A comparison of two methods for sampling assemblages of subterranean,wood-eating termites (Isoptera)

Abstract Within a 50 × 50 m area of wandoo Eucalyptus capillosa woodland in the Western Australian wheatbelt, the diversity and frequency of occurrence of wood-eating termite species was assessed at two food types. Over a 12 month period, monthly termite activity was determined: (i) at sound/undecayed artificial baits (seasoned wooden stakes of Jarrah, Karri, Pine, Batu, Oregon; Jarrah sawdust; paper rolls); and (li) at naturally occurring timber, fallen logs and branches of wandoo, in varying stages of decay. Termite diversity was 11 species at baits, 18 species at wandoo out of an overall site richness of 21 species. Karri attracted the most species (9); sawdust attracted none. At wandoo, Nasutitermes exitiosus, Coptotermes acinaciformis and Occasitermes occasus accounted for 59% of samples where termites were recorded. At baits, Heterotermes occiduus accounted for a mean of 80% of samples across bait types, but was rarely sampled at wandoo (5% of samples). Only H. occiduus, C. acinaciformis and Amitermes neogermanus ate bait. Pine, Oregon and paper rolls were most effective in attracting foraging termites in terms of highest per cent of replicates showing bait consumption and highest consumption rates. Jarrah and Batu were least attractive to foraging termites. Samples from wandoo underestimated the relative frequency of occurrence of H. occiduus within the study site. Coptotermes acinaciformis, which attack large food items, and certain species of Amitermes, which forage on subterranean food, may have been underestimated by both sampling methods. These findings indicate that a proper understanding of the structure of wood-eating termite assemblages within a given area requires a composite sampling strategy which addresses termites that eat sound or decayed wood, as well as surface and subsurface foragers.     

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.     

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.     

Assessing the Relative Efficiency of Termite Sampling Methods along a Rainfall Gradient in African Savannas

Although termites are ecosystem engineers in tropical and sub‐tropical environments, the study of termite ecology is often constrained by sampling difficulties and a lack of established sampling protocols, particularly for savannas. The efficiency and relevance of different methods along climatic gradients, even within a single biome, is largely unknown. Here, we compare the relative contribution of two commonly used sampling methods, cellulose baits and active searching transects, in quantifying savanna termite diversity along a rainfall gradient in South Africa; sampling was conducted during the wet season across four markedly different savanna types. We also assessed the usefulness of different forms of baiting techniques. The relative efficiency of sampling method varied with annual rainfall. In arid savannas, baiting was as effective as active searching transects at sampling termite diversity and we recommend the use of baiting rather due to it being less labor intensive. In savannas of moderately low to intermediate rainfall, baiting and transects sampled different termite species and so both are deemed necessary for an accurate assessment of termite diversity. In contrast, in wetter savannas transects gave a better assessment of diversity, with cellulose baits not contributing much to diversity assessment. The efficiency of baiting techniques differed across the rainfall gradient, with baits needing to be left in the field for a longer period in more arid savannas. We conclude that habitat type, even within a single biome, will determine the sampling method or methods necessary to quantify termite diversity accurately.     

The spatial pattern of soil-dwelling termites in primary and logged forest in Sabah, Malaysia

Abstract.  1. Primary and logged lowland dipterocarp forest sites were sampled for subterranean termites using soil pits located on a grid system in order to detect any patchiness in their distribution.
2. A spatial pattern in termite distributions was observed in the primary and logged sites, but the response differed between soil-feeding and non-soil-feeding termites.
3. Spatial analysis showed that soil-feeding termites were homogeneously distributed in the primary forest but significantly aggregated in the logged forest. This pattern was reversed for non-soil-feeding termites and may result from differences in resource provisioning between the two sites.
4. Gaps in termite distribution comprised a greater area than patches for both feeding groups and sites, but gaps dominated the logged site.
5. A significant association between soil-feeding and non-soil-feeding termite distributions occurred at both sites. This arose from an association between patches in the primary forest and between gaps in the logged forest.
6. Termite spatial pattern was optimally observed at a minimum extent of 64 m and lag of 2 m.
7. The spatially explicit SADIE (Spatial Analysis by Distances IndicEs) analyses were more successful than (non-spatially explicit) multivariate analysis (Canonical Correspondence Analysis) at detecting associations between termite spatial distributions and that of other biotic and abiotic variables.     

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.     

