Movement efficiency and behavior of termites in tunnels with changing width

Subterranean termites build extensive underground galleries that consist of elaborate tunnels and channels to forage for food resources. The changes in tunnel width along the length of the tunnel are related to both biotic (e.g., termite activity) and abiotic factors (e.g., soil density). Termites transport food through the tunnels from food sources to their nest. Thus, understanding the relationship between traveling behavior in the tunnels and changing width is important to comprehend the stability of the termite ecosystem. In the present study, we explored the traveling behavior of termites in terms of movement efficiency, where the movement efficiency was defined as the time (τ) needed for a termite to pass through a tunnel. To do so, we designed artificial tunnels with linearly changing width in a two-dimensional arena. The tunnel widths, W ₁ (for the entrance) and W ₂ (for the exit), were 2, 3, 4, 5, and 6 mm. We systematically observed the traveling behavior of the termites Reticulitermes speratus kyushuensis Morimoto (Isoptera: Rhinotermitidae) in the artificial tunnels and measured τ. The value of τ increased with the increase of W ₂, regardless of W ₁. τ was longer in the case of W ₁ < W ₂ than that of W ₁ > W ₂. The experimental results can be explained by behavioral differences observed in each case. The implications of the findings are briefly discussed in relation to termite foraging efficiency and the development of individual-based models for the construction of termite tunnels.     

Measurement of the time required for termites to pass each other in tunnels of different curvatures

Subterranean termites construct complex tunnel networks for foraging. During travel in the tunnels, termites often encounter one another when passing in opposite directions. Such encounters are likely to affect the “movement efficiency,” which is the time required for a termite to travel a certain distance in a tunnel. In this study, we explored how individual–individual encounters affect movement efficiency in tunnels by measuring the time (τ) taken by two termites to pass one another in tunnels of different curvatures. Artificial tunnels of 5 cm in length and variable widths (W) of 2, 3, or 4 mm were made. Tunnel distance (D) was 2, 3, 4, or 5 cm. When D had a higher value, curvature was lower. When W = 2, τ was significantly shorter in the tunnel with D = 5 than in tunnels of D = 2, 3, or 4, whereas τ was statistically the same for D = 2, 3 and 4. When W = 3, τ was shorter in the tunnel with D = 5 than for D = 3 and 4, while τ was longer in the tunnel with D = 2 than for D = 3 and 4. When W = 4, τ was longer in the tunnels with D = 2 and 3 than for D = 4 and 5. Based on these observations, 3 types of termite behavior were identified: biased walking, backward walking, and zigzag walking. We considered these results in relation to foraging efficiency.     

Directional selection by the subterranean termite Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae) at tunnel intersections

Subterranean termites construct complicated tunnel networks for foraging below ground. Thus, they often encounter tunnel intersections during foraging activity. Directional selection by termites at intersections is likely to affect foraging efficiency because depending on their selection, the path length from food resources to the nest can vary significantly. In order to understand how termites are guided to choose the most economical path without the use of pheromones, we artificially excavated two tunnels of varying widths (W₁ and W₂; 2, 3, or 4 mm) that intersected at a 90° angle in each of nine soil‐filled arenas. We observed the directional selection of termites at intersections in arenas with combinations of W₁ (mm) and W₂ (mm) (W₁, W₂). For (W₁, W₂) = (2, 2) and (4, 4), termites statistically equally chose the three directions left, right, and straight, while for the (3, 3) arena combination, termites preferentially decided to go straight. On the other hand, for (W₁, W₂) = (2, 3) (2, 4), and (3, 4), termites advancing from narrower tunnels into intersections tended to turn right or left, while termites coming from the wider tunnel were more inclined to go straight. On the assumption that a wider tunnel is more frequently used by termites in the field, we can deduce that the wider tunnel may represent the most efficient/shortest path. Thus, this simple selection mechanism can prevent termites from deviating from the most economical path.     

