Og DeSouza

Environmental heterogeneity as a strategy for pest management in Eucalyptus plantations

The effect of environmental heterogeneity on the distribution and abundance of Lepidoptera was tested in Eucalyptus plantations interwoven with natural vegetation. Collections were carried out in Aracruz, ES, Brazil, in five sites along a 1000 m transect starting in a native woodland and penetrating stands of Eucalyptus grandis and Eucalyptus saligna. Collections were undertaken with light traps, three times a month, from April to August, 1993, recording the number of individuals per morphospecies in each site. A positive correlation was found between the number of sites where each species was recorded and its mean local abundance (r2 = 0.45; P < 0.01; n = 790), indicating that the more ubiquitous Lepidoptera species are also those that were locally more abundant. No pest species was recorded among those very abundant. Pest species generally presented moderate abundances and only three out of ten were found at all sites. It seems, therefore, that pest species were constrained by the heterogeneity conferred by the coexistence of Eucalyptus plantations and remnants of native vegetation.

Diet Segregation between Cohabiting Builder and Inquiline Termite Species.

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.

Mutual use of trail-following chemical cues by a termite host and its inquiline.

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.

Individual basis for collective behaviour in the termite, Cornitermes cumulans

Abstract Interactions among individuals in social groups lead to the emergence of collective behaviour at large scales by means of multiplicative non-linear effects. Group foraging, nest building and task allocation are just some well-known examples present in social insects. However the precise mechanisms at the individual level that trigger and amplify social phenomena are not fully understood. Here we show evidence of complex dynamics in groups of the termite, Cornitermes cumulans (Kollar) (Isoptera: Termitidae), of different sizes and qualitatively compare the behaviour observed with that exhibited by agent-based computer models. It is then concluded that certain aspects of social behaviour in insects have a universal basis common to interconnected systems and that this may be useful for understanding the temporal dynamics of systems displaying social behaviour in general.

The nature of alarm communication in Constrictotermes cyphergaster (Blattodea: Termitoidea: Termitidae): the integration of chemical and vibroacoustic signals

ABSTRACT 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 workers 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. Summary: We inspected the functional significance of both vibroacoustic and chemical communication channels in Constrictotermes cyphergaster (Termitidae: Nasutitermitinae), confirming the hypothesis that these are not exclusive but rather complementary processes.

Lévy flights and self-similar exploratory behaviour of termite workers: beyond model fitting.

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.

Allometric Scaling of Patrolling Rate and Nest Volume in Constrictotermes cyphergaster Termites: Hints on the Settlement of Inquilines.

Structural and functional traits of organisms are known to be related to the size of individuals and to the size of their colonies when they belong to one. Among such traits, propensity to inquilinism in termites is known to relate positively to colony size. Larger termitaria hold larger diversity of facultative inquilines than smaller nests, whereas obligate inquilines seem unable to settle in nests smaller than a threshold volume. Respective underlying mechanisms, however, remain hypothetical. Here we test one of such hypotheses, namely, that nest defence correlates negatively to nest volume in Constrictotermes cyphergaster termites (Termitidae: Nasutitermitinae). As a surrogate to defence, we used ‘patrolling rate’, i.e., the number of termite individuals attending per unit time an experimentally damaged spot on the outer wall of their termitaria. We found that patrolling rate decayed allometrically with increasing nest size. Conspicuously higher patrolling rates occurred in smaller nests, while conspicuously lower rates occurred in larger nests presenting volumes in the vicinity of the threshold value for the establishment of inquilinism. This could be proven adaptive for the host and guest. At younger nest age, host colonies are smaller and presumably more vulnerable and unstable. Enhanced defence rates may, hence, prevent eventual risks to hosts from inquilinism at the same time that it prevents inquilines to settle in a still unstable nest. Conversely, when colonies grow and maturate enough to stand threats, they would invest in priorities other than active defence, opening an opportunity for inquilines to settle in nests which are more suitable or less risky. Under this two-fold process, cohabitation between host and inquiline could readily stabilize.

Resource suitability modulating spatial co‐occurrence of soil‐forager termites (Blattodea: Termitoidea)

Resource can regulate animal foraging range, which in turn determines the chances of species co‐occurrence. Here, we addressed the question of whether resource determines the co‐occurrence of soil‐forager termite species (i.e. those foraging in subterranean tunnels). Eight quadrats (4 × 4 m) were marked in seven sites of Brazilian Atlantic rainforests, giving a total area sampled of 896 m2. Inside each quadrat, we measured the co‐occurrence of soil forager species and the resource suitability (N:C ratio of the soil and litter biomass). The number of records of more than one soil‐forager termite species at a single foraging spot, relative to the total number of foraging spots detected in each forest, was taken as a surrogate for spatial co‐occurrence. We tested whether termite co‐occurrence was mediated by random or nonrandom processes. Data were subjected to linear regression to test how the termite species co‐occurrence responds to resources. We compared this method with a null model analysis. Soil‐forager termites comprised 885 records, 20 species and 14 genera. From those records, 29% indicated species co‐occurrence. Co‐occurrence was not random: occurred more frequently when resource suitability was very high or very low. This result suggests an optimised use of space by termite communities.

Top‐down network analysis characterizes hidden termite–termite interactions

Abstract The analysis of ecological networks is generally bottom‐up, where networks are established by observing interactions between individuals. Emergent network properties have been indicated to reflect the dominant mode of interactions in communities that might be mutualistic (e.g., pollination) or antagonistic (e.g., host–parasitoid communities). Many ecological communities, however, comprise species interactions that are difficult to observe directly. Here, we propose that a comparison of the emergent properties from detail‐rich reference communities with known modes of interaction can inform our understanding of detail‐sparse focal communities. With this top‐down approach, we consider patterns of coexistence between termite species that live as guests in mounds built by other host termite species as a case in point. Termite societies are extremely sensitive to perturbations, which precludes determining the nature of their interactions through direct observations. We perform a literature review to construct two networks representing termite mound cohabitation in a Brazilian savanna and in the tropical forest of Cameroon. We contrast the properties of these cohabitation networks with a total of 197 geographically diverse mutualistic plant–pollinator and antagonistic host–parasitoid networks. We analyze network properties for the networks, perform a principal components analysis (PCA), and compute the Mahalanobis distance of the termite networks to the cloud of mutualistic and antagonistic networks to assess the extent to which the termite networks overlap with the properties of the reference networks. Both termite networks overlap more closely with the mutualistic plant–pollinator communities than the antagonistic host–parasitoid communities, although the Brazilian community overlap with mutualistic communities is stronger. The analysis raises the hypothesis that termite–termite cohabitation networks may be overall mutualistic. More broadly, this work provides support for the argument that cryptic communities may be analyzed via comparison to well‐characterized communities.

Termite cohabitation: the relative effect of biotic and abiotic factors on mound biodiversity

1. Termites are important ecosystem engineers that improve primary productivity in trees and animal diversity outside their mounds. However, their ecological relationship with the species nesting inside their mounds is poorly understood.