Termite-mediated decomposition is an important, but often overlooked, component of the carbon cycle. Using a large-scale suppression experiment in Borneo, Griffiths et al. found that termites contribute between 58 and 64% of mass loss from dead wood.
The optimum maintenance conditions of the fungus-growing termite, Macrotermes gilvus (Hagen) (Blattodea: Termitidae), in the laboratory were studied. Termites were kept on a matrix of moist sand and with fungus comb as food. The survival of groups of termites was measured when maintained at different population densities by changing group size and container volume. Larger groups (≥0.6 g) were more vigorous and had significant higher survival rates than smaller groups (≤0.3 g). The population density for optimal survival of M. gilvus is 0.0025 g per container volume (ml) or 0.0169 g per matrix volume (cm(3)), i.e., 1.2 g of termites kept in a 480-ml container filled with 71 cm3 of sand. In termite groups of smaller size (i.e., 0.3 g) or groups maintained in smaller container (i.e., 100 ml) the fungus comb was overgrown with Xylaria spp., and subsequently all termites died within the study period. The insufficient number of workers for regulating the growth of unwanted fungi other than Termitomyces spp. in the fungus comb is the most likely reason. Unlike some other mound-building termite species, M. gilvus showed satisfactory survival when maintained in non-nutritious matrix (i.e., sand). There was no significant difference in the survival rate between different colonies of M. gilvus (n=5), with survival in the range of 78.5-84.4% after 4 wk. Advances in the maintenance of Macrotermes will enable researchers to study with more biological relevance many aspects of the biology, behavior, and management of this species.
Flight activities of two sympatric termite species, Macrotermes gilvus (Hagen) and Macrotermes carbonarius (Hagen), were studied in Penang Island, Malaysia. Herein, we present the first documentation of chronological reproductive isolation of M. gilvus and M. carbonarius. Flights of M. gilvus were recorded over a remarkably long 7-mo period from March to September, whereas swarming of M. carbonarius took place from November to January. Swarming events of M. gilvus and M. carbonarius occurred under atmospheric pressures of 1,005-1,011 and 1,006-1,010 hPa, respectively. Most flights of M. gilvus occurred on days with rain, whereas M. carbonarius avoided rain. Flight activity of M. gilvus was correlated significantly with atmospheric pressure and rainfall. The threshold temperature and relative humidity of M. gilvus flights were between 23 and 26 degrees C and 83 and 98% RH, respectively; M. carbonarius swarmed between 25 and 30 degrees C and 72 and 83% RH, respectively. The flight activity of M. gilvus concentrated in the warmer and humid months with a monthly total rainfall of 228 mm. Both species swarmed at distinct times of day during the limited field observations: Flights of M. gilvus began between 0300 and 0430 hours (light intensity <1 Lx), and flights of M. carbonarius lasted for only 4-10 min between 1900 and 1910 hours (at dusk; light intensity: 20-200 Lx). Windless conditions were preferred for the flights of both species.
Termites perform key ecological functions in tropical ecosystems, are strongly affected by variation in rainfall, and respond negatively to habitat disturbance. However, it is not known how the projected increase in frequency and severity of droughts in tropical rainforests will alter termite communities and the maintenance of ecosystem processes. Using a large-scale termite suppression experiment, we found that termite activity and abundance increased during drought in a Bornean forest. This increase resulted in accelerated litter decomposition, elevated soil moisture, greater soil nutrient heterogeneity, and higher seedling survival rates during the extreme El Niño drought of 2015-2016. Our work shows how an invertebrate group enhances ecosystem resistance to drought, providing evidence that the dual stressors of climate change and anthropogenic shifts in biotic communities will have various negative consequences for the maintenance of rainforest ecosystems.
