Alarm Reaction of European Reticulitermes Termites to Soldier Head Capsule Volatiles (Isoptera, Rhinotermitidae)

The alarm behavior of worker and soldier castes of four European subterranean termite species of the genus Reticulitermes (R. santonensis, R. lucifugus, R. grassei, R. banyulensis) is described. In petri dish bioassays we presented filter papers with squashed soldier heads as an odor source to trigger a alarm reaction. Untreated filter papers were used as controls. The behavioral patterns were alike for all four species. In the control situation (no alarm), only occasional workers and few soldiers were seen walking in the petri dish. When the odor source was presented, a general state of alertness was observed, showing as higher activity and presence, zigzag running, antennation of nestmates, jittering and jerking, mandible snapping, head-banging, and attraction to the odor source. Workers were attracted within a few seconds (6.9–13.4 s); soldiers followed much later (18.5–64.6 s). Both workers and soldiers inspected the odor source only briefly (workers for 8.7–12.8 s, soldiers for 10.0–19.5 s), with the exception of R. santonensis, where soldiers stayed an average of 209.8 s. The number of individuals, of 50 workers and 3 soldiers, near the odor source averaged 6.0–9.2 workers and 1.2–2.3 soldiers. The presence of workers is crucial: without workers, a significant alarm reaction in soldiers could not be induced. The roles of soldier and worker castes of Reticulitermes species within alarm communication and the origin and nature of possible alarm signals are discussed.     

Worker aggression of ant Lasius japonicus enhanced by termite soldier–specific secretion as an alarm pheromone of Reticulitermes speratus

The subterranean termite Reticulitermes speratus usually nests in rotten wood trunks, which may also be occupied by the Japanese garden ant Lasius japonicus. Few battles were observed between them under ordinary circumstances because they inhabit separate nesting sites. However, once the termite nesting sites were artificially broken, the ant workers invaded and hunted the termites, although the termite soldiers fought against the ants. This study aims to confirm intra‐ and inter‐specific chemical interactions between the termite and ant. Solid phase microextraction–gas chromatograph (SPME‐GC) analyses revealed that R. speratus soldiers secreted caste‐specific sesquiterpene hydrocarbon when they were irritated. Both the hexane extract of the soldiers and its hydrocarbon fraction, as well as the crushed soldier bodies, attracted the soldiers but dispersed the workers when presented on the trails. We also confirmed that the soldier chemicals enhanced aggressiveness of L. japonicus, which rushed around the odor sources and hunted any termites that were present. These findings suggest that: (i) the soldier–specific secretion might serve as an alarm pheromone in termite chemical communication, in which components recruit soldiers and also warn the other colony members away; and (ii) termite communication is eavesdropped on by L. japonicus workers to locate and hunt the termites.     

Caste development systems of the Neotropical termite <Emphasis Type="Italic">Constrictotermes cyphergaster</Emphasis> (Isoptera,Termitidae)

The caste developmental system of the Neotropical termite Constrictotermes cyphergaster was investigated in 12 natural colonies from the semi-arid Caatinga region of northeastern Brazil. Twelve measurements were taken of different body parts of the larvae, workers, presoldiers, soldiers, nymphs, and alates for morphometric analyses, and principal component analyses (PCA) were performed based on these parameters. The nymphal line comprised one larval instar, followed by five nymphal instars, and alates, in agreement with the developmental pattern described for Termitidae species. The apterous line included two larval instars, followed by workers, presoldiers, and soldiers. Both workers and soldiers were found to be monomorphic and composed only of male specimens, with only one instar phase each. C. cyphergaster, like most of the Nasutitermitinae, has only male soldiers. The presence of male workers, with only one instar, suggests a simplified system of differentiation in the apterous line of C. cyphergaster. These patterns, which had not previously been described for ‘full nasute’ Nasutitermitinae, are similar to the differentiation patterns of the ‘mandibulate nasute’ Cornitermes walkeri.     

Tests of four hypotheses on soldier production,by using wild colonies ofPheidole fervida F. Smith (Hymenoptera: Formicidae)

Following four hypotheses on the production of soldiers were tested in the Pheidole fervida colonies collected in the natural field:

1. The proportion of new soldiers is primarily dependent on the population size of old workers.
2. The abundance of old soldiers reduces the proportion of new soldiers.
3. The production of sexuals reduces the proportion of new soldiers.
4. The number of soldiers is correlated with the size of defense zone.

The present test suggested that the last idea was most likely to the wild colonies of P. fervida.     

