Feeding, uptake, and utilization of carbohydrates by western subterranean termite (Isoptera: Rhinotermitidae)

Western subterranean termite, Reticulitermes hesperus (Banks), prefers various mono-, di-, and trisaccharides, total feeding being the greatest on paper disks treated with 5% ribose followed by 3% xylose, 2% maltose, 2% fructose, 2% arabinose, and 2% ribose. In multiple choice tests, termites were not able to discriminate between 2% ribose, 2% fructose, 2% xylose, and 2% maltose. Termites readily take up [14C] sucrose in feeding studies. Most of the sucrose is used as an energy source for respiration (89.2%), a very small proportion remains within the termite (9.3%), and an even smaller amount is excreted as solid waste (1.5%). The amount of 14C label transferred to other colony members via trophallaxis, body contact, or grooming is small and directly dependent upon the time and numbers of donors and recipients. At day 15 postmixing, the percentage of transfer was highest, 14.4 and 15.1% for both 1:1 and 2:1 donor to recipient mixing ratios, respectively. The mean amount of labeled 14C received by recipients increased from seven disintegrations per minute (dpm) on day 2 to 30 dpm on day 15 for 1:1. Overall mean radioactivity recovered from recipient termites when mixed with donor termites at 1:1 ratio (20 dpm) was significantly less than (28 dpm) when mixed with donor termites at 2:1 ratio. Sugars act as phagostimulants to the termites at concentrations much higher to those that termites naturally encounter in wood. Termites readily metabolize carbohydrates such as sucrose, and thus their use in bait matrices may increase consumption and retention at bait stations.     

Efficacy of vetiver oil and nootkatone as soil barriers against Formosan subterranean termite (Isoptera: Rhinotermitidae).

Vetiver oil and its components nootkatone and cedrene were assessed as sand treatments for their efficacy to disrupt food recruitment by Coptotermes formosanus Shiraki. Termites were required to tunnel through sand treated with vetiver oil, nootkatone, cedrene, or untreated sand to reach a food source. Results showed that sand treated with vetiver oil or nootkatone disrupted termite tunneling behavior. As a consequence, after 21 d, wood consumption and termite survival were significantly lower compared with cedrene-treated or untreated sand treatments. Sand treated with vetiver oil or nootkatone at 100 microg/g substrate were effective barriers to termites.     

Laboratory assays evaluate the influence of physical guidelines on subterranean termite (Isoptera: Rhinotermitidae) tunneling, bait discovery, and consumption

Laboratory assays were conducted to determine whether physical guidelines could direct subterranean termite foraging behavior. Several materials (wood, plastics, and wood thermoplastic composites) were evaluated for their potential to serve as termite guidelines. Termite tunneling along the different types of guidelines was measured. The proportion of baits discovered when connected by a guideline was compared with the proportion of unconnected baits discovered. Termite consumption of baits also was quantified. Assay results indicated that the termites did not respond to all guideline materials in the same way. Termites built significantly longer tunnels along wood guidelines than they did along any of the plastic guidelines tested. However, tunnel length along the wood and the wood thermoplastic composites was not significantly different. The probability of two baits being discovered when they were connected by wood guidelines was significantly greater than when the baits were connected by plastic guidelines or left unconnected (no guideline). Pairs of baits connected by wood thermoplastic composites were also significantly more likely to be discovered than unconnected baits. Bait consumption was not enhanced by the presence of the guidelines. It is likely that guidelines made of wood competed with the baits as a termite food resource.     