Animal interactions play an important role in understanding ecological processes. The nature and intensity of these interactions can shape the impacts of organisms on their environment. Because ants and termites, with their high biomass and range of ecological functions, have considerable effects on their environment, the interaction between them is important for ecosystem processes. Although the manner in which ants and termites interact is becoming increasingly well studied, there has been no synthesis to date of the available literature. Here we review and synthesise all existing literature on ant-termite interactions. We infer that ant predation on termites is the most important, most widespread, and most studied type of interaction. Predatory ant species can regulate termite populations and subsequently slow down the decomposition of wood, litter and soil organic matter. As a consequence they also affect plant growth and distribution, nutrient cycling and nutrient availability. Although some ant species are specialised termite predators, there is probably a high level of opportunistic predation by generalist ant species, and hence their impact on ecosystem processes that termites are known to provide varies at the species level. The most fruitful future research direction will be to evaluate the impact of ant-termite predation on broader ecosystem processes. To do this it will be necessary to quantify the efficacy both of particular ant species and of ant communities as a whole in regulating termite populations in different biomes. We envisage that this work will require a combination of methods, including DNA barcoding of ant gut contents along with field observations and exclusion experiments. Such a combined approach is necessary for assessing how this interaction influences entire ecosystems.
The aim of our study was to investigate the intra- and interspecific agonistic behaviors exhibited by the worker and soldier castes of the subterranean termite Microcerotermes crassus Snyder (Isoptera: Termitidae). Aggression between M. crassus colonies from different field locations and also against three termite species--Coptotermes gestroi (Wasmann), Globitermnes sulphureus Haviland, and Odontotermes sp.--were observed in the laboratory. Termite responses were tested in paired combination of castes (soldiers versus soldiers, soldiers versus workers, and workers versus workers) consisting of 10 individuals each. Significant agonistic behaviors were observed only in encounters between pairings of different termite species. M. crassus was aggressive toward individuals from different species but not toward individuals from different M. crassus colonies. Mortality of M. crassus reached 100% in most of the interspecific encounters. However, no or low mortality was recorded in the intraspecific pairings.
Moisture is an important physical factor for the survival of termites. The effects of different moisture levels (0, 5, 10, 15, 20, and 25%) of a sand substrate on the behavior of laboratory groups of Microcerotermes crassus Snyder and Coptotermes gestroi (Wasmann) (Blattodea: Termitidae: Rhinotermitidae) were evaluated. Moisture content of sand affected wood consumption and influenced termite distribution across a moisture gradient for M. crassus. Changing the moisture parameters affected the location preference of C. gestroi, but the effect on wood consumption was not significant. Nonetheless, M. crassus and C. gestroi showed a similar distribution pattern of association with particular moisture levels.
A comparative field study was conducted to evaluate the ability of subterranean termites to damage a set of four different plastic materials (cable sheathings) exposed below- and above-ground. Eight pest species from six countries were included, viz., Coptotermes formosanus (Shiraki) in China, Japan, and the United States; Coptotermes gestroi (Wasmann) in Thailand and Malaysia; Coptotermes curvignathus (Holmgren) and Coptotermes kalshoveni (Kemner) in Malaysia; Coptotermes acinaciformis (Froggatt) with two forms of the species complex and Mastotermes darwiniensis (Froggatt) in Australia; and Reticulitermes flavipes (Kollar) in the United States. Termite species were separated into four tiers relative to decreasing ability to damage plastics. The first tier, most damaging, included C. acinaciformis, mound-building form, and M. darwiniensis, both from tropical Australia. The second tier included C. acinaciformis, tree-nesting form, from temperate Australia and C. kalshoveni from Southeast Asia. The third tier included C. curcignathus and C. gestroi from Southeast Asia and C. formosanus from China, Japan, and the United States, whereas the fourth tier included only R. flavipes, which caused no damage. A consequence of these results is that plastics considered resistant to termite damage in some locations will not be so in others because of differences in the termite fauna, for example, resistant plastics from the United States and Japan will require further testing in Southeast Asia and Australia. However, plastics considered resistant in Australia will be resistant in all other locations.