The effect of nonnestmates on foraging trail caste ratios ofNasutitermes corniger (Isoptera: Termitidae)

The mean number of workers on foraging trails ofNasutitermes corniger Motschulsky decreases after the introduction of nonnestmate conspecifics or congeneric heterospecifics. The mean number of soldiers on foraging trais does not vary when nestmates, conspecific nonnestmates or heterospecific workers are introduced. When heterospecific soldiers are introduced to the trail, however, the number ofN. corniger soldiers increases. Consequently, the ratio of soldiers to workers increases. These findings demonstrate that the addition of nonnestmates temporarily alters caste ratios on foraging trails, representing a shift in the distribution of castes in response to environmental perturbation.     

Effect of lectins from Opuntia ficus indica cladodes and Moringa oleifera seeds on survival of Nasutitermes corniger

Biodegradation by termites is a serious problem for wood and crop industries worldwide, and new environmentally friendly alternatives for termite control have been developed. This work investigated the effects of crude and purified preparations containing lectins from Opuntia ficus indica cladodes (OfiL) and Moringa oleifera seeds (WSMoL and cMoL) on Nasutitermes corniger workers and soldiers. Purified OfiL was more active than cladode extracts, showing a stronger termiticidal activity against workers (LC₅₀ of 0.116 mg ml⁻¹) than against soldiers. OfiL was active against soldiers only at 1.5 mg ml⁻¹. All preparations containing WSMoL and cMoL were active only at concentrations of 1.0 and 1.5 mg ml⁻¹. The tested preparations did not exert repellent activity against N. corniger. OfiL was able to kill workers and therefore is potentially a new tool for N. corniger control; as a consequence, this lectin could disturb organization, structure, and maintenance of termite colonies.     

Community structure and antifungal activity of actinobacteria in a fungus-growing termite

1. Fungus-growing termites cultivate the fungal mutualist Termitomyces as their main food source; however, how fungus-growing termites protect Termitomyces from threats is still unclear. In this study, we investigated the actinobacterial communities in Odontotermes formosanus individuals and their fungal combs.
2. Moreover, the antifungal activities of the isolated actinobacteria were tested. 16 S rRNA gene sequencing results indicated that the actinobacteria in O. formosanus and its fungal combs belong to 5 classes, 17 orders, 40 families, and 84 genera.
3. The relative abundance of Coriobacteriia in the nymphs, young workers, old workers, and soldiers was higher than that in the queens and fungal combs, and the relative abundance of class Actinobacteria in the queens and the fungal combs was higher than that in the nymphs, young workers, old workers, and soldiers.
4. Based on antifungal bioassays, 3 strains of Amycolatopsis and 2 strains of Streptomyces isolated from old workers had strong inhibitory activity against Xylaria angulosa but weak inhibitory activity against Termitomyces sp.
5. These results indicated that the actinobacteria of O. formosanus may contribute to protecting termite fungal food via their asymmetric antifungal activities.

     

Seed–litter–position drives seedling establishment in grassland species under recurrent drought

Changes in land use and climate interfere with grassland ecosystem processes. Here I experimentally investigated the combined effects of land‐use change related litter cover and contrasting water supply on seedling emergence. In this context, the role of the initial relative position of seeds, i.e. seeds on top of the litter versus seeds beneath the litter in interaction with water supply has not been investigated so far. I hypothesised that facilitative effects of litter on seedling emergence occur when seeds are covered by litter and deteriorate when litter covers the ground and seeds fall on it (seeds on top of the litter). Further, I hypothesised that the importance of seed position for seedling emergence will increase under conditions of recurrent drought. I performed a controlled pot experiment on seedling emergence of three common European grassland species (Pimpinella saxifraga, Leontodon autumnalis, Sanguisorba officinalis) by experimental manipulations of litter and water availability. Seedling emergence under moist conditions showed no significant differences between each litter position compared to the control across species. In contrast, under recurrent drought, seedling emergence was significantly higher below the litter compared to seeds on top of the litter and the control (i.e. no litter). In abandoned land, seedling emergence may be limited when seeds fall on ground‐covering litter. In contrast, in grasslands with regular low‐intensity land use, seedling emergence may be enhanced when a moderate level of litter covers seeds at the end of the growing season. Protective mechanisms that occur with seeds positioned beneath litter are particularly important under recurrent drought.     

