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Stable isotope ratios and uric acid preservation in termites belonging to three feeding habits in Thailand.

Nitrogen and carbon stable isotope ratios and uric acid concentrations in termites sampled from a dry evergreen forest in Thailand, were determined across three kinds of feeding habits. Feeding habits of Microcerotermes crassus, which is an abundant wood-feeder, and Dicuspiditermes makhamensis, a common soil-feeding termite, were confirmed by isotopic signatures. Lichen feeding termites (Hospitalitermes birmanicus, H. bicolor and H. ataramensis) were characterized by low delta15N values, suggesting that they assimilated nitrogen deposited from the atmosphere. There was also a significant difference in uric acid concentrations between termites representing different feeding habits. No significant relationships were found between uric acid concentrations and delta15N or delta13C in Hospitalitermes. However, delta15N values were correlated with C/N ratios in H. birmanicus, except in one colony of H. ataramensis. delta13C values in both species were negatively correlated with C/N ratios.

Animals↗

Do stable isotopes reflect the food web development in regenerating ecosystems?

We evaluated the use of delta15N- and delta13C-values to monitor the development of food web complexity and biodiversity in a regenerating ecosystem. Therefore a model food chain was established feeding cultivated woodlice (Porcellio dilatatus) on a cellulolytic fungus (Chaetomium globosum) grown on cellulose paper. Two diets of different quality (C:N ratios of 54 vs. 200) with different delta15N- (1.3% vs. 3.1%) but identical delta13C-values caused low and high dietary stress in animals of treatment A and B, respectively. After an incubation time of 7 weeks amount, elemental and isotopic composition of collected faeces and exuviae as well as woodlice and remaining food were determined. The increase of delta15N-values of woodlice relative to the diet was 5.7% and 2.5% in treatments A and B, respectively, whereas delta13C-shifts were 1.0% and 1.6%, showing a reverse relationship. Modelling of elemental and isotopic mass balances indicated that faeces recycling explains the unexpected high 15N-enrichments. Moreover, 13C-enrichments were positively correlated to the degree of starvation. Considering the effects of starvation and recycling of faeces, stable isotopes represent a useful tool to elucidate trophic interactions in regenerating food webs.

Animals↗

Beta-orcinol depsidones from the lichen Usnea sp. from Sri Lanka.

Two beta-orcinol depsidone lactones, the methyl ethers of menegazziaic acid and stictic acid were isolated along with glyceryl trilinolate and usnic acid from an Usnea sp. new to Sri Lanka growing on rotting trees of Acacia decurrans. Usnic acid exhibited potent antitermite activity against a common pest of tea, Glyptotermes dilatatus, at low elevations.

Animals↗

On the mound of Macrotermes michaelseni as an organ of respiratory gas exchange.

Patterns and rates of air movements in the mounds and nests of Macrotermes michaelseni were studied using tracer methods. Wind is a significant source of energy for powering nest ventilation, despite the mound being a completely enclosed structure. Nests are ventilated by a tidal movement of air driven by temporal variation in wind speed and wind direction. Density gradients sufficiently steep to drive bulk flow by natural convection will be rare. However, metabolism-induced buoyant forces may interact with wind energy in a way that promotes homeostasis of the mound atmosphere.

Air Movements↗

Termite digestibility and water and energy contents determine the water economy index of numbats (Myrmecobius fasciatus) and other myrmecophages.

Digestibility by captive numbats for termites was determined by feeding trials to be 81%+/-1.2% for Coptotermes sp. and 64%+/-3.3% for Nasutitermes sp. Water, ash, and energy content of both the Coptotermes (0.96+/-0.099 mg(dry mass) individual(-1), 78.0%+/-0.36% water, 5.8%+/-0.31% ash, 23.1+/-0.19 kJ g-1dry total energy) and Nasutitermes (0.91+/-0.046 mg(dry mass) individual(-1), 76.7%+/-3.09% water, 7.5%+/-1.10% ash, 22.7+/-0.36 kJ g-1dry total energy) were similar to values measured previously for other termites and for ants and insects in general. Numbats have a slow passage time for termites (20-30 h), presumably to enhance the digestion of termites. The water economy index (WEI) was 0.2 for captive numbats feeding on Coptotermes and 0.25 for Nasutitermes, whereas the WEI measured for wild, free-living numbats was 0.29, which corresponds to a digestibility of 58%. The WEI of a myrmecophage diet is determined by the energy and water contents and digestibilities of termites and ants, in the absence of drinking. The WEI for numbats, and other termitivorous mammals as well as reptiles, is higher than would be expected for an animal-based diet because of their relatively low digestibility (58%-81%) for termites. A high WEI preadapts myrmecophages to survival in arid environments without having to drink.