Estimating termite population size using spatial statistics for termite tunnel patterns

Subterranean termites build underground tunnels for foraging. The obtained food is transported to the nest through these tunnels, and consumed to maintain the termite colony. In this process, termites can cause damage to wooden structures. To develop effective control strategies to reduce termite damage, it is important to know the sizes of the termite populations in the tunnels. In this study, we proposed a method for estimating the termite population size using the spatial statistic indices including fractal dimension (FD), local density (LD), and join count statistic (JCS) for the tunnel patterns. However, the method needs further improvement to be applied in field conditions. For the method, we generated 8,000 tunnel pattern images (1,000 images for each N) using an agent-based model based on experimental data. Here, N (= 3, 4, ..., 10) represents the number of termites participating in tunnel construction in the simulation. Subsequently, we calculated the FD, LD and JCS values of the tunnel pattern and trained and verified the k-nearest neighbors (KNN) algorithm, using 5,600 and 2,400 images, respectively. The population size (N) was estimated based on the FD, LD and JCS using the KNN algorithm. The estimated accuracy for all N was 60% to 97% in the range of k = 1 to 300. If the model for tunnel pattern generation includes heterogeneous environmental conditions, the proposed method could be used to effectively estimate the actual number of termite populations. Finally, we briefly discuss the challenges affecting our model, and how these could be overcome.     

Traveling behavior of a termite in tunnels with different curvatures and base surface roughness

This study focused on understanding the termite traveling behavior. Artificial tunnels with different curvatures and base surface roughness were constructed. Each tunnel was 50 mm in length, with widths of W (W = 2, 3, or 4 mm). The distance between the two ends of the tunnel was D (D = 20, 30, 40, or 50 mm). A higher value of D means a lower curvature. The roughness, R (R = 60, 120, 240 and ∞), was generated by uniformly sanding the substrate with a sanding machine. A higher R value means a finer base surface. The symbol ∞ represents a smooth surface. Time (τ) taken by a termite to pass through the tunnel was measured. τ was longer in tunnels with a smooth surface compared with the tunnels with a rough surface. When W = 2, there was no effect of the roughness on speed. This is because the narrow tunnel width diluted the effects of the smoothness of the surface. When W = 3, τ was statistically shorter for R = 240, than for R = 60 and 120. This suggests that an appropriate surface roughness could positively contribute to the traveling speed.     

Effect of landscape heterogeneity on termite tunnel pattern: Simulation study

Although the environment in which termites live is very heterogeneous, most experimental studies on the termite tunnel patterns have been conducted on homogeneous sand substrates. In order to explore how the heterogeneity affects tunnel patterns, I developed an agent-based model to simulate termite tunneling behavior at the individual level. In this model, grid space consists of easy and difficult areas for tunneling. Heterogeneity, H, was defined as the degree of the mixture of the two areas. The tunnel patterns formed by changing the number of termites, N, and H were quantitatively characterized by territory circularity and the territory area. These patterns were categorized into two groups, one with a small territory area and high circularity (group 1) and the other with a large area and low circularity (group 2). Considering the previous study that the termite populations with high N values have high territorial scalability, it can be said that the territories belonging to group 2 have higher foraging abilities and viability than those belonging to group 1. The simulation results showed that the tunnel patterns generated for small N and high H belonged to group 2. This implies that the heterogeneity can make a positive contribution to the expansion of the foraging area by effectively focusing the foraging energy of a termite population. I briefly discussed the mechanism of this positive role and the limitations of this simulation study. In addition, I discussed issues that need to be resolved in the near future to overcome the limitations.     

Why is the number of primary tunnels of the formosan subterranean termite,Coptotermes formosanus Shiraki (Isoptera: Rhinotermidae), restricted during foraging?

Subterranean termites forage by digging a network of tunnels to come into contact with food sources. When 1000 termites (Coptotermes formosanus Shiraki) were placed in a laboratory arena, 6.7 primary tunnels were constructed. The aim of this study was to explain the empirical observation in which termites restrict the number of primary tunnels. To this end, we constructed a model to simulate termite tunnel patterns based on empirical data and to calculate food transportation efficiency, γ, for the tunnel patterns. The efficiency was defined as the ratio of the number of encountered food particles to the sum of the shortest length from the location of encountered food particles to the initial position of growth of the tunnel. The γ was maximized when the number of primary tunnels was 5 or 6, which was fairly consistent with the empirical number of primary tunnels. This result indicated that termites may restrict the number of their primary tunnels to improve the transportation efficiency, which is directly related to their survival.     

