocarbon profile independent of the treatment. These results indicate that the differentiation of the chemical signature of the worker caste is a dynamic process, depending only on time and not on colony membership, confirming that, for these termites, this signature has a lesser role in colony membership than caste membership, unlike the chemical signatures of other social insects. The temporal process of this cuticular change is also associated with a change in the alkene/methyl-branched alkane ratio.     

Field evaluation of the bait toxicant chlorfluazuron in eliminating Coptotermes acinaciformis (Froggatt) (Isoptera: Rhinotermitidae)

Two aspects of the Exterra Termite Interception and Baiting System (Ensystex, Fayetteville, NC) were evaluated in a field experiment using 13 termite mounds near Townsville, Australia. First, a cellulose-acetate powder containing either 0.05% wt:wt or 0.25% wt:wt chlorfluazuron (Requiem, Ensystex, Fayetteville, NC) was tested for its efficacy in eliminating colonies of the xylophagous mound-building subterranean termite Coptotermes acinaciformis (Froggatt). The moist bait matrix was replenished during the first inspection of 10 mounds (five mounds by two treatments) used in the experiment. Second, a single application of the moist bait matrix was used on three additional mounds to test termite responses and the effectiveness of 0.25% wt:wt chlorfluazuron. Although there was no evidence of repellence, there was little removal of replenished bait. Five colonies were eliminated by 0.05% wt:wt chlorfluazuron and five colonies by 0.25% wt:wt chlorfluazuron: another colony was moribund, and elimination appeared imminent. Colony decline was first suspected some 12 wk after bait application, and colony elimination was confirmed, by destructive sampling, about 5 wk later. Colony elimination may have occurred within 12 wk. One colony was an anomaly and did not succumb to the effects of the toxicant. Another colony was not eliminated because of invasion of the baiting system by ants. Ants, principally Iridomyrmex purpureus (F. Smith) group and Papyrius nitidus (Mayr) group, occurred commonly in the stations during the experiment. Microcerotermes sp. was found in five of the C. acinaciformis mounds, after colony elimination. Inspections of small sections of mounds and wooden dowels inserted into mounds were reliable methods for monitoring colony health.     

Phagostimulatory sugars enhance uptake and horizontal transfer of hexaflumuron in the western subterranean termite (Isoptera: Rhinotermitidae)

Western subterranean termite, Reticulitermes hesperus Banks (Isoptera: Rhinotermitidae), workers fed more on paper disks treated with the carbohydrates xylose, ribose, and fructose than on untreated disks. This feeding behavior of termites for certain carbohydrates was used to demonstrate an increase in the uptake and transfer of the insect growth regulator hexaflumuron among termites. The addition of 3% xylose to paper disks significantly increased the uptake of [14C]hexaflumuron and its subsequent transfer to other termites. Similarly, there was a significant increase in mortality of termites fed on paper disks treated with [14C]hexaflumuron (0.1 and 0.5%) in combination with 3% xylose for similar time periods compared with termites feeding on [14C]hexaflumuron (0.1 and 0.5%) alone. Overall percentage of mortality of termites feeding on [14C]hexaflumuron in combination with 3% xylose during the duration of the study (25-30 d) was approximately equal to 70%.     

Elimination and reinvasion studies with Coptotermes formosanus (Isoptera: Rhinotermitidae) in Louisiana

Three Formosan subterranean termite, Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae), colonies located inside the 12.75-ha Louis Armstrong Park, New Orleans, were selected for elimination by using the chitin synthesis inhibitor hexaflumuron. Once eliminated, each vacated foraging territory was monitored for reinvasion by neighboring C. formosanus colonies, Reticulitermnes flavipes (Kollar) (Isoptera: Rhinotermitidae) colonies, or both. Each selected colony was eliminated in approximately 3 mo by using baits containing hexaflumuron. Overall activity of each untreated colony in the park remained unchanged during the same period. New C. formosanus and R. flavipes activity was detected in two of the three vacated territories, and in both areas, within days of selected colony elimination. The third vacated territory was completely reoccupied by a new C. formosanus colony approximately 7 mo later. Mark-recapture studies and DNA fingerprinting confirmed the distinctness of the reinvaders from eliminated and neighboring colonies.     

