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The use of nonrepellent liquid termiticides against subterranean termites has long relied on the assumption that foraging termites in soils could transfer toxicants to nestmates to achieve population control. However, their dose-dependent lethal time can lead to rapid termite mortality in proximity of the treatment, triggering secondary repellency. The current study characterizes the dynamic nature of the "death zone," i.e. the area adjacent to soil termiticides that termites would avoid owing the accumulation of cadavers. Using whole subterranean termite laboratory colonies of Coptotermes gestroi (Wasmann) with 3 × 15 m foraging distances, fipronil was implemented at 1.5 m, 7.5 m, or 12.5 m away from colony central nests, emulating a corrective action against an termite structural infestation. For treatments at 7.5 m and 12.5 m, the death zone stabilized at an average of ~2.56 m away from the treatment after 40 d post-treatment, and colonies suffered as little as 1.5% mortality by 200 d post-treatment. Colonies located 1.5 m away from the treatment minimized the death zone to ~1.1 m and suffered as little as 23.5% mortality. Mortality only occurred within the first few days of treatment from initial exposure, as the rapid emergence of the death zone negated further transfer effects among nestmates over time. In some cases, foraging termites were trapped within the infested structure. While technically nonrepellent, fipronil becomes functionally repellent from the rapid mortality onset near the treatment. Even if diligently implemented to successfully protect structures, surrounding termite colonies are minimally impacted by fipronil soil treatments. [ABSTRACT FROM AUTHOR]

The article presents a study which investigated the efficiency of neem leaf extract on the control of wood termite, Coptotermes species, and gut cellulolytic bacteria. Topics discussed include isolation of cellulolytic bacteria from termite gut, identification of cellulolytic bacteria, and preparation of plant bio-pesticide.

The Asian subterranean termite, Coptotermes gestroi (Wasmann 1896) (Isoptera Rhinotermitidae), is a structural pest endemic to Southeast Asia that has been spread anthropogenically to much of the tropics worldwide. We report a land-based establishment of C. gestroi in Alexandria, Egypt, the northernmost locality of this species and over 4000 km from its nearest previous locality in northern India. An earlier study suggested that C. gestroi, misidentified as Reticulitermes lucifugus (Rossi 1792), was in Alexandria in 2010. We confirm C. gestroi in Alexandria based on soldier morphology and COII sequence data. We also report that Coptotermes alexandrinus Mohammad et Ramadan 2023, is a junior synonym of C. gestroi. The world distribution of C. gestroi is presented. [ABSTRACT FROM AUTHOR]

The effective monitoring and identification of existing subterranean termite populations within coarse woody debris and infested wood in service depend on accurate detection. These insects are often concealed within logs, wooden support structures, walls, and floorboards of buildings. In the absence of external mud tubes, termite infestations normally must be discovered through the destructive exploration of wooden structures to reveal the physical presence of these insect pests. Subterranean termite species are difficult to identify due to similarities in morphological features, but they may be readily distinguished by differences in volatile emissions from which they are divided into chemotaxonomic groups. Consequently, a more effective and nondestructive approach for detection and identification is to take advantage of unique species-specific emissions of volatile organic compounds (VOCs) from termite bodies which easily pass through wooden structures, allowing for detection without physical damage to wood and avoiding expensive DNA analysis. Electronic aroma detection analyses were conducted with an Aromascan A32S electronic-nose (e-nose) instrument, fitted with a 32-sensor conducting polymer (CP) sensor array, for discrimination between four common subterranean termite species based on differences in volatile emissions. Principal component analysis (PCA) of whole-body volatiles effectively distinguished between four termite species with the first two principal components accounting for more than 98% of sample variance (p < 0.01). Unique electronic aroma signature patterns (smellprints) were produced from e-nose sensor array outputs that allowed for the effective identification of termite species based on distinct differences in volatile metabolites released from their bodies. The e-nose methods were determined to be an improved means for rapidly detecting and monitoring termite species in wood. The method is cheaper than conventional detection methods and allows for the timelier discovery of species-specific termite infestations necessary for better management. The e-nose capability of detecting the Formosan termite in indoor living spaces was particularly significant due to the production of naphthalene, a volatile hazardous gas causing many adverse human health effects in enclosed environments. [ABSTRACT FROM AUTHOR]

The effective monitoring and identification of existing subterranean termite populations within coarse woody debris and infested wood in service depend on accurate detection. These insects are often concealed within logs, wooden support structures, walls, and floorboards of buildings. In the absence of external mud tubes, termite infestations normally must be discovered through the destructive exploration of wooden structures to reveal the physical presence of these insect pests. Subterranean termite species are difficult to identify due to similarities in morphological features, but they may be readily distinguished by differences in volatile emissions from which they are divided into chemotaxonomic groups. Consequently, a more effective and nondestructive approach for detection and identification is to take advantage of unique species-specific emissions of volatile organic compounds (VOCs) from termite bodies which easily pass through wooden structures, allowing for detection without physical damage to wood and avoiding expensive DNA analysis. Electronic aroma detection analyses were conducted with an Aromascan A32S electronic-nose (e-nose) instrument, fitted with a 32-sensor conducting polymer (CP) sensor array, for discrimination between four common subterranean termite species based on differences in volatile emissions. Principal component analysis (PCA) of whole-body volatiles effectively distinguished between four termite species with the first two principal components accounting for more than 98% of sample variance (p < 0.01). Unique electronic aroma signature patterns (smellprints) were produced from e-nose sensor array outputs that allowed for the effective identification of termite species based on distinct differences in volatile metabolites released from their bodies. The e-nose methods were determined to be an improved means for rapidly detecting and monitoring termite species in wood. The method is cheaper than conventional detection methods and allows for the timelier discovery of species-specific termite infestations necessary for better management. The e-nose capability of detecting the Formosan termite in indoor living spaces was particularly significant due to the production of naphthalene, a volatile hazardous gas causing many adverse human health effects in enclosed environments.