Utilizing rabbit immunoglobulin G protein for mark-capture studies on the desert subterranean termite, Heterotermes aureus (Snyder)

Mark-capture dispersal studies were conducted to investigate the feasibility of marking the southwestern desert subterranean termite, Heterotermes aureus (Snyder) with rabbit immunoglobulin G (IgG). In turn, short-range dispersal patterns of H. aureus were measured across a 20-m diameter desert landscape at three distinct field locations. Each location consisted of 51 termite feeding stations containing corrugated cardboard. The central feeding station (CFS) at each location was impregnated with rabbit IgG. A circular grid was then constructed around each CFS that consisted of 50 additional unmarked cardboard feeding stations strategically placed around the CFS at distances of 1.5, 2.0, 4.0, 7.0 or 10.0 m. Termites self-marked with rabbit IgG by feeding on the marked bait. The CFS and the 50 peripheral feeding stations were sampled for marked termites twice at each location 17–65 days after the marked bait was placed at the CFS to determine the spatial dispersal patterns of H. aureus within each research grid. Termites that self marked by feeding on rabbit IgG marked bait were detected by an anti-rabbit IgG enzyme-linked immunosorbent assay (ELISA). Generally, the CFSs contained the highest frequency of marked termites with 28.0% of the individuals assayed from the CFSs containing rabbit IgG. Over the course of the study, 39 of the unmarked peripheral feeding stations contained at least one marked termite. Of the termites assayed from the peripheral stations (n = 2,955), 124 or 4.2% of the individuals contained the mark. The average distance traveled by the marked termites collected at the peripheral feeding stations was 5.7 ± 3.3 m from the CFSs. We also examined single nucleotide polymorphisms (SNPs) from termites collected at each field site. Data indicated that each field site were genetically distinct and therefore non-related termites. We discuss the advantages and limitations of marking termites with rabbit IgG for dispersal studies.     

Directional change in tunneling of subterranean termites (Isoptera: Rhinotermitidae) in response to decayed wood attractants

Wood discs decayed with brown rot fungi and polymer discs impregnated with acetone extract of decayed wood were embedded in sand of a two-dimensional foraging arena to evaluate their attractant potential in directing termite tunnels toward them. Termites were released near one arena corner and were guided to follow the physical guideline of the arena edges. In the absence of the attractants, termites generally formed a relatively linear tunnel along the edges. When decayed wood discs or treated polymer discs were placed in wet sand near one side of the arena, termite tunnels departed from the arena edges and were oriented toward them. The attraction distance ranged from 12 to 18 cm. The attractant properties were most likely water soluble and permeated through wet sand to cause termites to change their orientation. The results demonstrated that when such attractants are placed near a bait station in the field, they may be used to direct termite foraging toward the station.     

The influence of soil heterogeneity on exploratory tunnelling by the subterranean termite Coptotermes frenchi (Isoptera: Rhinotermitidae)

The exploration of sand-filled arenas by workers of an entire colony of the Australian, subterranean foraging, tree-nesting termite, Coptotermes frenchi Hill was investigated under laboratory conditions. The first experiment tested whether termite exploration of sand was influenced by the presence of gaps or objects in the sand. Gaps and objects were chosen to represent soil heterogeneity in the urban environment: gaps to represent tunnels dug by other animals, perspex strips to represent cables and pipes, and wood strips to represent roots. Termites always chose to explore gaps thoroughly before they began tunnelling in the sand. Significantly more and longer tunnels were excavated from the end of gaps at the far end of the arenas, and relatively little tunnelling occurred around and along objects. Termite density was significantly greater around and along wood compared with perspex blocks. The second experiment tested whether termite exploratory tunnelling was influenced by soil moisture. The termites tunnelled slowly in dry sand, but after discovering a patch of wet sand, increased tunnelling five-fold until it was completely explored, after which activity declined. Energy and water conservation may be behind these patterns of exploratory tunnelling as well as those seen in large field studies, but caution is urged when interpreting small scale laboratory experiments to explain large scale field data.     

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.     