Minimizing moving distance in deposition behavior of the subterranean termite

Subterranean termite nests are located underground and termites forage out by constructing tunnels to reach food resources, and tunneling behavior is critical in order to maximize the foraging efficiency. Excavation, transportation, and deposition behavior are involved in the tunneling, and termites have to move back and forth to do this. Although there are three sequential behaviors, excavation has been the focus of most previous studies. In this study, we investigated the deposition behavior of the Formosan subterranean termite, Coptotermes formosanus Shiraki, in experimental arenas having different widths (2, 3, and 4 mm), and characterized the function of deposited particles. We also simulated moving distance of the termites in different functions. Our results showed that total amounts of deposited particles were significantly higher in broad (4 mm width) than narrow (2 mm) tunnels and most deposited particles were observed near the tip of the tunnel regardless of tunnel widths. In addition, we found that deposited particles followed a quadratic decrease function, and simulation results showed that moving distance of termites in this function was the shortest. The quadratic decrease function of deposited particles in both experiment and simulation suggested that short moving distance in the decrease quadratic function is a strategy to minimize moving distance during the deposition behavior.     

Effects of concentration,distance, and application methods of Altriset (Chlorantraniliprole) on eastern subterranean termite (Isoptera: Rhinotermitidae)

The effects of various concentrations, distance, and application methods of Altriset (Chlorantraniliprole) were investigated against one of the most destructive termites, the eastern subterranean termite, Reticulitermes flavipes Kollar. Three laboratory experiments were conducted. First, we examined the concentration effect of treating the soil contiguously to established foraging tunnels at a fixed 1 m distance. The results demonstrated 100% termite control in 19 d posttreatment at 100 and 50 μg/g and 27% termite mortality at 25 μg/g. Second, we tested the distance effect of the soil treatment (2 and 4 m) on the efficacy of Altriset to the satellite termite populations at a fixed 50 μg/g concentration. This resulted in 100% termite control in 22 d posttreatment at both 2 and 4 m. Third, we examined the effect of differing application methods using 12.5 and 25 μg/g prior to the establishment of foraging tunnels at a fixed 1m distance. This illustrated 100% termite control in 9 d posttreatment at 25 μg/g and 12 d posttreatment at 12.5 μg/g. The third experiment demonstrated soil treatments that were applied prior to termite tunnel establishment had greater efficacy than applications made post tunnel construction. Our results provide a comprehensive understanding about the efficacy of Altriset treatments on eastern subterranean termites.     

Surface irregularity induced-tunneling behavior of the formosan subterranean termite

Subterranean termites forage from place to place by tunneling through soil. In order to examine termite-tunneling responses to external factors, we designed a square arena that contains five introduction chambers connected with narrow paths, which was filled with sand. Triangle-shaped indentation with width W and height H was provided as surface irregularity on the sand facing the introduction chambers by using templates. After termites were introduced into the chambers, we measured elapsed time, τ, for a tunnel to reach the point at 3 mm away from the apex of the irregularity. We found that for W = 0 mm (the absence of irregularity), termites (Coptotermes formosanus Shiraki) did not tunnel for 7 h, whereas, for W = 1, 2, 3, and 4 mm, they exhibited the tunneling behavior within 20 min. The result indicated that the presence of surface irregularity is essential to induce termite tunneling. In addition, we found that W was correlated with τ, whereas H did not influence τ. This was briefly discussed in the context of individual movement behavior.     