In eusocial organisms, cooperative brood care within a colony represents a situation where the ancestral parental care duties have shifted away from the reproductive parent(s) towards their offspring. The shift to alloparental care was often instrumental in the initial emergence of eusociality, as it ultimately contributed to the establishment of the reproductive division of labour.Remarkably, eusocial taxa such as ants and termites, which still display an ancestral independent colony foundation phase, must go through an obligatory parental care period, as a temporary subsocial family unit. In termites specifically, an incipient colony inherently remains a woodroach family unit until alloparental care is established. Colony foundation success can then be limited by a series of factors that may include environmental, behavioural, symbiotic and physiological constraints.In this study, 450 incipient termite colonies (Coptotermes gestroi) were established to investigate the timing of physiological changes in founders during the transition from biparental to alloparental care. Results showed that the finite initial internal nutritional resources that alates carry during the dispersal flight are a primary limiting factor for successful colony establishment. The Coptotermes queen and king must rapidly establish (<150 days) their first cohort of offspring to reach alloparental care or simply run out of resources and die. Alates, therefore, carry just enough internal resources to produce the first few alloparents (< 15 workers) to prime the system towards colony ergonomic growth, with a definitive shift to solely reproductive functions.Eusocial insect primary reproductive traits were optimized for three successive functions within the life cycle of a colony: alate dispersal (sexual reproduction), colony foundation (parental care) and colony growth (increased egg production towards colony maturity). However, results suggest that trade‐offs involving these functions appear to primarily favour dispersal ones (quantity vs. quality of alates), as founder(s) carry minimal resources and have no room for parental care inefficiency and as they then fully rely on their alloparents for further reproductive output.The transition towards alloparental care during colony foundation of eusocial insects may, therefore, reflect on the initial evolutionary transition from ancestral subsociality to eusociality. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Information regarding the species composition and dispersal flight season of termites is crucial for termite management. The major obstacles to collecting such information are a lack of access to private buildings and shortage of workers to monitor and report on termite swarming. To overcome these difficulties, we launched a citizen science project in which members of the public and pest management professionals were invited to collect termite samples. We created the website, Taiwan Termite Identification Service, on which populace could log the collection information, and ship termite samples to our laboratory for identification. We also established a Facebook group, called the "Termite Forum," to publicize this project. A total of 3024 samples were collected from 2015 to 2020, and we identified the species of >93% of the samples. Based on 1499 samples collected from buildings, five structural termite pests were identified, and species composition in each county of Taiwan is available. According to 844 dispersal flight events, termite dispersal flight timing peak and degree of centralization were estimated using a Gaussian model. The collected data demonstrated that the invasive termite species, Coptotermes gestroi (Wasmann) (Blattodea: Rhinotermitidae), continued northward expansion. The first intercepted alate of Schedorhinotermes sp. (Blattodea: Rhinotermitidae) indicated that it may be a new invasive pest from Southeast Asia. This study reports on a successful case of a citizen science project where urban pest data were collected on a national scale. [ABSTRACT FROM AUTHOR]

In 1995, the launch of the first commercial chitin synthesis inhibitor (CSI) bait led to the transformation of the subterranean termite control industry around the world. Their slow mode of action, which relies on both their ability to be transferred among nestmates and termite molting biology, has made them cost-effective solutions for subterranean termite colony elimination while minimizing the introduction of pesticides into the soil toward an environmentally sustainable strategy. However, despite successful commercial implementations, the acceptance of their use varies within the pest control industry around the world. Notably, the nuanced complexity of how CSI baits lead to colony elimination upon feeding by termite foragers has, in part, remained elusive for the past 3 decades, allowing for long-lasting misconceptions to persist. A recent series of studies has since provided complementary elements of understanding how CSI baits utilize termites' inherent colony demography, behavior, and physiology to trigger colony elimination after a characteristic succession of events within the colony collapse process. I here provide a synthetic overview of subterranean termite colony characteristics when exposed to CSI baits using Coptotermes (Wasmann) (Blattodea: Heterotermitidae) as a primary model system. The changes in colony demography through the colony collapse reflect how the mode of action of CSI baits makes them a prime solution for sustainable subterranean termite pest management. Following decades of innovation, ongoing interactions among termite researchers, bait product manufacturers, and pest management providers must continue to bring solutions to existing and emerging termite pest problems around the world., (© The Author(s) 2024. Published by Oxford University Press on behalf of Entomological Society of America.)

During the last ten years, the problem of subterranean termites in Alexandria, Egypt has worsened, causing great economic losses. Throughout our study to survey the termite species in Alexandria, a species was found that differed from any of the previously recorded ones in Egypt. The aim of this study was to identify this invasive subterranean species in the region. Samples were collected from infested buildings and trees, and their morphological characteristics along with their DNA barcoding loci (COII, 18S and 28S) were determined. Moreover, phylogenetic comparison with other related species were also considered for the most accurate identification. The analysis of all the previously mentioned parameters proved that it is Coptotermes sp. This genus is identified and recorded for the first time in Egypt and North Africa. This study demonstrates that the combination of molecular and morphological methods is essential for accurate species identification, which aids in the correct control of destructive pests. [ABSTRACT FROM AUTHOR]

Aims: Arthropods guts, such as termite harbor diverse microorganisms including those that are capable of fixing atmospheric nitrogen (N2). Nitrogen-fixing bacteria can help termite to overcome their shortage of dietary N by providing fixed N2. Nitrogenase enzyme is responsible for this trait and encoded by nif genes which are highly conserved and are primarily used in the identification of N2-fixing microorganisms. Here, we characterized N2-fixing bacteria isolated from the hindguts of termite Coptotermes gestroi. Methodology and results: A total of 46 bacterial isolates were obtained after a primary screening based on their ability to grow on Burk's media. Subsequently, the nifH gene from two of these isolates, namely S7 and S20, were successfully amplified and sequenced. Molecular phylogenetic analysis of 16S rRNA gene sequence revealed that isolate S7 is closely related to Ralstonia pickettii ATCC27511 (99.34% similarity, 1059 bp), whereas isolate S20 is closely related to Microbacterium sp. NCCP-451 (LC488936) (99.06% similarity, 948 bp). Besides that, the recA gene of isolate S7 is closely related to Ralstonia pickettii 12D (CP001644) (100% similarity, 442 bp) and the type strain of Ralstonia pickettii (ATCC 27511) (NZ KN050646) (98.97% similarity, 438 bp). Meanwhile, nifH gene of isolate S7 showed highest similarity to the uncultured bacterium NR1606 (AF035490) (99.93% similarity, 277 bp). Moreover, the nifH gene of isolate S20 is clearly separated from Azoarcus sp. and distantly related to Microbacterium sp. The incongruence between the partial 16S rRNA and nifH gene sequences could indicate the possibility of horizontal transfer of nif genes. Conclusion, significance and impact of study: The phylogenetic incongruence between housekeeping genes (16S rRNA and RecA) and nifH gene in these bacteria provides new insight on potential horizontal gene transfer (HGT) activity taking place in bacterial communities particularly in the guts of arthropods. The finding of this study on potential HGT can also aid in the prediction of origins and evolution of gene transfer among bacteria. [ABSTRACT FROM AUTHOR]