Intracolonial differences in gut bacterial community between worker and soldier castes of Coptotermes formosanus

Abstract The establishment of symbiotic relationships with intestinal microorganisms enables termites to thrive on recalcitrant substrates such as cellulose and wood. A termite colony is composed of several different castes which have distinct feeding habits. The soldiers, for example, cannot feed by themselves and depend on workers, who feed them with digested or semi‐digested foods. To investigate the influence of feeding habits on the bacterial symbionts, a comparative study of gut bacteria between worker and soldier castes of the termite Coptotermes formosanus was conducted. The bacterial communities of both castes were investigated using denaturing gradient gel electrophoresis (DGGE) and clonal analysis of 16S ribosomal DNA (rDNA). Both methods indicated Bacteroidetes was the common predominant group; the common dominant phylotype was affiliated with a reported uncultured Bacteroidetes phylotype (BCf1–03). There were significant differences in Bacteroidetes and Spirochaetes between two castes. Compared to the gut bacteria of workers, those of soldiers were lower in abundance and diversity of Bacteroidetes and slightly higher in Spirochaetes. Two phylotypes (W8, W11) affiliated to Bacteroidetes and two (W26, W29) affiliated to Spirochaetes were exclusively found in the DGGE profile of the worker caste. Bacteroidetes are assumed to be involved in fermentation of sugars and nitrogenous compounds as well as degradation of uric acid. Spirochaetes are supposed to aid in the functions of acetogenesis and N₂‐fixation. The different feeding habits between workers and soldiers of C. formosanus may explain the observed differences in the gut bacterial community.     

Panic escape polyethism in worker and soldier Coptotermes formosanus (Isoptera: Rhinotermitidae)

Termites were the first animal to form societies. Two hundred million years of evolution provide for a multitude of innate social behaviors that can be experimentally dissected. These fine‐tuned patterns of behavior are especially interesting when observing group decision making in the panic mode. In this study, we examined behavioral patterns of termites under panic conditions to gain insight into how an escape flow self‐organizes. One hundred worker and 10 soldier Coptotermes formosanus were released into agar plates. After a disturbance was created most workers followed each other and ran along the wall of dishes, thus forming a unidirectional escape flow, whereas soldiers showed a significantly higher frequency of moving to the center of the arena or on periphery of the escape flow as compared to workers. Agonistic behavior was usually observed as soldiers moved to center or periphery. This is the first report on the behavioral repertoire of termites when panicked, with details on the behavioral polymorphism of workers and soldiers during an escape.     

Physical contact between soldier and worker is essential in soldier self-regulation of Coptotermes formosanus (Isoptera, Rhinotermitidae)

In a termite society, the soldier proportion is regulated at a species-specific level with a seasonal fluctuation. A feedback mechanism in soldier proportion regulation is well recognized, while the proximate means by which the presence of soldiers inhibits the new soldier formation from workers remains mostly elusive. In the present study, some possible means for this inhibition were tested with the Formosan subterranean termite, Coptotermes formosanus Shiraki. The results showed that neither soldier-originated volatiles nor non-volatile chemicals without physical contact were responsible for the inhibition of new soldier formation. The essentiality of physical contact between workers and soldiers in soldier regulation was established. However, by physical contacts, as the soldier-replacement experiment showed, workers could not individually recognize or “count” soldiers to maintain a set soldier proportion. The possible feedback mechanisms for soldier regulation are discussed.     

Ground vegetation dynamics in mountain spruce (<Emphasis Type="Italic">Picea</Emphasis> <Emphasis Type="Italic">abies</Emphasis> (L.) Karsten) forests recovering after air pollution stress impact

Our study focuses on the ground vegetation dynamics and its dependence on microsite conditions in declined climax mountain Norway spruce forests during the recovery period (1995–2006) following upon the considerable decrease of SO₂ pollution. We showed that ground vegetation development shifted from prevailing mosses and vegetation-free sites covered with spruce litter to dominance by Avenella flexuosa during the earlier period of massive decline of the observed ecosystems. The expansion of Vaccinium myrtillus seems to occur mainly under the gradually defoliating tree crowns whereas larger canopy gaps and quickly deforested areas are more successfully colonized by grasses, especially Calamagrostis villosa. The gradual spruce stand decline, as well as the corresponding ground vegetation dynamics, proceeded until the end of the twentieth century. Afterwards, the ground vegetation responded to the interruption of trees dying and stopped its expansion on spruce litter microsites. Retreat of both dominant grasses accompanied by the remarkable increase in cover of mosses occurred.     

