Sampling subterranean termite species diversity and activity in tropical savannas: an assessment of different bait choices

Abstract.  1. Cellulose baits are commonly used for semi-quantitative sampling of subterranean wood-feeding termites, with a single food choice sampling programme implemented most often. In most situations, however, the composition and feeding preferences of the subterranean termite assemblage remain unknown.
2. The diversity, frequency, foraging activity, and intensity of attack of termites were assessed regularly at 144 baits representing 12 different bait choices over 8.5 months, in two northern Australian tropical savanna sites that differed in vegetation structure (closed vs open). Baits differed in type (paper rolls, cardboard, wooden stakes), position (surface, buried), and moisture status at installation (wet, dry).
3. Sixteen species were recorded, including 11 wood-feeders. Average species diversity, foraging activity, and bait consumption were greater at buried baits than at surface baits. Wooden baits were most attractive early in the experiment, and paper baits more attractive later. Mean species diversity was greatest at wooden stakes in the closed site. Species frequency of occurrence varied across bait choices.
4. A composite bait sampling protocol of stakes and paper rolls installed above and below ground gave an accurate assessment of the activity, diversity, and structure of the termite guild sampled across all baits over 8.5 months.
5. The choice of bait, its presentation, and time of examination are critical to the success of a termite baiting programme. If the aim is to characterise the structure and foraging activity of the subterranean termite assemblages that are attracted to baits, composite baiting protocols should be implemented.     

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.     

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.     

Cryptic termites avoid predatory ants by eavesdropping on vibrational cues from their footsteps

Eavesdropping has evolved in many predator–prey relationships. Communication signals of social species may be particularly vulnerable to eavesdropping, such as pheromones produced by ants, which are predators of termites. Termites communicate mostly by way of substrate‐borne vibrations, which suggest they may be able to eavesdrop, using two possible mechanisms: ant chemicals or ant vibrations. We observed termites foraging within millimetres of ants in the field, suggesting the evolution of specialised detection behaviours. We found the termite Coptotermes acinaciformis detected their major predator, the ant Iridomyrmex purpureus, through thin wood using only vibrational cues from walking, and not chemical signals. Comparison of 16 termite and ant species found the ants‐walking signals were up to 100 times higher than those of termites. Eavesdropping on passive walking signals explains the predator detection and foraging behaviours in this ancient relationship, which may be applicable to many other predator–prey relationships.     

Abundance and diversity of termites (Isoptera) in imburnt versus burnt vegetation at the Barrens in Mediterranean Western Australia

Abstract Two years after an intense fire burnt large parts of the Barrens in Mediterranean Western Australia, its effects on wood-eating and litter-harvesting termites were investigated. Nine vegetation types varying in height, structure and floristics, were used for paired unburnt/burnt comparisons. Wood-eating termites were significantly less abundant and diverse in burnt stands, apparently by perishing in the fire rather than through food limitation. The harvester Tumulitermes westraliensis was not significantly affected by fire. Strategies enhancing persistence despite intense fire were construction of hard, protective clay mounds (Coptotermes frenchi, Amitermes obeuntis, T. westraliensis) and flexibility to site nests in diverse microhabitats, including the mounds of other termite species (Heterotermes).     

Effects of two locust control methods on wood-eating termites in arid Australia

Termites are ubiquitous detritivores and are a key influence on soil function and nutrient cycles, particularly in arid and semi-arid ecosystems. Locust control presents a unique hazard to termites and the effective functioning of ecosystems as a consequence of the overlap between pesticide applications and termite populations in grassland and desert landscapes. We monitored the effects of locust control methods using ultra-low-volume (ULV) barrier application of a chemical pesticide, fipronil, and a blanket application of a fungal biopesticide, Metarhizium acridum, on wood-eating termites in arid western New South Wales, Australia. We tested the hypothesis that spray applications decrease termite activity at wood baits using a BACI designed field experiment over 2 years. Our replicated control and treatment sites represented the spatial scale of Australian locust control activities. There was no detectable impact of either locust control treatment on termite activity, bait mass loss or termite community composition measures. Non-significant differences in termite survey measures among sites suggested that climate and environmental conditions were stronger drivers of our termite measures than the single, localized and unreplicated application of pesticides more commonly used in locust control operations in arid Australia. A lack of evidence for an impact of our fipronil or Metarhizium application methods supports their use as low hazard locust control options with minimal large scale and longer-term effects on termites in Australian arid rangelands. Future research would be necessary to determine the probable short-term impacts of treatments on individual termite colonies and the possible impacts on non-wood eating termite species in the arid-zone.     