Tunneling activity of subterranean termites (Isoptera: Rhinotermitidae) in sand with moisture gradients

Tunnel formation by Coptotermes formosanus and Reticulitermes flavipes was studied in a two-dimensional foraging arena with a moisture gradient. Moisture did not appear to affect tunneling when termite first emerged from the central release chamber. But as termites of both species moved further away from the chamber and into a moisture gradient, they tunneled significantly (P < 0.05) more in sand with a higher moisture content than in sand with a lower moisture content. Over the 10 d test period, both termite species tunneled more in sand with a higher moisture content. Fractal analysis indicated that regardless of the sand moisture content, termite tunnel geometry had a fractal dimension and termites generally tunneled more in higher moisture sand.     

Diversity and resilience of the wood‐feeding higher termite Mironasutitermes shangchengensis gut microbiota in response to temporal and diet variations

Termites are considered among the most efficient bioreactors, with high capacities for lignocellulose degradation and utilization. Recently, several studies have characterized the gut microbiota of diverse termites. However, the temporal dynamics of the gut microbiota within a given termite with dietary diversity are poorly understood. Here, we employed 16S rDNA barcoded pyrosequencing analysis to investigate temporal changes in bacterial diversity and richness of the gut microbiota of wood‐feeding higher termite Mironasutitermes shangchengensis under three lignocellulose content‐based diets that feature wood, corn stalks, and filter paper. Compositions of the predominant termite gut residents were largely constant among the gut microbiomes under different diets, but each diet caused specific changes in the bacterial composition over time. Notably, microbial communities exhibited an unexpectedly strong resilience during continuous feeding on both corn stalks and filter paper. Members of five bacterial phyla, that is, Spirochaetes, Firmicutes, Actinobacteria, Tenericutes, and Acidobacteria, were strongly associated with the resilience. These findings provide insights into the stability of the gut microbiota in higher termites and have important implications for the future design of robust bioreactors for lignocellulose degradation and utilization.     

Effect of seasonal changes in soil temperature and moisture on wood consumption and foraging activity of Formosan subterranean termite (Isoptera: Rhinotermitidae)

The objective of this study was to determine how seasonal changes affect the foraging activity and wood consumption of the Formosan subterranean termite, Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae), in New Orleans, LA. There was a significant correlation between wood consumption and air temperature, soil temperature, and soil moisture, but not precipitation or number of rainy days. In the first year of the study, wood consumption was the lowest in December, February, and March. Wood consumption in January was not significantly different from consumption during the rest of the year. There were no seasonal changes in the number of underground monitoring stations occupied by termites. In the second year of the study, wood consumption was lowest from January to March. There was a significant decrease in the number of monitoring stations occupied by termites during the winter. This study determined that C. formosanus will remain in monitoring stations and resume feeding during warmer periods of a mild winter if average soil temperatures remain above 15 degrees C. Only prolonged periods of cold weather, with average soil temperatures below 15 degrees C, caused a significant number of termites to abandon underground monitoring stations. Seasonal changes in foraging activity would probably only disrupt baiting programs during severe winters in New Orleans, LA.     

Measuring wood consumption by subterranean termites (Isoptera: Rhinotermitidae) with digitized images

A method is described in which consumption by termites of a feeding block composed of 36 thin (2-mm) wooden slats is estimated with a video image analysis. The significant regression between dry wood weight loss and decreased pixel count of digitized images demonstrates that the video image technique may be a useful tool to quantify termite feeding. Time required to measure wood consumption with the video image technique (approximately 11.5 min per sample) was significantly less than that for the manual method of washing, and weighing wooden blocks damaged by termite feeding (approximately 21.0 min per sample).     