Tunneling response of termites to a pre-formed tunnel

Subterranean termites move from place to place while foraging by tunneling through soil. During a period of foraging, they are likely to encounter a number of pre-formed tunnels created by, for example, tree roots or the breaking up of a zone of hard or compacted soil. We systematically observed the behavioral response of tunneling termites to such pre-formed, artificially constructed tunnels at widths, W, of 0.5, 1.0, 1.5, and 2.0mm, which mimicked pre-formed tunnels in the field. The two tunnels intersected at an angle, theta (=0 degrees , 10 degrees , 20 degrees , 30 degrees , 40 degrees , 50 degrees , and 60 degrees ) formed between the advance direction of a termite tunnel and the perpendicular direction of a pre-formed tunnel. For W=Wc (=0.5mm) and theta相似文献   

Farmers’ perception of termites in agriculture production and their indigenous utilization in Northwest Benin

Background

Although termites are considered as agricultural pests, they play an important role in maintaining the ecosystem. Therefore, it matters to investigate the farmers’ perception of the impacts of the termites on the agriculture and their indigenous utilization.

Methods

A semi-structured questionnaire was used to interview 94 farmers through 10 villages of Atacora department, in the northwestern region of Benin, to obtain information for the development of successful strategies of termite management and conservation. Their perceptions on the importance and management of termites along with the indigenous nomenclature and utilization of termite mounds were assessed. Termite species identified by farmers were collected and preserved in 80% alcohol for identification.

Results

Eight crops were identified by farmers as susceptible to termites with maize, sorghum, and yam as being the most susceptible. According to farmers, the susceptibility to termites of these crops is due to their high-water content and sweet taste. A total of 27 vernacular names of termites were recorded corresponding to 10 species, Amitermes evuncifer, Macrotermes subhyalinus, and Trinervitermes oeconomus being the most damaging termite species. All the names given to termite species had a meaning. The drought was identified by farmers as the main factor favouring termite attacks. Demolition of termite mounds in the fields was the most commonly reported control method. Salt and other pesticides were commonly used by farmers to protect stored farm products. The lack of effective control methods is the main constraint for termite management. In northwestern Benin, farmers reported different purpose utilizations of termite mounds and termites.

Conclusions

The study has shown that farmers perceived termites as pests of several agricultural crops and apply various indigenous control practices whose efficiency need to be verified. Utilization of termites and termite mound soil as food and medicinal resources underlines the need for a more focused approach to termite control for the conservation of non-pest termite species. The sensitization of farmers on the importance of termites as well as the development of an integrated control method to combat termite pests proved necessary.

     

Intra- and Interspecific Comparisons of Bacterial Diversity and Community Structure Support Coevolution of Gut Microbiota and Termite Host

We investigated the bacterial gut microbiota from 32 colonies of wood-feeding termites, comprising four Microcerotermes species (Termitidae) and four Reticulitermes species (Rhinotermitidae), using terminal restriction fragment length polymorphism analysis and clonal analysis of 16S rRNA. The obtained molecular community profiles were compared statistically between individuals, colonies, locations, and species of termites. Both analyses revealed that the bacterial community structure was remarkably similar within each termite genus, with small but significant differences between sampling sites and/or termite species. In contrast, considerable differences were found between the two termite genera. Only one bacterial phylotype (defined with 97% sequence identity) was shared between the two termite genera, while 18% and 50% of the phylotypes were shared between two congeneric species in the genera Microcerotermes and Reticulitermes, respectively. Nevertheless, a phylogenetic analysis of 228 phylotypes from Microcerotermes spp. and 367 phylotypes from Reticulitermes spp. with other termite gut clones available in public databases demonstrated the monophyly of many phylotypes from distantly related termites. The monophyletic “termite clusters” comprised of phylotypes from more than one termite species were distributed among 15 bacterial phyla, including the novel candidate phyla TG2 and TG3. These termite clusters accounted for 95% of the 960 clones analyzed in this study. Moreover, the clusters in 12 phyla comprised phylotypes from more than one termite (sub)family, accounting for 75% of the analyzed clones. Our results suggest that the majority of gut bacteria are not allochthonous but are specific symbionts that have coevolved with termites and that their community structure is basically consistent within a genus of termites.     

Tunnel formation by Reticulitermes flavipes and Coptotermes formosanus (Isoptera: Rhinotermitidae) in response to wood in sand.