Analysis of Variance↗

New insights into chimpanzees, tools, and termites from the Congo Basin.

The tool-using behaviors of wild chimpanzees comprise the most impressive assemblages and flexible repertoires of nonhuman material culture. We expand knowledge of the breadth and complexity of tool use in this species by providing the first descriptions of the form and function of two distinct tool sets used by chimpanzees in preying upon termites within the forests of the Goualougo Triangle, Republic of Congo. Further, we report the first application of remote video monitoring technology to record wild chimpanzee tool-using behavior. Based on tool assemblages recovered at termite nests, we hypothesized that chimpanzees were regularly visiting two forms of termite nests and using specific tools to extract termite prey depending on the structure of the nest. Six months of continuous remote video monitoring at six termite nests confirmed that chimpanzees use a tool set to puncture and fish at subterranean termite nests and another tool set to perforate and fish at epigeal (aboveground) nests. Our findings of strict adherence to tool forms at different nest types, tool material selectivity, repeated visits to nests with reusable wood tool assemblages, and differences in material culture between communities have broad implications for our understanding of the ecological and cultural factors that shape hominoid tool use.

Animals↗

Ability of canine termite detectors to locate live termites and discriminate them from non-termite material.

Dogs were trained to detect Eastern subterranean termites, Reticulitermes flavipes (Kollar), using the United States Customs method of scent detection dog training modified with a food reward. Dogs were tested with various numbers of Eastern subterranean termites placed in vented PVC containers. Trained dogs were 95.93% accurate in finding > or = 40 Eastern subterranean termite workers (positive indications) and incorrectly indicated the presence of termites in 2.69% of the containers without termites. Multiple species of termites [dark southern subterranean, R. virginicus (Banks); Formosan subterranean, Coptotermes formosanus Shiraki; powderpost, Cryptotermes cavifrons Banks; and southeastern drywood termites, Incisitermes snyderi (Light)], were similarly evaluated. Dogs trained to locate Eastern subterranean termites were also 100% accurate in finding dark southern subterranean termites, 98.89% accurate in finding Formosan subterranean termites, 97.33% accurate in finding powderpost termites, and 88.89% accurate in finding southeastern drywood termites. Dogs were able to discriminate live termites from non-termite material. Trained dogs' false response rate was 25.33% to Eastern subterranean termite-damaged wood, 6.67% to American cockroaches, Periplaneta americana (L.), and 2.67% to Florida carpenter ants, Camponotus floridanus Buckley.

Animals↗

Screening method for inhibitors against formosan subterranean termite beta-glucosidases in vivo.

Cellulose, a main structural constituent of plants, is the major nutritional component for wood-feeding termites. Enzymatic hydrolysis of cellulose to glucose occurs by the action of cellulases, a mixture of the three major classes of enzymes including endo-1,4-beta-glucanases, exo-1,4-beta-glucanases, and beta-glucosidase. Lower termites, such as the Formosan subterranean termite, Coptotermes formosanus Shiraki, require cellulolytic protozoa to efficiently digest cellulose for survival. Inhibitors developed against any of these cellulase system enzymes would be a potential termite treatment avenue. Our effort was to develop a screening system to determine whether termites could be controlled by administration of cellulase system inhibitors. Some reported compounds such as gluconolactone, conduritol B epoxide, and 1-deoxynojirimycin are potential beta-glucosidase inhibitors, but they have only been tested in vitro. We describe an in vivo method to test the inhibitory ability of the designated chemicals to act on beta-1,4-glucosidases, one member of the cellulase system that is the key step that releases glucose for use as an energy and carbon source for termites. Inhibition in releasing glucose from cellooligosaccharides might be sufficient to starve termites. Fluorescein di-beta-D-glucopyranoside was used as the artificial enzyme substrate and the fluorescent intensity of the reaction product (fluorescein) quantified with an automated fluorescence plate reader. Several known in vitro beta-1,4-glucosidase inhibitors were tested in vivo, and their inhibitory potential was determined. Endogenous and protozoan cellulase activities are both assumed to play a role.