Toxicity, repellency, and transfer of chlorfenapyr against western subterranean termites (Isoptera: Rhinotermitidae)

Chlorfenapyr is a slow-acting insecticide against western subterranean termite, Reticulitermes hesperus Banks, when applied to sand. The LD50 at day 7 for workers is 29.98 ng per termite and considerably higher than that of chlorpyrifos (14.01), cypermethrin (3.21), and fipronil (0.16). Brief exposures to sand treated with chlorfenapyr resulted in dose-dependent mortality over a broad range of concentrations. Brief 1-h exposures to > or =75 ppm provided >88% kill of termites at day 7. Chlorfenapyr deposits did not repel termites, even at 300 ppm. Termites tunneled from 0.1 to 1.8 cm into sand treated with 10- to 300-ppm chlorfenapyr deposits, resulting in > or =70% mortality. Within 1 h after being exposed to 50 ppm chlorfenapyr, approximately 17% of the termites exhibited impaired responses to synthetic trail pheromone. By 4 h, nearly 60% of the workers were not able to follow a 10 fg/cm pheromone trail. There was a direct linear relationship of the uptake of [14C]chlorfenapyr as concentration and duration of exposure increased. The percentage of chlorfenapyr transferred to recipients varied from 13.3 to 38.4%. Donors exposed for 1 h transferred a greater percentage of chlorfenapyr than did donors exposed for 4 h. A 1-h exposure on 100-ppm deposits provided sufficient uptake to kill 100% of the donors and sufficient transfer to kill 96% of the recipients. There was not enough transfer for recipients to serve as secondary donors and kill other termites. Horizontal transfer is limited to contact with the original donor and by the decreased mobility of workers within 4-8 h after exposure to treated sand. The effectiveness of chlorfenapyr barrier treatments is primarily due to its nonrepellency and delayed toxicity.     

Effect of a lignin-degrading fungus on feeding preferences of Formosan subterranean termite (Isoptera: Rhinotermitidae) for different commercial lumber

The feeding preferences of the Formosan subterranean termite, Coptotermes formosanus Shiraki, for commercial lumber Alaska yellow cedar, Chamaecyparis nootkatensis (D. Don) Spach; yellow birch, Betula alleghaniensis Britton; northern red oak, Quercus rubra L.; redwood, Sequoia sempervirers (D. Don) Endl; and spruce (Picea spp.) were examined to determine whether the presence of the lignin-degrading basidiomycete Marasmiellus troyanus (Murrill) Singer could alter the relative preference of termites for these wood species. In paired choice tests with fungus-inoculated sawdust versus control sawdust, termites showed a strong preference for the fungus-inoculated sawdust for all wood species tested, except for Alaska yellow cedar. In a multiple-choice test using sawdust without fungus, termites showed a very strong preference for red oak sawdust over the other three species. In a paired choice test using fungus-inoculated sawdust, termites showed a preference for redwood over red oak sawdust. In a feeding test using autoclaved wood blocks without fungal decay, there was no difference in termite consumption of birch, red oak, or redwood. The relative preference of termites for redwood increased when blocks were decayed by M. troyanus for 3 and 8 wk. These results indicate that chemical modifications due to fungal decay affected the feeding preference of termites for different commercial lumber.     

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.     

Iron reduction in the metal-rich guts of wood-feeding termites

Termites play important roles in lignocellulose and humus turnover in diverse terrestrial ecosystems, and are significant sources of global atmospheric methane and carbon dioxide. All known termite species engage in obligate, complex nutritional symbioses with their gut microbes to carry out such processes. Several hundred microbial species, representing a broad phylogenetic and physiological diversity, are found within the well‐bounded, microliter‐in‐scale gut ecosystem of a given termite. However, most of these species have never been obtained in laboratory culture, and little can be said about their functional roles in the gut community or symbiosis. Herein, an unappreciated facet of the gut chemistry and microbiology of wood‐feeding termites is revealed: the redox metabolism of iron. Gut fluids from field‐collected termites contained millimolar amounts of ferrous iron and other heavy metals. When iron(III) hydroxides were amended to a filter paper diet of Zootermopsis nevadensis, a dampwood termite collected in the San Gabriel Mountains of Southern California, the specimens accumulated high levels of iron(II) in their guts. Additionally, iron was reduced at rapid initial rates in anoxic gut homogenates prepared from field‐collected Z. nevadensis specimens. A Clostridium sp. and a Desulfovibrio sp. were isolated from dilution‐to‐extinction enrichments of Z. nevadensis gut contents and were found to reduce iron(III), as did the termite gut spirochete Treponema primitia. The iron in the guts of wood‐feeding termites may influence the pathways of carbon‐ and electron‐flow, as well as microbial community composition in these tiny ecosystems of global importance.     

