Abstract. While the key role of termites in the decomposition of litter in the tropics has been acknowledged for a long time, much less information exists on their importance in the recycling of dung of primary consumers, especially herbivores. A review of published studies shows that a diverse group of termites (at least 126 species) has been reported to feed on a wide range of mammalian dung (18 species). Predominantly, wood-feeding and polyphagous wood-litter feeding species were found to feed also frequently on dung. Moreover, we found that termites can quickly remove large amounts of mammalian dung, especially in the dry season, when on average about 1/3 of the dung deposited in a given habitat is removed by termites within one month (with the highest rates observed in savannas). No distinctive preference for mammalian dung over other organic food sources was observed for fungusgrowing termites (Macrotermitinae), whereas the majority of the non-fungus growing taxa studied prefer dung over other food. As termites bring large quantities of dung below the soil surface, disturb and enrich soils with nutrients, dung feeding by termites appears to be a previously underestimated process important in the functioning of tropical ecosystems.
Aim In this paper we aim to show that proportional sampling can detect species–area relationships (SARs) more effectively than uniform sampling. We tested the contribution of alpha and beta diversity in ant communities as explanations for the SAR. Location Tropical forest remnants in Viçosa, Minas Gerais, Brazil (20 °45′ S, 42 °50′ W). Methods We sampled 17 forest remnants with proportional sampling. To disentangle sampling effects from other mechanisms, species richness was fitted in a model with remnant size, number of samples (sampling effects) and an interaction term. Results A SAR was observed independent of the number of samples, discarding sampling effects. Alpha diversity was not influenced by remnant size, and beta diversity increased with remnant size; evidence to the fact that habitat diversity within remnants could be the dominant cause of the SAR. Such a relationship between beta diversity and remnant area may have also arisen due to the combined effects of territoriality and aggregation of ant species. Main conclusions The proposed model, together with proportional sampling, allowed the distinction between sampling effects and other mechanisms.
Alarm signalling is of paramount importance to communication in all social insects. In termites, vibroacoustic and chemical alarm signalling are bound to operate synergistically but have never been studied simultaneously in a single species. Here, we inspected the functional significance of both communication channels in Constrictotermes cyphergaster (Termitidae: Nasutitermitinae), confirming the hypothesis that these are not exclusive, but rather complementary processes. In natural situations, the alarm predominantly attracts soldiers, which actively search for the source of a disturbance. Laboratory testing revealed that the frontal gland of soldiers produces a rich mixture of terpenoid compounds including an alarm pheromone. Extensive testing led to identification of the alarm pheromone being composed of abundant monoterpene hydrocarbons (1S)-α-pinene and myrcene, along with a minor component, (E)-β-ocimene. The vibratory alarm signalling consists of vibratory movements evidenced as bursts; a series of beats produced predominantly by soldiers. Exposing termite groups to various mixtures containing the alarm pheromone (crushed soldier heads, frontal gland extracts, mixture of all monoterpenes, and the alarm pheromone mixture made of standards) resulted in significantly higher activity in the tested groups and also increased intensity of the vibratory alarm communication, with the responses clearly dose-dependent. Lower doses of the pheromone provoked higher numbers of vibratory signals compared to higher doses. Higher doses induced long-term running of all termites without stops necessary to perform vibratory behaviour. Surprisingly, even crushed worker heads led to low (but significant) increases in the alarm responses, suggesting that other unknown compound in the worker's head is perceived and answered by termites. Our results demonstrate the existence of different alarm levels in termites, with lower levels being communicated through vibratory signals, and higher levels causing general alarm or retreat being communicated through the alarm pheromone.
How do termite inquilines manage to cohabit termitaria along with the termite builder species? With this in mind, we analysed one of the several strategies that inquilines could use to circumvent conflicts with their hosts, namely, the use of distinct diets. We inspected overlapping patterns for the diets of several cohabiting Neotropical termite species, as inferred from carbon and nitrogen isotopic signatures for termite individuals. Cohabitant communities from distinct termitaria presented overlapping diet spaces, indicating that they exploited similar diets at the regional scale. When such communities were split into their components, full diet segregation could be observed between builders and inquilines, at regional (environment-wide) and local (termitarium) scales. Additionally, diet segregation among inquilines themselves was also observed in the vast majority of inspected termitaria. Inquiline species distribution among termitaria was not random. Environmental-wide diet similarity, coupled with local diet segregation and deterministic inquiline distribution, could denounce interactions for feeding resources. However, inquilines and builders not sharing the same termitarium, and thus not subject to potential conflicts, still exhibited distinct diets. Moreover, the areas of the builder’s diet space and that of its inquilines did not correlate negatively. Accordingly, the diet areas of builders which hosted inquilines were in average as large as the areas of builders hosting no inquilines. Such results indicate the possibility that dietary partitioning by these cohabiting termites was not majorly driven by current interactive constraints. Rather, it seems to be a result of traits previously fixed in the evolutionary past of cohabitants.
Termite nests are often secondarily inhabited by other termite species ( = inquilines) that cohabit with the host. To understand this association, we studied the trail-following behaviour in two Neotropical species, Constrictotermes cyphergaster (Termitidae: Nasutitermitinae) and its obligatory inquiline, Inquilinitermes microcerus (Termitidae: Termitinae). Using behavioural experiments and chemical analyses, we determined that the trail-following pheromone of C. cyphergaster is made of neocembrene and (3Z,6Z,8E)-dodeca-3,6,8-trien-1-ol. Although no specific compound was identified in I. microcerus, workers were able to follow the above compounds in behavioural bioassays. Interestingly, in choice tests, C. cyphergaster prefers conspecific over heterospecific trails while I. microcerus shows the converse behaviour. In no-choice tests with whole body extracts, C. cyphergaster showed no preference for, while I. microcerus clearly avoided heterospecific trails. This seems to agree with the hypothesis that trail-following pheromones may shape the cohabitation of C. cyphergaster and I. microcerus and reinforce the idea that their cohabitation is based on conflict-avoiding strategies.
This paper describes a study on termites, investigating the relationship between increasing group size and individual worker longevity under resource-deprived conditions. It was found that survival was significantly lower for isolated individuals and higher for individuals in bigger group sizes, suggesting that social interactions play an important role in the mechanisms leading to longer survival. A computer model, incorporating individual interactions among mobile cellular automata is presented along with experiments.
Animal movements have been related to optimal foraging strategies where self-similar trajectories are central. Most of the experimental studies done so far have focused mainly on fitting statistical models to data in order to test for movement patterns described by power-laws. Here we show by analyzing over half a million movement displacements that isolated termite workers actually exhibit a range of very interesting dynamical properties –including Lévy flights– in their exploratory behaviour. Going beyond the current trend of statistical model fitting alone, our study analyses anomalous diffusion and structure functions to estimate values of the scaling exponents describing displacement statistics. We evince the fractal nature of the movement patterns and show how the scaling exponents describing termite space exploration intriguingly comply with mathematical relations found in the physics of transport phenomena. By doing this, we rescue a rich variety of physical and biological phenomenology that can be potentially important and meaningful for the study of complex animal behavior and, in particular, for the study of how patterns of exploratory behaviour of individual social insects may impact not only their feeding demands but also nestmate encounter patterns and, hence, their dynamics at the social scale.
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