Termites and ants contribute more to animal biomass in tropical rain forests than any other single group and perform vital ecosystem functions. Although ants prey on termites, at the community level the linkage between these groups is poorly understood. Thus, assessing the distribution and specificity of ant termitophagy is of considerable interest. We describe an approach for quantifying ant-termite food webs by sequencing termite DNA (cytochrome c oxidase subunit II, COII) from ant guts and apply this to a soil-dwelling ant community from tropical rain forest in Gabon. We extracted DNA from 215 ants from 15 species. Of these, 17.2 % of individuals had termite DNA in their guts, with BLAST analysis confirming the identity of 34.1 % of these termites to family level or better. Although ant species varied in detection of termite DNA, ranging from 63 % (5/7; Camponotus sp. 1) to 0 % (0/7; Ponera sp. 1), there was no evidence (with small sample sizes) for heterogeneity in termite consumption across ant taxa, and no evidence for species-specific ant-termite predation. In all three ant species with identifiable termite DNA in multiple individuals, multiple termite species were represented. Furthermore, the two termite species that were detected on multiple occasions in ant guts were in both cases found in multiple ant species, suggesting that ant-termite food webs are not strongly compartmentalised. However, two ant species were found to consume only Anoplotermes-group termites, indicating possible predatory specialisation at a higher taxonomic level. Using a laboratory feeding test, we were able to detect termite COII sequences in ant guts up to 2 h after feeding, indicating that our method only detects recent feeding events. Our data provide tentative support for the hypothesis that unspecialised termite predation by ants is widespread and highlight the use of molecular approaches for future studies of ant-termite food webs.
The majority of true parasitoids manipulate their host's physiology for their own benefit. In this study, we documented the physiological changes that occurred in major soldiers of the subterranean termite Macrotermes gilvus (Hagen) (Isoptera: Termitidae) parasitized by the koinobiont larval endoparasitoid Misotermes mindeni Disney and Neoh (Diptera: Phoridae). We compared the metabolic rate, body water content, body water loss rate, cuticular permeability, and desiccation tolerance between parasitized and unparasitized major soldiers. The metabolic rate of parasitized hosts was significantly higher than that of unparasitized termites. Mean total body water content of parasitized major soldiers (64.73±3.26%) was significantly lower than that of unparasitized termites (71.99±2.23%). Parasitized hosts also had significantly lower total body water loss rates (5.72±0.06%/h) and higher cuticular permeability (49.37±11.26 μg/cm/h/mmHg) than unparasitized major soldiers (6.75±0.16%/h and 60.76±24.98 μg/cm/h/mmHg, respectively). Parasitized major soldiers survived almost twice as long as unparasitized termites (LT(50)=6.66 h and LT(50)=3.40 h, respectively) and they had significantly higher tolerance to water loss compared to unparasitized termites (45.28±6.79% and 32.84±7.69%, respectively). Body lipid content in parasitized hosts (19.84±6.27%) was significantly higher than that of unparasitized termites (6.17±7.87%). Finally, parasitized hosts had a significantly lower percentage of cuticular water content than unparasitized major soldiers (10.97±1.84% and 13.17±2.21%, respectively). Based on these data, we conclude that the parasitism-induced physiological changes in the host are beneficial to the parasitoids as the alterations can clearly increase the parasite's chances of survival when exposed to extreme environmental conditions and ensure that the parasitoids are able to complete their larval development successfully before the host dies.