All termites have established a wide range of associations with symbiotic microbes in their guts. Some termite species are also associated with microbes that grow in their nests, but the prevalence of these associations remains largely unknown. Here, we studied the bacterial communities associated with the termites and galleries of three wood-feeding termite species by using 16S rRNA gene amplicon sequencing. We found that the compositions of bacterial communities among termite bodies, termite galleries, and control wood fragments devoid of termite activities differ in a species-specific manner. Termite galleries were enriched in bacterial operational taxonomic units (OTUs) belonging to Rhizobiales and Actinobacteria, which were often shared by several termite species. The abundance of several bacterial OTUs, such as Bacillus, Clostridium, Corynebacterium, and Staphylococcus, was reduced in termite galleries. Our results demonstrate that both termite guts and termite galleries harbor unique bacterial communities. [ABSTRACT FROM AUTHOR]

Background: Coptotermes sjostedti Holmgren (Isoptera:Rhinotermitidae) is one of the major termite species found infesting woods and other plant materials in Africa. Bioassays were carried out at the Arboretum of the Department of Forestry and Wildlife Resources Management, University of Calabar, Nigeria to investigate the effect of oils from the kernel of Jatropha curcas L. and Azadirachta indica oils A. Juss against the African subterranean termite (Coptotermes sjostedti Holmgren). Treatments comprised of the plant oils and Solignum (a synthetic termiticide, serving as positive control) applied at the rate of 5, 10 and 15 mL per 100 cm3 of the wood from seven plant species. Untreated woods also served as control. Seven wood species were placed in a test arena and artificially infested with 50 termites per unit. The setup was a randomized complete block design with four replicates. Data on termite mortality and wood weight loss were recorded. Results: The use of 10 and 15 mL of J. curcas oil caused more than 90% mortality of C. sjostedti, after 96 h, kept wood consumption by the insect below 6.0% after 3 months, and these outcomes were similar to that of Solignum at the same concentrations. About 35.0–65.2% reduction in weight was observed among untreated woods from the tested woods, indicating their susceptibility to C. sjostedti attacks. Correlation analysis shows a significant (r > − 0.900; P < 0.0001) negative association between termite mortality and the rate of wood consumption. Conclusion: The use of J. curcas oil at 10 mL per 100 cm3 of wood could be a potent alternative to Solignum for the control of C. sjostedti. [ABSTRACT FROM AUTHOR]

Microbial communities in the immediate environment of socialized invertebrates can help to suppress pathogens, in part by synthesizing bioactive natural products. Here we characterized the core microbiomes of three termite species (genus Coptotermes) and their nest material to gain more insight into the diversity of termite-associated bacteria. Sampling a healthy termite colony over time implicated a consolidated and highly stable microbiome, pointing toward the fact that beneficial bacterial phyla play a major role in termite fitness. In contrast, there was a significant shift in the composition of the core microbiome in one nest during a fungal infection, affecting the abundance of well-characterized Streptomyces species (phylum Actinobacteria) as well as less-studied bacterial phyla such as Acidobacteria. High-throughput cultivation in microplates was implemented to isolate and identify these less-studied bacterial phylogenetic group. Amplicon sequencing confirmed that our method maintained the bacterial diversity of the environmental samples, enabling the isolation of novel Acidobacteriaceae and expanding the list of cultivated species to include two strains that may define new species within the genera Terracidiphilus and Acidobacterium.

Microbial communities in the immediate environment of socialized invertebrates can help to suppress pathogens, in part by synthesizing bioactive natural products. Here we characterized the core microbiomes of three termite species (genus Coptotermes) and their nest material to gain more insight into the diversity of termite-associated bacteria. Sampling a healthy termite colony over time implicated a consolidated and highly stable microbiome, pointing toward the fact that beneficial bacterial phyla play a major role in termite fitness. In contrast, there was a significant shift in the composition of the core microbiome in one nest during a fungal infection, affecting the abundance of well-characterized Streptomyces species (phylum Actinobacteria) as well as less-studied bacterial phyla such as Acidobacteria. High-throughput cultivation in microplates was implemented to isolate and identify these less-studied bacterial phylogenetic group. Amplicon sequencing confirmed that our method maintained the bacterial diversity of the environmental samples, enabling the isolation of novel Acidobacteriaceae and expanding the list of cultivated species to include two strains that may define new species within the genera Terracidiphilus and Acidobacterium. [ABSTRACT FROM AUTHOR]

The Formosan subterranean termite, Coptotermes formosanus Shiraki, 1909, is a highly destructive structural pest endemic to East Asia. We report a land-based establishment of C. formosanus in Petah Tikva, Israel, over 6000 km from its nearest previous locality in China. The species' identity was confirmed by soldier morphology and by COI sequence data. In addition, a population discovered in 1992 in suburban San Diego, California, USA, remains viable. Marathon hosts the first infestation of C. formosanus in the Florida Keys. The world distribution of C. formosanus is presented, and the biogeography and mechanisms of spread of this termite are discussed. [ABSTRACT FROM AUTHOR]

Cellulose and lignocellulose, are abundantly available biomass, derived from plant sources. Apart from this, lignocellulose, is also known to be the main component of municipal solid waste, which needs bioremediation. Because of their bioethanol production capabilities, these polysaccharides, may also be used as renewable energy sources, replacing the fossil fuels. Biodegradation of cellulose and lignocelluloses to simpler sugars and ethanol production may be processed by an important enzyme, cellulase. This enzyme, occurs naturally in the gut of isopteran insects, termites. The gut microbiota, of these white ants synthesize cellulase and are capable of digesting cellulose and lignocelluose. Cellulases derived from microbes of termites, are considered to be natural and cost effective, enzyme reserves. In this context, the present study, was attempted to extract cellulase from gut bacteria of termites. White ants, were collected from wood, and their gut microbes, were isolated, identified and the cellulase was extracted. The bacterial species which was isolated in the present study was identified as Pseudomonas aeruginosa. Gram's staining and different biochemical tests, were performed, to ascertain its identity. The cellulase enzyme, which was extracted was partially purified, by the method of ammonium sulphate precipitation. This study, reiterates, that termites, may be considered as the best economical source for cellulase extraction, which can be used for degradation of cellulose and lignocelluloses, and also for bioethanol production. [ABSTRACT FROM AUTHOR]