The tunneling responses of two subterranean termite species, Coptotermes formosanus Shiraki and Reticulitermes flavipes (Kollar), to the presence of sound wood in laboratory arenas were studied. Branching pattern and the speed of tunnel construction between R. flavipes and C. formosanus also were compared. Patlak's residence index (rho) was generated using the length, width, speed of construction, and area of the primary tunnels built by termites. In the same allotted time, C. formosanus built wider and shorter primary tunnels, whereas R. flavipes built thinner and longer primary tunnels. The presence of wood did not affect termite tunnel formation. This lack of variation in tunnel formation parameters was evidenced by the inability of the termites to locate wood sources over distance, even as short as 2.5 mm, and by the similar tunneling behaviors in areas of the arena with or without wood. Patlak's model predicted the densities of tunnels with an error between 9 and 28%. in experiments with R. flavipes exposed to a range of 0-8,000 g of wood, and between 61 and 87% in experiments with C. formosanus. These results indicated that the residence index can provide a qualitative measure of the effect of habitat heterogeneity on the individual termite tunnels. The tunneling constructions strategy of these subterranean termites is discussed.     

Tunnel geometry of the subterranean termite Reticulitermes grassei （Isoptera： Rhinotermitidae）in response to sand bulk density and the presence of food

The cryptic habits of subterranean termites restricts detailed analysis of their foraging patterns in situ, but the process is evidently dominated by tunnel constructions connecting the nest with woody resources discovered within the territory of each colony. In this study, tunnel formation and orientation were studied experimentally in the termite Reticulitermes grassei （Clement）, using 2-dimensional laboratory foraging arenas con- taining fine sand as the substratum. The building of exploratory tunnels over a 10-day period and the geometry of the resulting network are described. Fractal analysis showed that tunnel geometry had a fractal dimension, regardless of the total length tunnelled whether foragers encountered the food source or not. The bulk density of the sand in the arenas affected the distances tunnelled, with higher density reducing construction, but did not affect tunnel geometry. Tunnels were not discernibly orientated with respect to the positioning of the food source, even in a situation where termites had failed to find the food source at a distance of less than 50 mm, suggesting that volatiles from wood are not attractants.     

Hydrogen emission by three wood-feeding subterranean termite species (Isoptera: Rhinotermitidae): Production and characteristics

Abstract Hydrogen emission by wood-feeding termites, Coptotermes formosanus, Reticulitermes flavipes and Reticulitermes virginicus, was investigated upon a cellulosic substrate as their food source. The emission rates among the three species tested were significantly different and R. virginicus demonstrated the greatest H₂ emission at 4.78 ± 0.15 μmol/h/g body weight. In a sealed test apparatus, H₂ emission for each termite species showed a quick increase at the initial incubation hours (3–6 h), followed by a slower growth, possibly due to the feedback inhibition by gas accumulation. Further investigation revealed that continuous H₂ emission could be maintained by reducing the H₂ partial pressure in the sealed container. The bioconversion of cellulose to molecular H₂ by the subterranean termites tested could reach as high as 3 858 ± 294 μmol/g cellulose, suggesting that the termite gut system is unique and efficient in H₂ conversion from cellulosic substrate.     

Estimation of food composition of Hodotermes mossambicus (Isoptera: Hodotermitidae) based on observations and stable carbon isotope ratios

Abstract The diet of the harvester termite Hodotermes mossambicus was investigated at two sites with distinct dietary components: C₄ grasses (δ¹³C isotope values, ?13.8‰ to ?14.0‰) and C₃ plants (δ¹³C isotope values, ?25.6‰ to ?27.1‰). By comparing observations of food items carried into the colony by the termites and carbon isotope ratios of whole termites (that determined assimilated carbon), the relative proportion of the C₃ and C₄ plant food components of the termite diet was estimated. There was agreement between the observational data and stable carbon isotopic data, with grass representing approximately 93% of the diet of H. mossambicus at two study sites (urban and rural) on the South African highveld. However, when correcting for mass of food items, that is, C₃ and C₄, carried by termites, the proportion of grass (C₄) in the diet may be underestimated.     