How small is too small for small animals? Four terrestrial arthropod species in different-sized remnant woodlands in agricultural Western Australia

Island biogeography theory, and the 50/500 rule of genetics, have effectively devalued small habitat fragments for species conservation. Metapopulation theory has given new value to small remnants but data on species persistence are scarce. This study examined the capacity of very small and sheep-grazed remnants of eucalypt woodland in agricultural Western Australia to support remnant-dependent terrestrial arthropods. We surveyed 53 sheep-grazed remnants of wheatbelt wandoo Eucalyptus capillosa for the presence of four species of arthropod with different dispersal strategies (terrestrial versus aerial) and diet (predaceous vs. herbivorous): the harvester and mound-building termite Drepanotermes tamminensis, the wood-eating and mound-building termite Amitermes obeuntis, the predaceous and burrowing scorpion Urodacus armatus and the predaceous 'bull' ant Myrmecia nigriceps. All species with the exception of the scorpion disperse aerially, and all construct above-ground structures that are easily recognized. Remnants ranged in size from 50 m² to 21 000 m² (mean 1791 m²), in spatial isolation (distance to the nearest vegetation remnant) from 10 m to 500 m (mean 123 m) and in a length-to-width ratio (shape) from circular (mean ratio 1.0) to linear (mean ratio 4.0). Observations in small and grazed remnants were compared with observations made in six wandoo woodland sites within a large (1040 ha) and ungrazed remnant. The total number of target species was highly correlated with remnant area (r = 0.68). Remnant isolation and remnant shape had no apparent influence on the total number of target species. The minimum area of grazed remnants in which individual species were recorded followed the large predator Urodacus armatus (4515 m²) > smaller predator Myrmecia nigriceps (300 m²) > harvester termites Drepanotermes tamminensis (102 m²) > wood-eating termites Amitermes obeuntis (50 m²). With the exception of U. armatus which occurred only in three of the four largest grazed remnants, the occurrence of all other species increased from small to large grazed remnants, suggesting a remnant-size effect for all species. Remnant isolation or remnant shape had no apparent influence on the occurrence of any one species. The terrestrially dispersing scorpion persisted in remnants despite their isolation from other remnants from 200 m to 500 m. For both termite species, mound heights were significantly greater in large, ungrazed woodlands than in small and grazed woodlands. The incidence of mound abandonment in smaller and grazed remnants was considerably higher for harvester than for wood-eating termite colonies. This suggests differences in spatial requirements and possibly diet-related susceptibilities to fluctuations in food availability. The diameter of Myrmecia nigriceps nests showed no relationship with remnant size or isolation. This study demonstrated that even very small remnant woodlands on farms may play an important role in sustaining small native animals, either as stepping-stones for dispersing individuals (termites, ants) or in providing adequate habitat to sustain populations for longer periods (all four species).     

Estimating relative decline in populations of subterranean termites (Isoptera: Rhinotermitidae) due to baiting.

Although mark-recapture protocols produce inaccurate population estimates of termite colonies, they might be employed to estimate a relative change in colony size. This possibility was tested using two Australian, mound-building, wood-eating, subterranean Coptotermes species. Three different toxicants delivered in baits were used to decrease (but not eliminate) colony size, and a single mark-recapture protocol was used to estimate pre- and postbaiting population sizes. For both species, the numbers of termites retrieved from bait stations varied widely, resulting in no significant differences in the numbers of termites sampled between treatments in either the pre- or postbaiting protocols. There were significantly fewer termites sampled in all treatments, controls included, in the postbaiting protocol compared with the pre-, suggesting a seasonal change in forager numbers. The comparison of population estimates shows a large decrease in toxicant treated colonies compared with little change in control colonies, which suggests that estimating the relative decline in population size using mark-recapture protocols might to be possible. However, the change in population estimate was due entirely to the significantly lower recapture rate in the control colonies relative to the toxicant treated colonies, as numbers of unmarked termites did not change between treatments. The population estimates should be treated with caution because low recapture rates produce dubious population estimates and, in some cases, postbaiting mark-recapture population estimates could be much greater than those at prebaiting, despite consumption of bait in sufficient quantities to cause population decline. A possible interaction between fat-stain markers and toxicants should be investigated if mark-recapture population estimates are used. Alternative methods of population change are advised, along with other indirect measures.     