Animals↗

Colony social organization and population genetic structure of an introduced population of formosan subterranean termite from New Orleans, Louisiana.

The Formosan subterranean termite, Coptotermes formosanus Shiraki, is an invasive species in many parts of the world, including the U.S. mainland. The reasons for its invasive success may have to do with the flexible social and spatial organization of colonies. We investigated the population and breeding structure of 14 C. formosanus colonies in Louis Armstrong Park, New Orleans, LA. This population has been the focus of extensive study for many years, providing the opportunity to relate aspects of colony breeding structure to previous findings on colony characteristics such as body weight and number of workers, wood consumption, and intercolony aggression. Eight colonies were headed by a single pair of outbred reproductives (simple families), whereas six colonies were headed by low numbers of multiple kings and/or queens that were likely the neotenic descendants of the original colony (extended families). Within the foraging area of one large extended family colony, we found genetic differentiation among different collection sites, suggesting the presence of separate reproductive centers. No significant difference between simple family colonies and extended family colonies was found in worker body weight, soldier body weight, foraging area, population size, or wood consumption. However, level of inbreeding within colonies was negatively correlated with worker body weight and positively correlated with wood consumption. Also, genetic distance between colonies was positively correlated with aggression levels, suggesting a genetic basis to nestmate discrimination cues in this termite population. No obvious trait associated with colony reproductive structure was found that could account for the invasion success of this species.

Animals↗

Nesting ecology and cuticular microbial loads in dampwood (Zootermopsis angusticollis) and drywood termites (Incisitermes minor, I. schwarzi, Cryptotermes cavifrons).

Termites form one-piece nests in wood that can vary in their moisture content and degree of decomposition, and thus microbial richness. To estimate the microbial load of nests and the potential risk they pose for colony members, we quantified the number of microbes in the nest and on the cuticle of the dampwood termite, Zootermopsis angusticollis, and three drywood termites, Incisitermes minor, I. schwarzi, and Cryptotermes cavifrons. The number of colony forming units (CFUs) cultured from nest material samples and washes of the cuticle of larvae and nymphs were determined. CFUs recorded from nest material was low (fewer than 60 CFUs/g) in the drywood termites and comparatively high in the dampwood species, as more than 800 bacterial and fungal CFUs/g were cultured from the nest material of Z. angusticollis. Similarly, cuticular microbial loads were negligible in the drywood termites sampled, ranging from 0.5 to fewer than 16 CFUs/cm2, whereas approximately 200 CFUs/cm2 were cultured from Z. angusticollis. The nesting and feeding habits of these basal species likely influence colony microbial load and the degree of pathogen exposure, which in turn could favor adaptations to resist disease that vary with termite nesting biology.

Animals↗

Neotenic formation in laboratory colonies of the termite Coptotermes gestroi after orphaning.

The termite Coptotermes gestroi (Wasmann 1896) (Rhinotermitidae: Coptotermitinae) is an exotic species in Brazil and information concerning its reproductive developmental biology is scarce. We induced the formation of neotenics in laboratory colonies through orphaning experiments. Orphaning experiments were conducted in three-year old colonies of C. gestroi kept under laboratory conditions. After three months, eight nymphoid neotenics were observed in one colony after queen removal. Histological analysis showed that these neotenics were non-functional. The results suggest that these individuals may have arisen from the first nymphal instar (N1) or from an early N1 instar after one or two larval moults. Neotenics also were recorded on two incipient colonies of C. gestroi that lost the queen naturally.

Animals↗

Duplication and diversifying selection among termite antifungal peptides.