Soldier‐biased gene expression in a subterranean termite implies functional specialization of the defensive caste

In a termite colony, reproduction is typically monopolized by a small number of sexuals that are supported by reproductively altruistic soldiers and workers. We expect caste differentiation to be associated with clear‐cut differences in gene expression, and for these differences to reflect caste function and development. Here, we use RNA‐Sequencing to compare the gene expression profiles of sexual nymphs and two non‐reproductive helper castes (i.e., workers and soldiers) of the Eastern subterranean termite Reticulitermes flavipes. We found that of n = 93 genes that are strictly expressed as a function of caste, a majority (78%) show a soldier‐specific pattern. This conspicuous soldier‐bias in genome‐wide expression suggests that this defensively specialized caste is functionally well‐differentiated from both the reproductive and the other non‐reproductive caste of this species, despite a shared developmental program with workers. Gene ontology analysis supports the notion of functional specialization by soldiers, as soldier‐biased gene sets are enriched for novel biological processes. Whether this pattern reflects ancient or more recent bouts of selection for caste novelty at the gene‐regulatory level is not known, but because soldiers are sterile and thus have no direct fitness, any selection for novelty must have been mediated indirectly, through reproducing relatives.     

It is not only group size: Soldiers also modulate the tolerance to insecticide in termites (Blattodea: Isoptera)

The survivorship of social insects is known to increase with group size, even in situations of stress, such as starvation and exposure to insecticides. However, in termites, studies have been undertaken only with workers, disregarding the possible effect of soldiers. The role of soldiers in the termite colonies goes beyond defense, mainly in Nasutitermes species. It is already known that soldiers initiate the foraging as well as improve the decision-making of food resources. Here, we evaluated the effect of group size and the presence of soldiers on exposure to sublethal doses of the insecticide imidacloprid in Nasutitermes corniger (Termitidae: Nasutitermitinae). To do so, toxicity bioassays were undertaken initially to determine the dose of the insecticide required to kill 50% of the N. corniger population (LD₅₀) to be used in the main experiments. Survival bioassays were then carried out with termite groups, with and without soldiers, in different sizes (6, 10, 14, 22, 26, 30 and 60), exposed and nonexposed to insecticide. In general, the mean time to death of termites increases linearly with group size. However, the mean time to death in groups with soldiers was significantly longer only in groups exposed to the insecticide. Our results indicate that soldiers can help to increase the tolerance of nasute termite groups to insecticides, in addition to the group size, as already shown in the previous study. The size of the group and social context could, therefore, modulate behavioral and/or physiological responses that enhance the ability to survive under stressful situations.     

Inter- and intraspecific alarm response in the termiteMacrotermes subhyalinus (Rambur)

Summary In laboratory studies we analysed the reaction of aMacrotermes subhyalinus (Rambur) colony in contact with a small group of test termites placed at the head of a gallery system. As test termites we usedMacrotermes subhyalinus of the same nest,Macrotermes subhyalinus of a foreign nest and the related sympatric speciesMacrotermes bellicosus (Smeathman).Contact with an introduced group of foragers evokes a significant recruitment of workers and minor soldiers. The intensity of the response depends on the test groups we introduced. The highest recruitment results withM. bellicosus andM. subhyalinus from a foreign nest. During the recruitment the increase in the number of minor soldiers is greater than the increase in the number of workers, and as a result, the relative proportion of the soldiers in the population leaving the nest is significantly higher than before.Direct observations at the place of contact show thatM. bellicosus individuals are immediately attacked and killed. In the case ofM. subhyalinus from a foreign nest, not all individuals are killed; some are only intensively examined with the antennae or seized with the mandibles.M. subhyalinus from the same nest are rarely attacked. Mostly they are antennally inspected or enter the gallery system unhindered.The recruitment of termites from the nest is released by major workers which, after contact with the test termites, run back into the nest and provoke an alarm by tactile stimuli. On their way back into the nest the major workers, in all probability, lay a pheromone trail which leads the outgoing alarmed termites towards the place of contact.     

Fire ant predation on native and introduced subterranean termites in the laboratory: effect of high soldier number in Coptotermes formosanus

Abstract.

• 1 Native Reticulitermes spp. Holmgren and the introduced Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae) were exposed to predation in the laboratory by Solenopsis invicta Buren (Hymenoptera: Formicidae), a known predator of Reticulitermes.
• 2 In one experiment, both taxa contained the 2.4% soldiers commonly found in Reticulitermes. In a second experiment, Reticulitermes contained 2.4% and C.formosanus a more normal 18.3% soldiers.
• 3 When soldier proportions were equal, C.formosanus soldiers suffered less of a decrease in survivorship due to predation compared with Reticulitermes soldiers. Little difference was found between taxa in the effect of predation on termite workers or in the effect of termites on predator survivorship.
• 4 When soldier proportions were unequal, both C.formosanus castes suffered less from predation than those of Reticulitermes. Again, little difference was found in the effect of termites on ants.
• 5 C.formosanus showed less movement into new containers when soldier number was abnormally low, compared with both taxa with normal caste proportions.