Congeneric species that live in sympatry may have evolved various mechanisms that maintain reproductive isolation among species. However, with the spread of invasive organisms owing to increased global human activity, some species that evolved in allopatry can now be found outside their native range and may have the opportunity to interact, in the absence of mechanisms for reproductive isolation. In South Florida, where the Asian subterranean termite, Coptotermes gestroi (Wamann), and the Formosan subterranean termite, Coptotermes formosanus Shiraki (Blattodea: Rhinotermitidae) are invasive, the two species can engage in heterospecific mating behavior as their distribution range and their dispersal flight season both overlap. Termites rely on semiochemicals for many of their activities, including finding a mate after a dispersal flight. In this study, we showed that females of both species produce (3Z,6Z,8E)-dodeca-3,6,8-trien-1-ol (DTE) from their tergal glands as a shared sex pheromone. We suggest that both species primarily rely on an inundative dispersal flight strategy to find a mate, and that DTE is used as a short distance pheromone or contact pheromone to initiate and maintain the tandem between males and females. The preference of C. gestroi males for C. formosanus females during tandem resulted from the relatively high amount of DTE produced by tergal glands of C. formosanus females, when compared with those of C. gestroi females. This results in confusion of mating in the field during simultaneous dispersal flights, with a potential for hybridization. Such observations imply that no prezygotic barriers emerged while the two species evolved in allopatry for ~18 Ma. [ABSTRACT FROM AUTHOR]

Entomopathogenic fungi have the potential to infect most arthropods including termites which are economically important major insect pests of wood, wood products and building structures. However, the application of this fungus in the field has not shown satisfactory results yet, one of which is constrained in mass production of conidia. The purpose of this study was to evaluate 16 types of biodegradable products and waste as substrates for mass production of conidia using solid state fermentation method and two types of inoculum (solid and liquid inoculum). Toxicity tests were carried out on subterranean termites (Coptotermes sp.) based on JIS K 1571, 2010. The parameters observed were the number and dry weight of the conidia produced, conidial viability, nutrient content of the substrate, and percentage of termite mortality. The results showed that rice, sorghum and corn were the best media for the growth of entomopathogenic fungi based on the number of conidia and dry weight of the conidia produced. Metarhizium sp. T4.B23 produced the highest number of conidia, 1.12 x 1011 conidia/100 g substrate and yielded 180.9 ± 0.623 g dry conidia/kg of rice; followed by Metarhizium sp. B2.2 grown on sorghum that resulted in 1.11 x 1010 conidia/100 g substrates and 127.8 ± 0.570 g/kg sorghum; and B. bassiana produced 8.3 x 109 conidia/100 g substrate and 31.24 ± 0.407 g/kg sorghum. Metarhizium sp. B2.2 showed the highest toxicity to termites with 100% mortality that was observed within the second day of testing. The novelty of this research is we found two strains of Metarhizium indigenous Indonesia (T4.B23 and B2.2) that isolated from soil in Wangameti, Sumba, East Nusa Tenggara and Mount Gandang Dewata, West Sulawesi, respectively. In this study we not only can produce the high number of conidia from these fungi using cheap media but also it has high toxicity to termites. Furthermore, the conidia of these fungi are very potential to be developed as a biopesticide for termite control, so the use of chemical pesticides can be reduced. [ABSTRACT FROM AUTHOR]

Termites feed on a carbon-rich but nitrogen-poor diet and evolved efficient nitrogen conservation strategies. It was previously suggested that during time of low access to resources (i.e., starvation), subterranean termites (Rhinotermitidae) colonies would adopt an energy conservation strategy by cannibalizing soldiers, a dependent caste. However, such hypothesis was tested with relatively small groups of foragers, which may not have reflected how resources are reallocated in whole colonies when food (carbon) is scarce. The current study subjected 2-year-old colonies (≈ 3000 termites with all castes and instars) to starvation, in order to reexamine if cannibalism is part of an active energy conservation strategy. Within 12 days of starvation, eggs and larvae were all cannibalized, followed by young workers. Soldiers then died marginally faster than old workers. By 22 days, cannibalism was no longer observed, and the king and queen were among the last individuals to die. Termites that engaged in the process of ecdysis with no energetic resources, failed to molt, died in the process, and the subsequent cannibalism was a passive mortality-driven process. Therefore, cannibalism was primarily the result of the inherent termite behavior for recycling nitrogen resources. The hemimetabolous developmental pathway, the difference of timing in the molting cycle of termite instars and the relatively rapid exhaustion of soldiers, indirectly determined the sequence of starvation-induced mortality and subsequent cannibalism. Although termites have evolved efficient nitrogen conservation strategies, they have not evolved an efficient carbon conservation strategy, which is essentially limited to keeping the primary reproductives alive as long as possible. [ABSTRACT FROM AUTHOR]

In termites, primary reproductives proceed through dispersal flight with finite metabolic reserves. During colony foundation, the mated pair requires just enough resources to produce the first few workers, through biparental care. When alloparental care is established, the primary reproductives are fully supported by their own offspring. It was, therefore, argued that the pressure to accumulate large quantities of metabolic reserves in termite imagoes was relaxed over evolutionary time, which resulted in a progressive reduction in alate size compared to ancestral wood roaches. However, such directional reduction in size may partially be countered if mated pairs with relatively large internal metabolic resources are more successful than mated pairs with relatively low internal resources. This hypothesis was tested with Coptotermes gestroi by establishing incipient termite colonies with a wide range of combinations of female and male initial weights in laboratory conditions. Both females and males depleted most of their internal resources within 9 months, and the combined initial weight of female and male explained 27% of the variation in incipient colony growth. Mature colonies that can invest into high-quality alates may have a slight fitness advantage; however, this advantage may be secondary to other environmental factors, as during large Coptotermes dispersal flights, a vast majority of alates die within the first few days, which would increase the fitness of colonies that invested in quantity over quality. Within a given termite species, the relative size of imagoes may, therefore, reflect a reproductive strategy trade-off emerging from the life history of the species. [ABSTRACT FROM AUTHOR]

A corrigendum for the article "Ecological diversification of the Australian Coptotermes termites and the evolution of mound building" is presented with the modification of two pairs of environmental variables.