We have identified and analyzed the mRNA sequence of 20 new defensin-like peptides from 11 Australian termite species of Nasutitermes and from an outgroup, Drepanotermes rubriceps. The sequence was amplified by reverse transcriptase PCR with a degenerate primer designed from termicin, an antifungal peptide previously characterized from the termite Pseudocanthotermes spiniger. All 20 genes show high sequence identity with P. spiniger termicin and have duplicated repeatedly during the radiation of Nasutitermes. Comparison of the relative fixation rates of synonymous (silent) and nonsynonymous (amino acid altering) mutations indicates that the Nasutitermes termicins are positively selected. This positive selection appears to drive a decrease in termicin charge. In termites with two genes, the decrease in charge is predominantly restricted to one termicin. Furthermore, the spread of charge is significantly greater within species than across species among amino acid sites that appear to be under strong positive selection and this spread is attributable to only three sites. Our results suggest that after termicin duplication, certain critical sites have maintained a positive charge in one duplicate and evolved towards neutrality in the other and that positive selection has directed these changes repeatedly and independently. This diversification among duplicated genes may be a counter-response to the evolution of fungal resistance in social insects that are particularly vulnerable to fungal epidemics.

Amino Acid Sequence↗

Sociality and the rate of molecular evolution.

The molecular clock does not tick at a uniform rate in all taxa but may be influenced by species characteristics. Eusocial species (those with reproductive division of labor) have been predicted to have faster rates of molecular evolution than their nonsocial relatives because of greatly reduced effective population size; if most individuals in a population are nonreproductive and only one or few queens produce all the offspring, then eusocial animals could have much lower effective population sizes than their solitary relatives, which should increase the rate of substitution of "nearly neutral" mutations. An earlier study reported faster rates in eusocial honeybees and vespid wasps but failed to correct for phylogenetic nonindependence or to distinguish between potential causes of rate variation. Because sociality has evolved independently in many different lineages, it is possible to conduct a more wide-ranging study to test the generality of the relationship. We have conducted a comparative analysis of 25 phylogenetically independent pairs of social lineages and their nonsocial relatives, including bees, wasps, ants, termites, shrimps, and mole rats, using a range of available DNA sequences (mitochondrial and nuclear DNA coding for proteins and RNAs, and nontranslated sequences). By including a wide range of social taxa, we were able to test whether there is a general influence of sociality on rates of molecular evolution and to test specific predictions of the hypothesis: (1) that social species have faster rates because they have reduced effective population sizes; (2) that mitochondrial genes would show a greater effect of sociality than nuclear genes; and (3) that rates of molecular evolution should be correlated with the degree of sociality. We find no consistent pattern in rates of molecular evolution between social and nonsocial lineages and no evidence that mitochondrial genes show faster rates in social taxa. However, we show that the most highly eusocial Hymenoptera do have faster rates than their nonsocial relatives. We also find that social parasites (that utilize the workers from related species to produce their own offspring) have faster rates than their social relatives, which is consistent with an effect of lower effective population size on rate of molecular evolution. Our results illustrate the importance of allowing for phylogenetic nonindependence when conducting investigations of determinants of variation in rate of molecular evolution.

Animals↗

Variation in positive selection in termite GNBPs and Relish.

Social insects are model organisms for investigating molecular evolution in the innate immune system. Their diversity affords comparative analysis among closely related species, and group living is likely to contribute to the pathogen stress imposed on the immune system. We used different models of nucleotide substitution at nonsynonymous (amino acid altering) and synonymous (silent) sites to compare the different levels and type of selection among three immunity genes in 13 Australian termite species (Nasutitermes). The immunity genes include two encoding pathogen recognition proteins (gram-negative bacterial-binding proteins) that duplicated and diverged before or soon after the evolution of the termites and a transcription factor (Relish), which induces the production of antimicrobial peptides. A comparison of evolutionary models that assign four unrestricted classes of dN/dS (the ratio of the nonsynonymous to synonymous substitution rate) to different Nasutitermes lineages revealed that the occurrence of positive selection (dN/dS > 1) varies among lineages and the three genes. Positive selection appears to have driven the evolution of all three genes in an ancestral lineage of three subterranean termites. It had previously been suggested that there was a transition along this ancestral lineage to termite morphology and ecology associated with a diet of decayed wood, a diet that may expose termites to elevated levels of fungal and bacterial pathogens. Relish appears to have experienced the highest levels of selective pressure for change among all three genes. Positively selected sites in the molecule are located in regions that are important for its activation, which suggests that amino acid substitutions at these sites are a counter response to pathogen mechanisms that disrupt the activation of Relish.