The foraging patterns of termites are strongly related to physiological limits in overcoming desiccation stress. In this study, we examined moisture preferences and physiological characteristics of Macrotermes carbonarius (Hagen) and M. gilvus (Hagen) as both exhibit conspicuous patterns of foraging activity. Despite both species showing no significant differences in calculated cuticular permeability, and percentage of total body water, they differed greatly in rate of water loss and surface area to volume ratio. For example, M. carbonarius which had a lower surface area to volume ratio (29.26-53.66) showed lower rate of water loss and percentage of total body water loss. This also resulted in higher LT(50) when exposed to extreme conditions (≈2% RH). However, contrasting observations were made in M. gilvus that has smaller size with higher surface area to volume ratio of 40.28-69.75. It is likely that the standard equation for calculating insect surface areas is inadequate for these termite species. The trend was further supported by the result of a moisture preference bioassay that indicated M. carbonarius had a broader range of moisture preference (between 5% and 20%) than M. gilvus which had a relatively narrow moisture preference (only 20%). These results explain why M. carbonarius can tolerate desiccation stress for a longer period foraging above-ground in the open air; while M. gilvus only forages below ground or concealed within foraging mud tubes.
Termites are the major decomposers in tropical region but yet their occurrences in oil palm plantation especially in peat soil are generally treated as pest. Study of termite species in peat land was conducted in selected oil palm plantations in North Sarawak with 5-7 years old palms and South Sarawak with 13-15 years old palms with two sites in each area. Results of quadrate (25 x 25 x 30 cm) sampling showed termite was significantly higher in relative density with increasing depth of soil (0-10 = 21.23, 10-20 = 42.52 and 20-30 cm = 81.12%) which could be advantaged from being predated by ants (Hymenoptera: Formicidae) which were higher in density from soil surface to 10 cm soil depth with relative density of 31.84%. Modified transect sampling (50x6 m) had successfully sampled 18 species of termites from 2 families (Rhinotermitidae and Termitidae), 5 subfamilies (Rhinotermitinae, Coptotermitinae, Termitinae, Macrotermitinae and Nasutitermitinae) and 11 genera (Coptotermes, Schedorhinotermes, Termes, Macrotermes, Nasutitermes, Globitermes, Amitermes, Parrhinotermes, Pericapritermes, Havilanditermes and Prohamitermes). Both plantation sites have termite dominantly feeding on rotten wood as a result of abundant dead woods. However, Coptotermes curvignathus Holmgren was identified to feed on the living tissues of oil palm causing damage or death of the tree. Study showed higher encounter of soil-feeding termite in longer established plantation. It indicates the gradually shifting of soil condition towards a stabilized environment which favors the successful settlement of soil feeder termite species. Termite control should be more targets specific to avoid harming beneficial termites.
Coptotermes gestroi (Wasmann) or the Asian subterranean termite is a serious structural pest in urban settlements in Southeast Asia that has been introduced to other parts of the world through human commerce. Although mitochondrial DNA markers were previously used to shed light on the dispersal history of the Asian subterranean termite, there were limited attempts to analyze or include populations of the termite found in the wild in Southeast Asia. In this study, we analyzed the 16S ribosomal RNA (16S rRNA) and cytochrome c oxidase subunit 1 (cox1) genes of Asian subterranean termite colonies found in mangrove swamps, beach forests, plantations, and buildings in semi-urban and urban areas to determine the relationship between colonies found in the wild and the urban habitat, and to investigate the possibility of different ecotypes of the termite in Peninsular Malaysia. Our findings show that the 16S rRNA haplotypes recovered from this study clustered into eastern, western, and southern populations of the termite, while the cox1 haplotypes were often specific to an area or site. The 16S rRNA and cox1 genes or haplotypes showed that the most abundant haplotype occupied a wide range of environments or habitats. In addition, the cox1 tree showed evidence of historical biogeography where basal haplotypes inhabited a wide range of habitats, while apical haplotypes were restricted to mangrove swamps and beach forests. Information on the haplotype-habitat association of C. gestroi will enable the prediction of habitats that may harbor or be at risk of invasion in areas where they have been introduced.