Abstract This study evaluated the termite susceptibility of Papua New Guinea (PNG) balsa (Ochroma pyramidale) against Australia's most economically important wood-destroying termite species, Coptotermes acinaciformis. Samples of envelope treated PNG balsa with the insecticide bifenthrin and non-treated balsa, radiata pine (Pinus radiata) sapwood and southern blue gum (Eucalyptus globulus) heartwood were exposed to seven different colonies of C. acinaciformis at a field site in the Northern Territory, Australia. The results indicated that balsa is susceptible to termite attack, with a mean mass loss of 30.9% (range 3.8–99.9%). Mean mass losses for radiata pine and southern blue gum were 90.3 and 98.9% respectively. It was evident that balsa can be successfully protected from termite attack with envelope-treatments of bifenthrin, with the latter proving to be highly efficacious at mass/volume retentions much lower than those specified in AS 1604.1 and typically used in the Australian timber preservation industry. Highlights • PNG balsa is susceptible to wood-destroying termite attack in Australia, with a mean mass loss of 30.9% (range 3.9–99.9%). • Similarly, as a control in the study, the mean mass losses for radiata pine and southern blue gum were 90.3% and 98.9%. • Balsa can be successfully protected from termite attack with envelope-treatments of the insecticide bifenthrin. • The paper reviews the literature for balsa susceptibility to termite species and details the origin of balsa and methods used. [ABSTRACT FROM AUTHOR]

This study examined the tunneling activity and the behaviour of two subterranean termites, Globitermes sulphurues and Coptotermes gestroi treated with four plant extracts, namely Pyllanthus niruri, Azadirachta indica, Leucaena leucocephala and Andrographis paniculata. All plants were extracted with three different solvents (methanol, hexane and water) and were diluted into three concentrations (500, 5,000 and 10,000 ppm). A group of 50 workers and 2 soldiers were tested and observed daily during the exposure in order to determine their survivorship and behaviour. Both sections were scored to determine their tunneling activities after seven days. There was a significant difference of tunneling activities of C. gestroi on sand treated with plant extracts (X²(2) = 31.790, p < 0.0001) with a mean rank of 8.50 for methanolic extracts and 32.50 for both hexane and water extracts. Meanwhile, no significant difference was observed on tunneling activity of G. sulphureus on treated sand (X²(2) = 2.200, p = 0.333) with a mean rank of 20.72 for methanolic extracts, 26.31 for water extracts and 26.47 for hexane extracts. Results showed that plants extracted with methanol demonstrated strong repellent properties with 0 tunneling activity on the treated sand and low survivorship of both termites. Moreover, both termites did not survive (0%) when they were treated with methanolic extracts at 10,000 ppm. They also displayed a different behaviour post-exposure such as avoidance, gradually losing the ability to walk and low feeding consumption. These results indicate that there is a strong termiticidal activity of plants extracted with methanol against C. gestroi and G. sulphureus. [ABSTRACT FROM AUTHOR]

The presence of biofertilizers can affect the life and movement of subterranean organisms in the soil. In the current study, the effect of three biofertilizers (Unigrow, Rhizogold and Rhizogold Plus) alone and in combination with diatomaceous earth (DE) and insecticides on mortality and movement of subterranean termites, Coptotermes heimi (Wasmann), was investigated under laboratory conditions. The mortality of termites in soil mixed with biofertilizers was low as compared to the control treatment, which led to significant mortality. Length of galleries formed by termites with the addition of Unigrow in the soil was significantly higher as compared to the other two biofertilizers and control treatments. Mortality increased in the presence of insecticides and DE in the soil substrate. Unigrow mixed soil having chlorfenapyr had maximum mortality and minimum gallery length followed by Rhizogold and Rhizogold Plus. The addition of DE and insecticide further enhances mortality, and we recommend that some toxic elements should be incorporated in biofertilizers to control subterranean termites. [ABSTRACT FROM AUTHOR]

Amongst the termite genera within Rhinotermitidae, the genus Coptotermes is probably regarded as one of the most destructive pest in Malaysia as it contributes to more than 90% of the damages in buildings and structures. In this study, 17 parameters were used for morphometric analysis. Partial sequence of mitochondrial DNA cytochrome oxidase subunit II (COII) was obtained from 21 populations of C. curvignathus, seven populations of C. sepangensis and two populations of C. kalshoveni in Sarawak. In addition, 11 sequences of Coptotermes spp. were obtained from GenBank and included in the analysis. Based on Discriminant Function Analyses (DFA), the maximum width of postmentum (MxWPt) is identified as the best morphological character for differentiating the four species of Coptotermes in this study, proved by canonical discriminant function. Phylogenetic analyses of COII found that C. sepangensis and C. kalshoveni are nested within a single clade. Morphological comparison also showed these two species to be identical in terms of body measurements and diagnostic features. These evidence indicates that C. sepangensis and C. kalshoveni are the same species, thus we suggest C. sepangensis is a junior synonym of C. kalshoveni. [ABSTRACT FROM AUTHOR]

Mutualistic associations between insects and microorganisms must imply gains for both partners, and the emphasis has mostly focused on coevolved host–symbiont systems. However, some insect hosts may have evolved traits that allow for various means of association with opportunistic microbial communities, especially when the microbes are omnipresent in their environment. It was previously shown that colonies of the subterranean termite Coptotermes formosanus Shiraki (Blattodea: Rhinotermitidae) build nests out of fecal material that host a community of Streptomyces Waksman and Henrici (Actinomycetales: Streptomycetaceae). These Actinobacteria produce an array of bioactive metabolites that provides a level of protection for termites against certain entomopathogenic fungi. How C. formosanus acquires and maintains this association remains unknown. This study shows that the majority of Streptomyces isolates found in field termite fecal nest materials are identical to Streptomyces isolates from soils surrounding the nests and are not vertically inherited. A survey of Streptomyces communities from C. formosanus fecal nest materials sampled at 20 locations around the world revealed that all nests are reliably associated with a diverse Streptomyces community. The C. formosanus fecal nest material therefore provides a nutritional framework that can recruit beneficial Streptomyces from the soil environment, in the absence of long-term coevolutionary processes. A diverse Streptomyces community is reliably present in soils, and subterranean termite colonies can acquire such facultative symbionts each social cycle into their fecal nest. This association probably emerged as an exaptation from the existing termite nest structure and benefits both the termite and the opportunistic colonizing bacteria. [ABSTRACT FROM AUTHOR]