Colony interactions in <Emphasis Type="Italic">Reticulitermes grassei</Emphasis> population assessed by molecular genetic methods

The cryptic habits of subterranean termites impair studies of colony delimitation and the organization of foraging. Whereas feeding sites can be identified and the movements of foragers between them can be assessed using traditional mark-release techniques, the assignment of the termites found at feeding sites to their parent colonies remains problematical. Thus the extent and overlap of individual colony foraging territories are unknown. We used microsatellite markers to delineate colonies of R. grassei and compared the results with data from a mark-release-recapture study (Nobre et al., 2007) carried out in the same population and over the same period of time. The majority of colonies exhibited only a single-feeding site, but presented, even at a local scale, a high degree of variability in colony structure and no evidence of inbreeding. F-statistics for some colonies were consistent with pleometrosis. The population as a whole contained individual colonies that were separately identified as Mendelian, mixed family or pleometrotic families. Furthermore, microsatellite data suggest that what was represented by the MRR technique as a single foraging group could be drawn from more than one colony. Colony delineation is discussed as a tool for understanding overall population structure and termite feeding strategies. Received 5 July; revised 3 October 2007; accepted 19 October 2007.     

Fox‐baiting in agricultural landscapes in south‐eastern Australia: a case‐study appraisal and suggestions for improvement

Summary In south‐eastern Australia, the introduced Red Fox (Vulpes vulpes) is a major predator of native wildlife and livestock. Fox control in agricultural landscapes is heavily reliant on the laying of poisoned baits by private landholders, yet there have been few assessments of the application or success of landholder‐baiting practices. We evaluated a community‐based fox‐baiting campaign, typical of programs employed throughout the agricultural regions of south‐eastern Australia to control foxes. We recorded the spatial coverage of 1080 baits deployed by landholders, assessed baiting procedures, monitored the survival of six radio‐collared foxes during and after baiting, and compared the spatial coverage and likely effectiveness of the baiting program with two alternative (theoretical) baiting strategies. Relative to other baiting programs, coordination among neighbours was reasonably high, with 37.5% of baited properties (n = 40) adjoining ≥3 neighbouring properties that also contained baits. Nevertheless, the maximum distance from the centre of a baited property to the nearest edge of an unbaited property was <750 m (mean = 380 m ± 147 m SD). On average, 33% (±17% SD) of each fox’s home range overlapped with baited properties, but only two foxes died during the baiting program. The remaining four foxes were still alive 10 weeks after baiting ceased. Modelling of simulated fox home ranges showed that 13.5% contained no bait stations based on the community baiting program, whereas alternative roadside‐ and grid‐baiting strategies (theoretically) delivered baits to all simulated home ranges. Some landholders employed practices that could reduce the effectiveness of baiting programs such as not removing decayed baits before deploying new ones or placing bait stations too close together. Our research illustrates the difficulties of managing a coordinated baiting program on private land that effectively controls foxes. Alternative baiting strategies such as roadside baiting need to be considered to improve fox control in agricultural landscapes.     

Asexual queen succession in the subterranean termite Reticulitermes virginicus

Termite colonies are founded by a pair of primary reproductives. In many species, including subterranean termites (family Rhinotermitidae), the primary king and queen can be succeeded by neotenic reproductives that are produced from workers or nymphs within the colony. It is generally believed that these neotenics inbreed within the colony, sometimes for many generations. Here, we show that primary queens of the North American subterranean termite, Reticulitermes virginicus, are replaced by numerous parthenogenetically produced female neotenics. We collected functional female neotenics from five colonies of R. virginicus in North Carolina and Texas, USA. Genetic analysis at eight microsatellite loci showed that 91-100% of the neotenics present within a colony were homozygous at all loci, indicating that they were produced through automictic parthenogenesis with terminal fusion. In contrast, workers, soldiers and alates were almost exclusively sexually produced by mating between the female neotenics and a single king. This is the second termite species shown to undergo asexual queen succession, a system first described in the Japanese species, Reticulitermes speratus. Thus, the conditional use of sexual and asexual reproduction to produce members of different castes may be widespread within Reticulitermes and possibly other subterranean termites.     

Feeding site selection by workers of the Formosan subterranean termite Coptotermes formosanus (Isoptera: Rhinotermitidae) - a re-analysis of field data from a mark-recapture study

During the early stages of the development of termite baits, dyed paper was placed in specified feeding stations to ascertain whether a slow-acting toxicant could be placed in a few bait stations to be delivered to the entire colony members of the Formosan subterranean termite, Coptotermes formosanus Shiraki. Feeding frequency data, as measured by the dye concentration in individual termites, suggested the absence of feeding site fidelity. However, these results were often misinterpreted as random movement of termites that were marked and released for population estimate studies, or the random search of food in soil by subterranean termites. A computer simulation program was constructed to re-examine this feeding frequency data, and confirmed the earlier conclusion that the lack of feeding site fidelity was the most likely explanation for the data.