     

Hunting strategy of a generalist ant species proposed as a biological control agent against termites

We studied the hunting behaviour of Myrmicaria opaciventris (Hymenoptera: Formicidae) in order to evaluate if it can be used as a biological control agent against the termites that damage sugarcane plantations. Hunting workers foraged in groups and recruited nestmates at short-range when they encountered large termite soldiers or groups of small termite workers. Differences in prey capture concerned the: (1) means of detection (from a distance or by contact); (2) termite body part seized (small termites seized by the body; large termites by an appendage); (3) percentages of prey abandoned; and (4) use of venom. The sting of the workers is spatulated implying a topical application of the venom on the prey. Large termites were stretched by several workers whose adherence to the substrate is facilitated by well-developed arolia and claws on the legs while others spread venom on the body and carved it up. An adaptation to termite capture was noted with a distribution of tasks between the workers which subdued prey, and those which transported it. In the former case, the workers easily eliminated termite soldiers, successively attacked several termite workers and even captured new individuals while holding the first ones captured between their mandibles before retrieving them all at once. The remaining individuals were retrieved by the transporting workers. Given this particularly effective predatory strategy, we concluded that, under certain conditions, M. opaciventris can be used as a biological control agent against termites.     

Biological changes in the Eastern subterranean termite, <Emphasis Type="Italic">Reticulitermes flavipes</Emphasis> (Isoptera,Rhinotermitidae) and its protozoa profile following starvation

The effects of starvation on survival, body mass, movement, other behavior, and the symbiotic protozoan community in Reticulitermes flavipes (Kollar) were investigated in a 40-day assay. Groups of 100 termites in their natural worker/soldier ratio (98/2) were tested. Starvation resulted in significantly greater mortality and induced more cannibalism of workers than in the control. Worker survival rate gradually declined to 58% along with an increasing rate of cannibalism during the first 30 days, and then quickly decreased to 5% along with an increasing rate of necrophagia after 30 days. In contrast, starvation had no significant effect on the survival of soldiers (60%) as compared to the control (90%) and starved workers did not cannibalize soldiers. Ten protozoan species residing in the hindgut were identified. When compared with field termites, the 40-day starvation eliminated 5 species (Trichonympha agilis, Pyrsonympha vertens, and P. major, D. gracilis, Holomastigotes elongatum), significantly reduced the numbers of two species (Dinenympha fimbriata, Spironympha kofoidi), had no effect on two species (Trichomitus trypanoides and Spirotrichonympha flagellata), and led to proliferation of one species (Monocercomonas sp.), whereas feeding on filter paper reduced the populations of five species (T. agilis, D. fimbriata, P. vertens, P. major, and S. flagellata) to different degrees. Workers surviving starvation had similar body mass and short-range movement speed to workers fed on filter paper and workers freshly collected from natural setting. The energy-demanding survival and walking indicated that they might obtain sufficient nutrients from cannibalizing other workers. The observed behaviors are important in helping to understand termite survival strategies and the mechanisms by which termites maintain their social structure under stressful conditions.     

Fine root decomposition rates do not mirror those of leaf litter among temperate tree species

Elucidating the function of and patterns among plant traits above ground has been a major research focus, while the patterns and functioning of belowground traits remain less well understood. Even less well known is whether species differences in leaf traits and their associated biogeochemical effects are mirrored by differences in root traits and their effects. We studied fine root decomposition and N dynamics in a common garden study of 11 temperate European and North American tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pseudotsuga menziesii, Quercus robur, Quercus rubra and Tilia cordata) to determine whether leaf litter and fine root decomposition rates are correlated across species as well as which species traits influence microbial decomposition above versus below ground. Decomposition and N immobilization rates of fine roots were unrelated to those of leaf litter across species. The lack of correspondence of above- and belowground processes arose partly because the tissue traits that influenced decomposition and detritus N dynamics different for roots versus leaves, and partly because influential traits were unrelated between roots and leaves across species. For example, while high hemicellulose concentrations and thinner roots were associated with more rapid decomposition below ground, low lignin and high Ca concentrations were associated with rapid aboveground leaf decomposition. Our study suggests that among these temperate trees, species effects on C and N dynamics in decomposing fine roots and leaf litter may not reinforce each other. Thus, species differences in rates of microbially mediated decomposition may not be as large as they would be if above- and belowground processes were working in similar directions (i.e., if faster decomposition above ground corresponded to faster decomposition below ground). Our results imply that studies that focus solely on aboveground traits may obscure some of the important mechanisms by which plant species influence ecosystem processes.