Effect of orange oil extract on the formosan subterranean termite (Isoptera: Rhinotermitidae)

The Formosan subterranean termite, Coptotermesformosanus Shiraki (Isoptera: Rhinotermitidae), accidentally brought into the United States, has become a major urban pest, causing damage to structures and live trees. Because of increasing restrictions on the use of conventional termiticides, attention is focused on finding safer alternative methods for termite management. Oil from citrus peel, referred to here as orange oil extract (OOE), contains -92% d-limonene, and it is generally known to be toxic to insects. In laboratory experiments, 96 and 68% termites were killed in 5 d when OOE at 5 ppm (vol:vol) was dispensed from the top or bottom, respectively, with termites held at the opposite end of a tight-fitting plastic container. Apart from high mortality, workers exposed to vapor consumed significantly less filter paper than controls. However, when termites were exposed to OOE vapor, even at 10 ppm, in the void of a model wall, there was very little mortality. Termites did not tunnel through glass tubes filled with sand treated with 0.2 or 0.4% OOE. Sand treated with OOE was extracted each week for 8 wk to determine the remaining amount of d-limonene. Results indicated that there was a sharp decline in the quantity of d-limonene during the first 3 wk to a residual level that gradually decreased over the remaining period. With a suitable method of application and in combination with other control practices, OOE can be effectively used for the control of subterranean termites.     

Effects of sublethal exposure to imidacloprid on subsequent behavior of subterranean termite Reticulitermes virginicus (Isoptera: Rhinotermitidae)

Experiments were conducted to determine whether subterranean termites, Reticuliternes virginicus (Banks), previously exposed to sublethal doses of imidacloprid (Premise), and allowed to recover for 1 wk, demonstrated behavioral aversion to a subsequent exposure. Worker termites experiencing a previous sublethal but debilitating exposure to imidacloprid-treated sand (either 10 or 100 ppm for 4 h) showed no apparent aversion to a second encounter with imidacloprid-treated sand under conditions of this experiment. If these laboratory results hold in the field and termites traveling through a zone of soil treated with imidacloprid are impaired but subsequently recover, they will be just as likely as their naive nestmates to reenter the treated area if their travels take them through the nonrepellent application a second time. Our results also indicate that a sublethal exposure to imidacloprid can affect termite tunneling behavior. Many worker termites that received an initial 4-h exposure to 100 ppm imidacloprid-treated sand died, but those that survived tunneled significantly less than did their naive nestmates, as did some termites exposed to 10 ppm imidacloprid.     

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.     

Feeding preference of Coptotermes formosanus (Isoptera: Rhinotermitidae) for gamma-irradiated wood impregnated with benzoylphenylurea compounds under laboratory conditions

The feeding preference of Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae) for 200-kGy gamma-irradiated Cryptomeria japonica D. Don (Japanese cedar) sapwood impregnated with benzoylphenylurea compounds such as hexaflumuron and noviflumuron was examined by three laboratory tests. Although termites were not deterred from feeding on gamma-irradiated wood samples that had been impregnated with hexaflumuron or noviflumuron, termite mortality was significantly higher compared with solvent controls in the no-choice test. All live termites were transferred to paper disks immediately after the no-choice test to investigate changes in mortality with time, and this test also confirmed the effects of hexaflumuron and noviflumuron on worker termites, which showed a significant feeding preference for gamma-irradiated wood. Only the 1480 ppm noviflumuron-impregnated gamma-irradiated wood specimens showed significant differences in mortality in the two-choice test. These results suggest that gamma-irradiated C. japonica wood, which is locally abundant in Japan, may have potential as a bait substrate for benzoylphenylurea compounds.