The Selection of an Isolate of the Hyphomycete Fungus,Metarhizium anisopliae,for Control of Termites in Australia

Ninety-three isolates ofMetarhizium anisopliae,mostly derived from a survey of termite material, were screened for activity againstNasutitermes exitiosusandCoptotermes frenchiorC. acinaciformisusing a grooming assay technique. Twenty-six of the most promising isolates were further evaluated by bioassay againstN. exitiosusandC. acinaciformis.All isolates were pathogenic withCoptotermesspp. being more susceptible thanN. exitiosus.A group of nine isolates, chosen for their level of pathogenicity for one or other genus of termites and to represent a genetically diverse group, was finally compared in a minicolony test using termite colonies in 1 liter jars. The isolate, code-named FI-610 (derived from nest-mound material ofC. lacteusin SE New South Wales), was one of the most effective isolates against termites from both of the two colonies tested. This isolate also grew relatively well on agar plates at 36°C. FI-610 was thus selected for field trials and was found to be effective in killing colonies ofC. acinaciformiswhen 10 g (=3 × 10¹¹conidia) or more of conidial powder was blown into the center of the large mound colonies.     

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.     

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.     

Toxic effects of 2-deoxy-D-galactose on Coptotermes formosanus (Isoptera: Rhinotermitidae) and symbionts

In the interest of developing interventions to infestations by Formosan subterranean termites, Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae), several rare sugars were tested for effects on the termites and symbionts. Among these, the D-galactose analog, 2-deoxy-D-galactose (2deoxyGal) showed promise as a potential control chemical. At a test concentration of 2deoxyGal (320.4 microg/mm3) in water applied to 5-cm filter paper, in bioassays with 20 termite workers, we found that worker termite mortality was significantly affected over a 2-wk period. Subsequent dose-mortality feeding studies confirmed these findings. In addition, consumption of the sugar-treated filter paper by termites caused a significant decrease in hindgut protozoan populations. 2deoxyGal caused dose-dependent termite mortality, taking on average 1 wk to begin killing workers, indicating that it may have promise as a delayed action toxin, which, if added to baits, could allow time after bait discovery for an entire colony to be affected.     

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.     

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.     

Interspecific Agonistic Behavior of <Emphasis Type="Italic">Macrotermes gilvus</Emphasis> (Isoptera: Termitidae): Implication on Termite Baiting in the Philippines

The study investigated interspecific agonistic behavior of Macrotermes gilvus Hagen (Isoptera: Termitidae: Macrotermitinae) against three economically important subterranean termites in the Philippines, viz., Coptotermes gestroi Wasmann, Nasutitermes luzonicus Oshima and Microcerotermes losbanosensis Oshima. Termite-termite interactions after a 1:1 pairing experiment showed that M. gilvus workers and soldiers were highly aggressive against C. gestroi, N. luzonicus and M. losbanosensis leading to severe injury or death of the opponent termite species in a short period of time. The levels of agonism were caste and species specific. Worker termites of M. gilvus showed an equally aggressive behavior as soldiers contributing to the high mortality of opponent species used in this study. It is likely that the highly aggressive behavior of M. gilvus limits foraging activity of C. gestroi, N. luzonicus and M. losbanosensis around in-ground bait stations contributing to the low success of termite baits containing chitin synthesis inhibitors in the Philippines.     

Escape Behavior of the Formosan Subterranean Termite (Isoptera: Rhinotermitidae) in Response to Disturbance

Disturbances to monitors and baits have often been noted as a confounding factor for the success of termite baiting systems. We studied escape behavior of Coptotermes formosanus Shiraki to a food source disturbance in the laboratory. Four peripheral chambers connected to the central food chamber through tubes served as escape routes from which escape patterns, escape and return times and differences in soldier and worker responses were recorded. We used two sand conditions: moist and dry to test whether favorable versus unfavorable conditions change escape behavior. The post disturbance responses were video recorded for 1 h. C. formosanus quickly escaped from the central chamber when disturbed. The time taken to reach the maximum number of termites that escaped ranged from 20 s to 210 s followed by a gradual return to the disturbed chamber. The majority of the escapees returned to the disturbed area within a few minutes after evacuation in both dry and wet sand conditions. A significantly higher percentage of termites escaped in wet sand arenas (43%) than in the dry sand arenas (31%). Our experiments demonstrate that C. formosanus is highly unlikely to abandon bait stations for extensive periods of time as a result of mechanical disturbances regardless of habitat quality.     