Research Highlights: Tree size and wood characteristics influenced the susceptibility of five Amazonian timber tree species to heartwood decay and colonization by termites. Termites occurred in the heartwoods of 43% of the trees, with Coptotermes testaceus the most abundant species. Background and Objectives: Hollows and rotten cores in the stems of living trees have ecological and economic impacts in forests managed for timber. The decision on whether to cut or maintain hollow trees in such forests must account for the susceptibility of different tree species to decay. We investigated tree and wood characteristics of living trees of five commercial timber species in the eastern Amazon that influenced the likelihood of heartwood decay and the occurrence of termite nests inside the rotten cores. Materials and Methods: We used Pearson’s correlations and one-way analysis of variance (ANOVA) to explore relationships among tree basal area and hollow area. We used principal components analysis (PCA) to analyze the variation of wood anatomical traits, followed by a linear regression to explore the relationships between PCA scores, and heartwood hollow area. We used a logistic model to investigate if the probability the occurrence of colonies of C. testaceus inside tree cores varied with tree and species characteristics. Results: Heartwood hollow areas increased with stem basal area. Larger hollows were more likely to occur in species with higher vessel and ray densities, and smaller diameter vessels. Termites occurred in the hollows of 43% of the trees sampled, with C. testaceus the most common (76%). The probability of encountering termite nests of C. testaceus varied among tree species and was positively related to wood density. Conclusions: This study shows that given the increased likelihood of stem hollows and rotten cores in large trees, tree selection criteria in managed tropical forests should include maximum cutting sizes that vary with the susceptibility of different tree species to stem decay.

Surveys of the conditions of termite attack were conducted in two regencies, Pidie and Greater Aceh, Aceh Province, Indonesia (40 houses in each location). Interviews were also conducted with home owners to collect data on the building history; culture, such as daily life in the house; the frequency and intensity of termite attacks; and traditional knowledge for avoiding and/or suppressing termite attacks. We found that 51% of traditional houses were infested by two termite species: Coptotermes gestroi and Nasutitermes matangensis. The lower parts of traditional houses were frequently attacked and severely damaged by termites. Previous land use and the ages of the traditional houses affected the intensity of the termite attacks. Several measures for avoiding and/or suppressing termite attacks on cultural heritage buildings are also proposed.

Subterranean termites often transport soil into bait stations. In this study, we hypothesize that adding soil or clay material in the bait may affect preference and performance of termites. Choice and no-choice tests were conducted in the laboratory to investigate the aggregation and feeding preference, survivorship, wood consumption, and body water percentage of termites in response to food containers (here we simulated the bait station by placing a wood block into a bigger plastic box with termite-entering holes on the wall) with the void space filled with soil (sandy clay loam), clay material (sodium bentonite), or remained unfilled. Choice tests showed that under low-moisture conditions, food containers filled with clay attracted significantly more termites (Coptotermes formosanus Shiraki (Blattodea: Rhinotermitidae) and Reticulitermes guangzhouensis Ping (Blattodea: Rhinotermitidae)) compared to food containers filled with soil, or unfilled. Under medium-moisture conditions, however, the percentages of termites that aggregated in the food containers filled with soil or clay were similar, and both were significant higher than the percentages in unfilled ones. In no-choice tests, the highest survivorship and wood consumption in C. formosanus were recorded under medium-moisture conditions and when food containers were filled with clay, whereas the lowest survivorship and wood consumption were recorded under low-moisture conditions and when food containers were filled with soil. Interestingly, presence of clay increased the body water percentage of termites. Our study enhances the understanding of the foraging ecology of subterranean termites, and may contribute to the improvement of termite-baiting technologies. [ABSTRACT FROM AUTHOR]

This study evaluated the impact of a non-repellent liquid termiticide (fipronil) and a chitin synthesis inhibitor (CSI) termite bait (noviflumuron) on whole colonies of Coptotermes gestroi (Wasmann) (Blattodea: Isoptera: Rhinotermitidae) in laboratory conditions, over a 12-m foraging distance. The protocol simulated the implementation of remedial treatment of an infested structure, where the colony has multiple access routes, and where only a portion of the population was directly exposed to the treatment. Within 2 wk after the implementation of fipronil, all termites within 1.5 m away from the treatment died. The accumulation of cadavers near the treated area resulted in secondary repellency and the colonies avoided the treated area for the remaining 10 wk of the experiment, using alternative foraging galleries. At the end of the 12 wk, colonies exposed to fipronil did not have any difference in population size compared with control colonies. Comparatively, colonies exposed to noviflumuron had no change in foraging activity for the first ≈40 d, but then termites progressively ceased their activity throughout their foraging territory. By 12 wk, noviflumuron-exposed colonies were near-elimination, with only a few workers, soldiers remaining, and all colonies were eliminated by 95 d. This study shows that subterranean termite colonies with access to CSI baits are inevitably eliminated, regardless of the position of the bait, while colonies exposed to fipronil are only locally excluded from the area near the treatment, but may maintain their foraging activity in untreated areas, and retain their potential risk for structural damage in the long term. [ABSTRACT FROM AUTHOR]

Termites and their nests are potential resources for a wide assemblage of taxa including nematodes. During dispersal flight events from termite colonies, co-occurring nematodes in the nest may have phoretic opportunities to use termite alates as transportation hosts. The two subterranean termite species Coptotermes gestroi (Wasmann) and Coptotermes formosanus Shiraki are both invasive and established in south Florida. Alates of both species (n = 245) were collected during dispersal flight events in 2015–2016 from six locations, of which 30 (12.2%) were associated with one or more species of nematodes. Species of Bunonema Jägerskiöld (Rhabditida: Bunonematidae), Halicephalobus Timm (Rhabditda: Panagrolaimidae), and Poikilolaimus regenfussi (Sudhaus) Sudhaus and Koch (Rhabditida: Rhabditidae) were isolated from 5.3, 4.9, and 0.4% of termite alates, respectively, and Bunonema and Halicephalobus were concomitant in 1.6% of alates. Additional C. formosanus alates were field-collected to establish laboratory colonies in sterilized rearing containers (SRC) to determine if alate-associated nematodes would colonize the newly established nest and/or brood. Among 1-yr-old termite colonies reared in SRCs, 26.9% of the colonies were positive for nematodes confirming that within-colony transmission of nematodes occurred. All three isolated nematode genera are free-living bacterivores capable of asexual reproduction. This suggests that these common co-occurring, termite-associated nematodes are opportunistic and facultative symbionts that receive increased opportunities of geographical dispersion through phoresy during termite dispersal flight events. [ABSTRACT FROM AUTHOR]