Effects of nutrients on feeding activities of the pest termite Heterotermes tenuis (Isoptera: Rhinotermitidae)

Heterotermes tenuis is a native termite of large negative impact on the Brazilian economy. Many studies have focused on optimizing the baiting method for termite control by improving its use and its efficacy. In order to maximize termite bait consumption, this study aimed to select nutrient solutions which are able to enhance termite feeding. The effects of some sugars and urea on feeding preference of the subterranean termite H. tenuis were determined by double choice feeding tests, and the consumption was evaluated by variation in dry mass of filter paper after 28 days of termite feeding. Most of the solutions tested did not significantly influence feeding of H. tenuis . Termites fed preferentially on filter paper treated with 0.03 g/ml trehalose and 0.015 g/ml urea solutions. The data indicate that trehalose and urea solutions are phagostimulants to H. tenuis , and their future use in bait matrices may increase consumption of baits and improve this control technique.     

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.     

Estimation of foraging territories of Reticulitermes grassei through mark–release–recapture

Subterranean termites have highly cryptic life habits and their foraging activities are as a rule confined to below-ground level gallery systems. Mark–release–recapture (MRR) using fat-soluble histological dyes is a candidate method for the study of foraging dynamics and territories, but has not hitherto been standardized experimentally. A wide range of dye types and concentrations is reported in the literature. In this study, six potential dyes were evaluated at different concentrations for marking workers of Reticulitermes grassei (Clément) (Isoptera: Rhinotermitidae), under laboratory and simulated field conditions. Neutral red (1% wt/wt) was considered the most effective while showing acceptably low toxicity. In a subsequent field trial using dye impregnated into a wooden bait, the MMR procedure was used to map the foraging territory perimeter of a single colony. Assumptions inherent in the interpretation of MMR data are reviewed. To map the foraging territory perimeter by this method, two theoretical approaches are defined (a conservative and a non-conservative hypothesis). We show that the approach adopted may affect the estimate obtained by as much as 100%. Results are discussed in the context of the ecology and behaviour of subterranean termite colonies.     

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.     

The effect of bait design on bait consumption in termites (Isoptera: Rhinotermitidae)

The efficacy of baiting as a pest control method relies on the bait appealing to the pest species. In the case of wood-eating termites, bait stations should be designed to encourage termite presence and to maximize their consumption of bait matrix in order to expedite control in minimal time. A field experiment examined the effect of bait size (one large bait or four small baits of equivalent total size, with commensurate inspection and replacement schedules), compaction (tightly rolled or loosely folded) and composition (paper only or paper plus wood) on termite presence and on untreated bait paper removal rates over four months. All three factors were significant, with bait size the most important factor, followed by compaction and then composition. The least effective baits were small, compacted (rolled) paper-only baits with monthly inspections; these had the highest abandonment rate (70%) and had the least paper removed (mean of 24 g). The most effective baits were large, folded paper-plus-wood baits with inspections at two months; these had the lowest abandonment rate (20%) and had the highest paper removal (mean of 112 g). The more than four-fold difference between these baits types demonstrates that bait efficacy can be altered considerably merely by changing bait design without adding new ingredients to the bait matrix.     

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.     

Response of the Formosan subterranean termites (Isoptera: Rhinotermitidae) to baits or nonrepellent termiticides in extended foraging arenas

Distance effects of three treatments, noviflumuron, fipronil, and thiamethoxam, against laboratory populations of the Formosan subterranean termite, Coptotermesformosanus Shiraki, were tested in extended foraging arenas with foraging distances of 50 m. The results showed that during the 10-wk test period, all termites were killed by noviflumuron baits, whereas the nonrepellent termiticides fipronil and thiamethoxam divided the laboratory populations into two groups after causing 25-35% worker mortality. The horizontal transfer of lethal effects of fipronil was < or = 5 m. For thiamethoxam, the distance of transfer was substantially shorter. Because of their dose-dependent lethal time, the nonrepellent termiticides did not fulfill the requirements of a liquid bait model.     

The termite (Isoptera) fauna of a monsoonal rainforest near Darwin, northern Australia

Abstract  Termite species richness, frequency of occurrence and functional diversity at Holmes Jungle Nature Reserve, a monsoon rainforest patch in northern Australia, was investigated at the end of the wet season in 2003. A sampling protocol that employed direct search, soil pits and baiting techniques was used to sample litter, wood, mound, soil and arboreal nest microhabitats for termites. Five species from five genera and three families (Mastotermitidae, Rhinotermitidae and Termitidae) were recorded. This included the first record of Ephelotermes taylori (Hill) from monsoon forest in Australia. The family Termitidae was dominant and represented 70% of termite occurrences. Termites were most frequently encountered in carton runways on tree trunks, followed by lying dead wood and baits. Four nesting habits were represented: arboreal, epigeal, hypogeal and within wood. The arboreal nest-builder Nasutitermes graveolus (Hill) accounted for 61% of termite encounters. Epigeal mound-building species were rare. Wood-feeders were the only trophic group represented. Relatively high activity of Mastotermes darwiniensis Froggatt was recorded at baits within 2.5 months. The low species richness recorded at Holmes Jungle is consistent with the low diversity reported for Australian rainforests generally, but remains relatively depauperate compared with other monsoon forest and savanna habitats of the Northern Territory.     