Amino Acid Substitution↗

As you reap, so shall you sow: coupling of harvesting and inoculating stabilizes the mutualism between termites and fungi.

At present there is no consensus theory explaining the evolutionary stability of mutualistic interactions. However, the question is whether there are general 'rules', or whether each particular mutualism needs a unique explanation. Here, I address the ultimate evolutionary stability of the 'agricultural' mutualism between fungus-growing termites and Termitomyces fungi, and provide a proximate mechanism for how stability is achieved. The key to the proposed mechanism is the within-nest propagation mode of fungal symbionts by termites. The termites suppress horizontal fungal transmission by consuming modified unripe mushrooms (nodules) for food. However, these nodules provide asexual gut-resistant spores that form the inoculum of new substrate. This within-nest propagation has two important consequences: (i) the mutualistic fungi undergo severe, recurrent bottlenecks, so that the fungus is likely to be in monoculture and (ii) the termites 'artificially' select for high nodule production, because their fungal food source also provides the inoculum for the next harvest. I also provide a brief comparison of the termite-fungus mutualism with the analogous agricultural mutualism between attine ants and fungi. This comparison shows that--although common factors for the ultimate evolutionary stability of mutualisms can be identified--the proximate mechanisms can be fundamentally different between different mutualisms.

Agaricales↗

Limited food induces nepotism in drywood termites.

The evolution of cooperation and altruistic behaviour where individuals forego their own reproduction to help others reproduce can be explained by kin selection. Depending on the costs and benefits provided, altruism can be evolutionarily favoured if it is directed at close relatives. A considerable body of data supports the role of relatedness as a key determinant of cooperation and conflict within societies. However, the role of ecological factors and, in particular, how these costs and benefits interact with relatedness remains poorly understood. By studying 16 colonies, here I show that in a drywood termite ecological factors determine the importance of relatedness. In colonies with limited food supply, nestmates restrict cooperative interactions mainly to close relatives, while non-discriminative cooperation occurs when food is abundant. This shows for the first time directly the interaction between ecological conditions and relatedness in shaping cooperation.

Animals↗

Soldier morphogenesis in a nasute termite: discovery of a disc-like structure forming a soldier nasus.

Nasute termites belonging to the subfamily Nasutitermitinae, have a soldier caste that possesses a frontal projection (nasus) on the head, from which defensive substances are secreted. In the course of caste differentiation of the processional nasute termite Hospitalitermes medioflavus, the most dynamic morphogenesis occurs in the stage of moulting from male minor worker to presoldier (the stage preceding the soldier stage). We examined the presumptive nasus epithelium in minor workers and determined that the nasus develops rapidly just prior to the moulting to presoldiers. The rapid growth is associated with two folding layers of cuticle and epithelium, which we termed the soldier-nasus disc, and resembles the imaginal discs found in holometabolous insects.

Animals↗

The importance of leaf- and litter-feeding invertebrates as sources of animal protein for the Amazonian Amerindians.

At least 32 Amerindian groups in the Amazon basin use terrestrial invertebrates as food. Leaf- and litter-consuming invertebrates provide the more important, underestimated food sources for many Amerindian groups. Further, litter-consuming earthworms are also an important food resource for the Ye'Kuana (also known as Makiritare) in the Alto Orinoco (Amazonas, Venezuela). By selecting these small invertebrates the Amerindians are choosing their animal food from those food webs in the rainforest which have the highest energy flow and which constitute the greatest renewable stock of readily available nutrients. Here we show that the consumption of leaf- and litter-feeding invertebrates as a means of recovering protein, fat and vitamins by the forest-living peoples offers a new perspective for the development of sustainable animal food production within the paradigm of biodiversity maintenance.

Animals↗