Abstract: Coptotermes gestroi forages on wooden structures and causes substantial damage in urban areas of Brazil. The survival of foragers may be influenced by either group size or caste composition. It was hypothesised that workers survive at higher rates in larger groups and consequently exhibit higher rates of food consumption and that soldiers represent a nutritional resource for foragers during starvation periods. To test the first hypothesis, groups of 100, 200 and 400 workers were placed in experimental arenas containing wood as food. After 28 days, the units were evaluated and the number of living termites and wood consumption were estimated and compared. An analysis of the data found that there were no significant differences within the group sizes on termite survival but that relative wood consumption was significantly higher in the smaller groups. To test the second hypothesis, groups containing workers and soldiers (9:1 ratio) or only workers were placed in experimental arenas with food present or absent. Survival for each caste was monitored at 3 day intervals over 9 days to estimate the cannibalism rate. Cannibalism was significantly higher in the starved groups, but the presence of soldiers did not influence the rate of cannibalism. Colony fragmentation and starvation are stressful conditions that lead foragers of C. gestroi to adaptive behaviours in order to enhance their survival. [ABSTRACT FROM AUTHOR]

Various methods for termite detection have been developed, one of which is purely based on their acoustic signals. However, this method has a weakness, as it is difficult to separate the signals generated by the termites from noise in the environment. A combination of the feature extraction of the acoustic signals and a classification model is expected to overcome this weakness. In this investigation, we inserted 220 subterranean termites Coptotermes curvignathus into pine wood for feeding activity and observed their acoustic signals. In addition, three acoustic features (shortterm energy, entropy and zero moment power) were proposed to recognize the termites' acoustic signals. Subsequently, these features were analyzed and combined with discriminant analysis to produce a robust classification model. According to the numerical results, the integrated discriminant analysis and the acoustic feature in our termite detection system has an accuracy of 83.75%. [ABSTRACT FROM AUTHOR]

Laboulbeniopsis termitarius , an ectoparasitic fungus of termites, was harvested from the body surface of Reticulitermes speratus (Isoptera: Rhinotermitidae), collected in Uji, Kyoto Prefecture, Japan. This is the first record of this fungus from Japan. Three to eighteen termite workers from 500 individuals were infected among the eight colonies investigated. From our results, several hundred termite are required for the survey of this ectoparasitic fungi in Japan. The temperature effect on the relationship between the infection rate and the host activity was discussed. [ABSTRACT FROM AUTHOR]

Abstract: Spirotrichonymphea is a class of hypermastigote parabasalids defined by their spiral rows of many flagella. They are obligate hindgut symbionts of lower termites. Despite more than 100 yr of morphological and ultrastructural study, the group remains poorly characterised by molecular data and the phylogenetic positions and taxonomic validity of most genera remain in question. The genus Spirotrichonympha has been reported to inhabit several termite genera, including Reticulitermes, Coptotermes, and Hodotermopsis. The type species for this genus, Spirotrichonympha flagellata, was described from Reticulitermes lucifugus but no molecular data are yet available for this species. In this study, three new Spirotrichonympha species are described from three species of Reticulitermes. Their molecular phylogenetic position indicates that the genus is not monophyletic, as Spirotrichonympha species from Coptotermes, Paraneotermes, and Hodotermopsis branch separately. In contrast, the genus Holomastigotoides is monophyletic, as demonstrated using new sequences from Holomastigotoides species. The presence of Holomastigotoides in Prorhinotermes and the distinct phylogenetic positions of Spirotrichonympha from Reticulitermes and Coptotermes are consistent with a previously proposed symbiont fauna replacement in the ancestor of Reticulitermes. [ABSTRACT FROM AUTHOR]

Coptotermes suzhouensis (Isoptera: Rhinotermitidae) is a significant subterranean termite pest of wooden structures and is widely distributed in southeastern China. The complete mitochondrial DNA sequence of C. suzhouensis was analyzed in this study. The mitogenome was a circular molecule of 15,764 bp in length, which contained 13 protein-coding genes (PCGs), 22 transfer RNA genes, two ribosomal RNA genes, and an A+T-rich region with a gene arrangement typical of Isoptera mitogenomes. All PCGs were initiated by ATN codons and terminated by complete termination codons (TAA), except COX2, ND5, and Cytb, which ended with an incomplete termination codon T. All tRNAs displayed a typical clover-leaf structure, except for tRNASer(AGN), which did not contain the stemloop structure in the DHU arm. The A+T content (69.23%) of the A+T-rich region (949 bp) was higher than that of the entire mitogenome (65.60%), and two different sets of repeat units (A+B) were distributed in this region. Comparison of complete mitogenome sequences with those of Coptotermes formosanus indicated that the two taxa have very high genetic similarity. Forty-one representative termite species were used to construct phylogenetic trees by maximum likelihood, maximum parsimony, and Bayesian inference methods. The phylogenetic analyses also strongly supported (BPP, MLBP, and MPBP = 100%) that all C. suzhouensis and C. formosanus samples gathered into one clade with genetic distances between 0.000 and 0.002. This study provides molecular evidence for a more robust phylogenetic position of C. suzhouensis and inferrs that C. suzhouensis was the synonymy of C. formosanus. [ABSTRACT FROM AUTHOR]

In addition to being a structural pest, the Asian subterranean termite, Coptotermes gestroi (Wasmann) (Blattodea [Isoptera]: Rhinotermitidae), appears to be a serious pest of the native urban tree canopy of southeastern Florida. Asian subterranean termite colonies have the ability to cause feeding damage to a wide range of trees comprising the urban canopy, some potentially to a lethal level. Slash pine, Pinus elliottii Engelm. (Pinaceae), appears to be particularly susceptible to C. gestroi feeding damage, as the termites feed primarily on the outer layers of the wood, immediately beneath the bark, wounding live tissues and ultimately girdling the tree. In comparison, hardwood trees such as live oak, Quercus virginiana Mill. (Fagaceae), sustain damage to the heartwood at the center of the tree, resulting in a central cavity, but leaving the live tissue intact. Our results suggest that C. gestroi has the potential to kill pine trees within the local urban canopy, which may irreversibly alter the urban forest composition. In addition, many large trees in our survey have been consumed partially by mature C. gestroi colonies, potentially compromising their structural integrity. Some large oak trees that were extensively hollowed out by C. gestroi collapsed in 2017 during hurricane Irma. We discuss different potential approaches for protecting susceptible trees, as a comprehensive IPM strategy is needed for this invasive termite pest species, especially in the complex and extensive urban forest of southeastern Florida. [ABSTRACT FROM AUTHOR]