Field Evaluations of Subterranean Termite Preference for Sap-Stain Inoculated Wood

Few studies have focused on interactions between subterranean termites and the ophiostomatoid fungal associates of pine bark beetles or root feeding weevils. Field stake tests were employed at four locations throughout Mississippi to determine the feeding preference of subterranean termites for blue-stained, unstained, and partially decayed southern pine sapwood stakes. This study also utilized wood decayed by Gloeophyllum trabeum, a fungus previously shown to elicit a positive subterranean termite feeding response, as a positive control. Stakes inoculated with G. trabeum received significantly more attacks than all other treatments after 16 weeks. Of the stakes attacked by subterranean termites, stakes inoculated with Ophiostoma minus were degraded faster than any other treatment. Subterranean termite preference for stakes treated with either of two Leptographium spp. and the untreated negative controls did not differ; however, each was fed upon less than all other treatments. The feeding rate on stakes inoculated with O. ips and G. trabeum being fed upon by subterranean termites was not significantly different. These results represent the first evidence of wood containing non-structurally degrading fungi (O. ips and O. minus) eliciting a feeding preference from subterranean termites greater than that of decayed wood. The implications of these results are particularly relevant to pine forest ecology, nutrient cycling, subterranean termite control, and the utilization of blue-stained southern pine building products in the southeastern U.S.     

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.     

Field observations of intercolony aggression and territory changes inHeterotermes aureus (Isoptera: Rhinotermitidae)

Intercolony aggression was observed for a southwestern desert subterranean termite, Heterotermes aureus(Snyder), in bait plots in a Sonoran Desert grassland in Arizona. Intermittent conflict took place at the sites of corrugatedpaper rolls on the boundary between two colonies. Rows of dead termites locked in combat were found within the corrugations, with additional dead termites usually in the soil beneath the roll. Dead soldiers constituted 64 to 100% of termites recovered from sites of intercolony conflict. Ongoing intercolony aggression is likely involved in the maintenance of discrete H. aureusterritories.     

Responses of Coptotermes formosanus and Reticulitermes flavipes (Isoptera: Rhinotermitidae) to three types of wood rot fungi cultured on different substrates

This study examined the responses of two termite species, the Formosan subterranean termite, Coptotermes formosanus Shiraki, and the eastern subterranean termite, Reticulitermes flavipes (Kollar), to three types of wood decay fungi: a brown rot fungus, Gloeophyllum trabeum (Persoon: Fries) Murrill; a white rot fungus, Phanerochaete chrysosporium Burdsall; and a litter rot fungus, Marasmiellus troyanus (Murrill) Singer. We also examined the responses of termites to these three types of fungi grown on different substrates. For all three fungal species, both termite species showed a strong preference for fungus-infected sawdust over uninfected sawdust. In choice tests, both termite species preferred sawdust infected with either M. troyanus or P. chrysosporium over G. trabeum. However, termites did not show any preference for fungus-infected potato dextrose agar over uninfected potato dextrose agar. Tunneling activity of C. formosanus was greater in sand treated with methanol extracts of fungus-infected sawdust than in sand treated with extracts of uninfected sawdust. Because chemicals in the fungal extracts caused termites to tunnel further into treated sand than untreated sand, these chemicals could potentially be used to direct termite foraging toward bait stations in the field.     

Use of a bait impact index to assess effects of bait application against populations of Formosan subterranean termite (Isoptera: Rhinotermitidae) in a large area

The effects of bait applications on the overall activity of the Formosan subterranean termite, Coptotermes formosanus Shiraki, in a town of 1.2 km2 were assessed between 2001 and 2003 by using a bait impact index. The index incorporated the amount of baits applied and the distance between bait application loci and the monitoring stations from which termite activity was measured. Even with the collaboration of only 35% of the residents and incomplete treatment records, the bait impact index demonstrated that more baits applied in proximity to monitoring loci significantly caused the decline of C. formosanus 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.