Cloning and Heterologous Expression of Insecticidal-Protein-Encoding Genes from Photorhabdus luminescens TT01 in Enterobacter cloacae for Termite Control

Enterobacter cloacae, one of the indigenous gut bacteria of the Formosan subterranean termite (Coptotermes formosanus), was genetically modified with a transposon Tn5 vector containing genes (tcdA1 and tcdB1) encoding orally insecticidal proteins from the entomopathogenic bacterium Photorhabdus luminescens subsp. laumondii TT01, a symbiont of the entomopathogenic nematode Heterorhabditis bacteriophora, for termite control. In the laboratory, termites were fed filter paper inoculated with the recombinant bacteria. The chromosomal expression of the introduced genes showed that there were insecticidal activities against termite workers and soldiers challenged with the transformed bacteria. After termites were fed recombinant bacteria, the termite mortality was 3.3% at day 5, and it increased from 8.7% at day 9 to 93.3% at day 29. All the dead termites contained the recombinant bacteria in their guts. Transfer of the recombinant bacteria occurred between donor workers (initially fed recombinant bacteria) and recipient workers (not fed). More than 20% of the recipient termites ingested recombinant bacteria within 2 h, and 73.3% of them had ingested recombinant bacteria after 12 h. The method described here provides a useful alternative for sustainable control of the Formosan subterranean termite (C. formosanus) and other social insects, such as the imported red fire ant (Solenopsis invicta).     

Review of bees as guests in termite nests,with a new record of the communal bee, <Emphasis Type="Italic">Gaesochira obscura</Emphasis> (Smith, 1879) (Hymenoptera,Apidae), in nests of <Emphasis Type="Italic">Anoplotermes banksi</Emphasis> Emerson, 1925 (Isoptera,Termitidae, Apicotermitinae)

Associations between bees and termites are documented infrequently, but records are available for bee species ranging in behavior from solitary to highly eusocial. The subtribe Meliponina (stingless bees) is the most common bee group reported in termite nests, and, for some species, the occupancy of termite nests may be obligatory. The records of solitary bees nesting within termite nests include species of the tribes Emphorini, Centridini, Megachilini, and Paracolletini. Most of these bees can probably nest in other substrates, and their relationships with termite nests are presumably opportunistic. This study provides a review of published records of bees as guests in termite nests, and also describes the aggregation of nests of Gaesochira obscura within one nest of Anoplotermes banksi in Brazilian Amazonia. One termite nest contained at least ten nest entrances of G. obscura, with burrows 4–6 mm in diameter and about 10 cm long. Each burrow ended in brood cells in different stages of food provisioning and larval development. As commonly reported for other associations of this nature, there was no connection between the tunnels of bees and those of termites. This record adds important data on the biology of A. banksi. Because this is a single record, it is impossible to classify G. obscura either as a termitophile or termitariophile; this species may be opportunistic in relation to nesting substrate.     

Fipronil enhanced natural occurrence of Fusarium solani (Hypocreales: Nectriaceae) on building infesting termite Heterotermes indicola Wasmann (Blattodea: Rhinotermitidae)

The Southeast Asian subterranean termite, Heterotermes indicola Wasmann (Blattodea; Rhinotermitidae), is recognized as a building infesting lower termite species in urban environment. The extensive use of chemical termiticides against aerial mud tubes and underground nests of H. indicola beneath the buildings could not suppress its infestation; however, it enhanced the environmental contamination and insecticide resistance. In the present study, we tried to control termites using naturally occurring entomopathogenic fungi Fusarium solani (Mart.) Sacc., along with sublethal concentrations of termiticide fipronil in no-choice feeding pathogenecity bioassay for 20?days. Termite mortality after 20?days of continuous exposure to highest fungal treatment 1?×?10⁹?conidia/mL was 10% exclusively, whereas 100% mortality was calculated just after 16?days of concurrently exposure to 5?ppm of fipronil and highest rate of fungus 1?×?10⁹?conidia/mL. These results indicated that insecticidal stress declined the immune response of termites and reduced the repellency of termites against fungal conidia by breaching the primary defense mechanism (allogrooming). This co-application of F. solani at suitable sublethal concentrations of fipronil showed the promising potential against termites that may reduce the selection pressure of pesticides and resistance risk by targeted pests, but further investigations are necessary for developing field trials.