Feeding responses of subterranean termites Coptotermes curvignathus (Holmgren) and Coptotermes gestroi (Wasmann) (Blattodea: Rhinotermitidae) to bait matrices supplemented with various sugars, amino acids, and cassava were evaluated both in the laboratory and field. The results indicated that the two termite species consumed significantly different amount of filter papers that had been treated with various types and concentrations of sugars and amino acids. Based on consumption and survival data, filter papers with 3% glucose and 3% xylose were among the most consumed by C. curvignathus and C. gestroi, respectively. Both termite species consumed more of the filter papers treated with 3% casein than filter papers treated with L-alanine. Both species had a comparable survival rate compared with those in the controls. Results from laboratory and field trials on bait prototypes indicated that C. gestroi consumed more bait prototypes containing cellulose, 3% xylose, 3% casein, and cassava, whereas C curvignathus consumed more bait prototype containing cellulose, 3% glucose, and cassava, than on pure crystalline cellulose baits. Thus, with an improved and cost-effective bait formulation, a much wider control of subterranean termite colonies could be achieved.
Dual choice bioassays were used to evaluate the antifeedant property of essential oil and methanolic extract of Alpinia galanga (L.) (locally known as lengkuas) against two species of termites, Coptotermes gestroi (Wasmann) and Coptotermes curvignathus (Holmgren) (Isoptera: Rhinotermitidae). A 4-cm-diameter paper disc treated with A. galanga essential oil and another treated with either methanol or hexane as control were placed in a petri dish with 10 termites. Mean consumption of paper discs (miligram) treated with 2,000 ppm of essential oil by C. gestroi was 3.30 ± 0.24 mg and by C. curvignathus was 3.32 ± 0.24 mg. A. galanga essential oil showed significant difference in antifeedant effect, 2,000 ppm of A. galanga essential oil was considered to be the optimum concentration that gave maximum antifeedant effect. The essential oil composition was determined using gas chromatography-mass spectrometry. The major component of the essential oil was 1,8-cineol (61.9%). Antifeedant bioassay using 500 ppm of 1,8-cineol showed significant reduction in paper consumption by both termite species. Thus, the bioactive agent in A. galangal essential oil causing antifeeding activity was identified as 1,8-cineol. Repellent activity shows that 250 ppm of 1,8-cineol caused 50.00 ± 4.47% repellency for C. gestroi, whereas for C. curvignathus 750 ppm of 1,8-cineol was needed to cause similar repellent activity (56.67 ± 3.33%). C. curvignathus is more susceptible compare to C. gestroi in Contact Toxicity study, the lethal dose (LD50) of C. curvignathus was 945 mg/kg, whereas LD50 value for C. gestroi was 1,102 mg/kg. Hence 1,8-cineol may be developed as an alternative control against termite in sustainable agriculture practices.
Coptotermes curvignathus Holmgren is capable of feeding on living trees. This ability is attributed to their effective digestive system that is furnished by the termite's own cellulolytic enzymes and cooperative enzymes produced by their gut microbes. In this study, the identity of an array of diverse microbes residing in the gut of C. curvignathus was revealed by sequencing the near-full-length 16S rRNA genes. A total of 154 bacterial phylotypes were found. The Bacteroidetes was the most abundant phylum and accounted for about 65% of the gut microbial profile. This is followed by Firmicutes, Actinobacteria, Spirochetes, Proteobacteria, TM7, Deferribacteres, Planctomycetes, Verrucomicrobia, and Termite Group 1. Based on the phylogenetic study, this symbiosis can be a result of long coevolution of gut enterotypes with the phylogenic distribution, strong selection pressure in the gut, and other speculative pressures that determine bacterial biome to follow. The phylogenetic distribution of cloned rRNA genes in the bacterial domain that was considerably different from other termite reflects the strong selection pressures in the gut where a proportional composition of gut microbiome of C. curvignathus has established. The selection pressures could be linked to the unique diet preference of C. curvignathus that profoundly feeds on living trees. The delicate gut microbiome composition may provide available nutrients to the host as well as potential protection against opportunistic pathogen.