To preserve natural forests, forest plantations with fast-growing species are promoted. In the present work, the heartwood durabilities of Dyera costulata, Neolamarckia cadamba and Khaya ivorensis, from both plantations and natural forests, have been tested in comparison with rubberwood in laboratory and above-ground tests against the subterranean termite Coptotermes curvignathus. Both young trees from forest thinning and mature trees were tested. The results show that there is no significant difference in weight loss between planted trees and trees from a natural forest. The results were also compared with those of rubberwood. [ABSTRACT FROM AUTHOR]

Termites belong to the infraorder Isoptera, which contains almost 3,000 described species worldwide. These social insects cause substantial damage globally leading to billions of dollars of losses annually. Damage can occur to timber, wooden goods, paper, cotton, certain plastics, trees and many crops. Consequently, termite control and management is a major sector in the global pest-control industry. However, economic losses due to termite damage have not been quantified in the Fiji Islands to date. A review of published literature was conducted to assess the geographic range of Asian subterranean termites that occur in the Fiji Islands and to identify existing and potential control measures. The most common termite species that is known to cause millions of dollars of damage each year in the Fiji Islands is Coptotermes gestroi. This species is currently controlled primarily using the chemical fipronil but integrated termite management is the preferred long-term solution. Other possible control methods include physical, cultural, chemical and biological options. [ABSTRACT FROM AUTHOR]

BACKGROUND The lethal time of a chitin synthesis inhibitor (CSI)-baited colony of Coptotermes formosanus Shiraki largely depends on the time taken for workers to molt after consuming a lethal dose. Given that termite molting biology can regulate the time to elimination of a baited termite colony, the objective of this study was to determine how the fasting period (an important element of the worker molting cycle) and acquisition of the lethal dose can impact the total elimination period for a colony. RESULTS Noviflumuron eliminated the foraging population over a 30 m-long foraging arena within 9 weeks. However, mortality in treatment groups did not begin until day 11. The 10-day lag post treatment corresponds to the fasting period during which workers to prepare for ecdysis on the 11th day, at which time mortality occurs due to the effect of CSI. CONCLUSION This study shows that, with the exception of successful molting events during the first 10 days after baiting, mortality among workers is inevitable in the next molt with lethal doses of noviflumuron. The 10-day fasting period can comprise at least 22% of a molt cycle (45 days for a fourth worker instar or lower), and can provide a window during which a worker escapes the effect of CSI bait, thereby extending the total time needed for elimination of a colony. © 2017 Society of Chemical Industry [ABSTRACT FROM AUTHOR]

1. In incipient termite colonies, biparental brood care rapidly shifts towards alloparental brood care. This transition was suggested to recapitulate the evolutionary trajectory from subsocial wood roach ancestors to eusociality in termites. 2. Incipient colonies of the subterranean termite Coptotermes gestroi ( Wasmann) were investigated to determine if this transition was irreversible and if the burden of brood care on the first alloparents could be measured. To do so, the initial size of the work force necessary for an incipient colony to survive once the brood care became alloparental was determined. 3. The results of the study show that within 5 months after foundation, brood care duties were fully transferred to workers and the primary reproductives became irreversibly dependent on these workers for survival, reproduction, and colony growth. 4. Once the brood care became strictly alloparental, the presence of a single worker was enough to maintain the survival of the king and queen, confirming that 'reversed parental care' was also achieved. However, major brood loss and suppressed egg-laying activity from the queen was observed, suggesting that the burden of brood care was too high for a single worker to absorb. Therefore, once brood care has shifted to alloparents, a critical number of workers is necessary to prevent brood loss and initiate colony growth. 5. As the initial cost of brood care is rapidly absorbed with colony growth in termites, the performance in brood care of the first few alloparents in a subsocial wood roach ancestor may have contributed to the emergence of eusociality in this clade. [ABSTRACT FROM AUTHOR]

Termite baits using chitin synthesis inhibitors can protect structures from subterranean termite damage by eliminating colonies. One of the potential shortcomings of baits is that termites may have to feed extensively on a bait station for the colony to be eliminated. If disturbances occur during this critical feeding time, termites may abandon the site, which could prevent colony elimination. We exposed whole laboratory colonies of Coptotermes gestroi (Wasmann) containing an average of 59,000 termites to 0.5% noviflumuron baits, from 1 to 10 d, to simulate a temporary access to the bait as a result of disturbances. Our results show that colonies which fed for a single day and consumed as little as 1.1 g of bait matrix (=5.5 mg noviflumuron) were eliminated within 90 d. This study indicates that only a fraction of the bait matrix is necessary for elimination of large laboratory colonies, and that hypothetical disturbance reducing the access to a bait station would likely have little impact on the effect of the bait. Within 1 d, colonies had already acquired a lethal dose, which is shared by trophallaxis to all individuals of the colony, and colonies inevitably died despite short exposure to noviflumuron. It is estimated that if a field mature colony of 1 million individuals consumes more than 18.6 g of bait (=93 mg noviflumuron) in a relatively short period, it would already have acquired a colony-wide lethal dose and reached a point of no-return for colony elimination. [ABSTRACT FROM AUTHOR]

Competition is a fundamental process in ecology and helps to determine dominance hierarchies. Competition and dominance hierarchies have been little investigated in wood-eating termites, despite the necessary traits of similar resources, and showing spatial and temporal overlap. Competition and dominance between five species of wood-eating termites found in Huangzhou, China, was investigated in three laboratory experiments of aggression and detection, plus a year-long field survey of termite foraging activity. Dominance depended on body size, with largest species winning overwhelmingly in paired contests with equal numbers of individuals, although the advantage was reduced in paired competitions with equal biomass. The termites could detect different species from used filter papers, as larger species searched through paper used by smaller species, and smaller species avoided papers used by larger species. The largest species maintained activity all year, but in low abundance, whereas the second largest species increased activity in summer, and the smallest species increased their activity in winter. The termite species displayed a dominance hierarchy based on fighting ability, with a temporal change in foraging to avoid larger, more dominant species. The low abundance of the largest species, here Macrotermes barneyi, may be a function of human disturbance, which allows subordinate species to increase. Thus, competitive release may explain the increase in abundance of pest species, such as Coptotermes formosanus, in highly modified areas, such as urban systems.