Bacteria from the Gut of Australian Termites

The major gut bacteria of the worker caste of nine species of Australian termites, belonging to four families, were isolated and identified to generic level. All species were either facultative anaerobes or strict aerobes. A correlation appears to exist between the major gut bacterium and the family to which the termite belongs. The major bacterium from the two lowest termites, Mastotermes darwiniensis (family Mastotermitidae) and Cryptotermes primus (family Kalotermitidae), was Streptococcus; from four species belonging to the Rhinotermitidae (Heterotermes ferox, Coptotermes acinaciformis, C. lacteus, Schedorhinotermes intermedius intermedius) it was Enterobacter; and from three species of the Termitidae (Nasutitermes exitiosus, N. graveolus, N. walkeri) it was Staphylococcus. Enterobacter was a minor symbiont of M. darwiniensis, C. primus, and N. graveolus; Streptococcus was a minor symbiont of H. ferox, C. lacteus, S. intermedius intermedius, and N. exitiosus; and Bacillus was a minor symbiont of C. acinaciformis and S. intermedius intermedius. M. darwiniensis possessed another minor symbiont tentatively identified as Flavobacterium. C. acinaciformis from three widely separated locations possessed a similar microbiota, indicating some form of control on the composition of the gut bacteria. Bacteria, capable of growth on N-free medium in the presence of nitrogen gas, were isolated from all termites, except N. exitiosus and N. walkeri, and were identified as Enterobacter. No cellulose-degrading bacteria were isolated.     

Termites Improve the Horizontal Movement of Carbonized Particles: A Step towards Sustainable Utilization of Biochar

Soil amendments containing carbonized materials increase the soil carbon reservoir, influence plant productivity, and, ultimately, help to clean the environment. There is data on the effect of such additions on soil physicochemical properties or plant growth, but few studies have focused on how these carbonized materials are distributed by termite species in the soil ecosystem. It is the first comprehensive study of the transportation of biochar (BC) by termite species under tropical environmental conditions in Pakistan. The present study was carried out to test the hypothesis that if termite species I) were involved in the distribution of biochar particles II) if yes, then how far these particles were transported during the study period (10 days) and III) check their preference between the enriched BC (EBC) and non-enriched BC. BC was enriched with the cattle slurry after its pyrolysis in the study. The results showed that EBC particles were significantly more widely distributed than non-enriched BC particles, but both types of BC were transported more than 4 cm (ring 4) within 10 days (at the end of the experiment). The current study also revealed that EBC was easily attached to the setae, cuticle, and legs of termites, implying that it could potentially be transported over a greater distance. Furthermore, transportation of EBC over larger distances indicated a potential preference of termite species between the EBC and BC particles. During the study, however, the preference among the termite species was also observed. Under the prevailing study conditions, the Coptotermes heimi and Heteroterme indicola species transported the EBC further than Microtermes obesi and Odontotermes obesus. These findings revealed that transportation preferences were observed among the four termite species. In conclusion, the current study found that termites were involved in the distribution of BC particles, with a preference for EBC and that these have the potential to transport BC particles more than 4 cm within 10 days. Furthermore, two species Coptotermes heimi and Heteroterme indicola may be more suitable candidates for EBC transpiration in Pakistani soils. It was necessary to conduct additional research into the effect of temperature on the transportation process.     

Studies on the Natural Durability of Tropical Timbers to termite attack

Small samples of 28 tropical timbers were tested with the dry-wood termite Cryptotermes cynocephalus and the subterranean termite Coptotermes curvignathus using several methods of testing. The results revealed that four among 28 species of wood were completely durable to the two species of termite. They are Dalbergia latifolia, Eusideroxylon zwageri, Intsia bijuga and Tectona grandis. Agathis alba, Dipterocarpus spp. and Mangifera spp. were classified as the most susceptible wood species to termites.     

白蚁纤维素粉饵料成型技术研究

为研究白蚁饵料成型工艺,比较了9种胶黏剂对白蚁纤维素饵料成型效果、耐水性能以及对白蚁取食的影响。结果表明:20%、40%剂量糊精;50%、100%剂量三聚氰胺甲醛树脂,10%、20%、40%田菁胶、卡拉胶、壳聚糖、明胶;10%、50%、100%聚乙烯醇、硅酸钠对微晶纤维素的成型效果较好,经上述剂量胶黏剂处理后,纤维素饵块的邵氏硬度(HD)极显著高于对照。耐水性能试验至第30天时,50%聚乙烯醇、100%聚乙烯醇、100%三聚氰胺甲醛树脂处理的纤维素饵块的溃散程度指数分别为1.33、1.00、2.00,其余饵块的溃散程度指数均达3级。生测结果显示,在7 d的室内强迫取食试验中,白蚁对50%聚乙烯醇、100%聚乙烯醇、100%三聚氰胺甲醛树脂处理的块状纤维素饵料的取食率均极显著低于对照,说明饵块中添加的上述胶黏剂对白蚁的取食具有一定的影响。综上,50%聚乙烯醇、100%聚乙烯醇、100%三聚氰胺甲醛树脂适用于白蚁纤维素饵料成型,但若想获得白蚁喜食的饵块仍需对配方做进一步的优化。