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"Termites Habitations."
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Termites
\"Carefully leveled text and vibrant photographs introduce early readers to termites and the structures they build together\"-- Provided by publisher.
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Nonaggressive behavior: A strategy employed by an obligate nest invader to avoid conflict with its host species
In addition to its builders, termite nests are known to house a variety of secondary opportunistic termite species so‐called inquilines, but little is known about the mechanisms governing the maintenance of these symbioses. In a single nest, host and inquiline colonies are likely to engage in conflict due to nestmate discrimination, and an intriguing question is how both species cope with each other in the long term. Evasive behaviour has been suggested as one of the mechanisms reducing the frequency of host‐inquiline encounters, yet, the confinement imposed by the nests' physical boundaries suggests that cohabiting species would eventually come across each other. Under these circumstances, it is plausible that inquilines would be required to behave accordingly to secure their housing. Here, we show that once inevitably exposed to hosts individuals, inquilines exhibit nonthreatening behaviours, displaying hence a less threatening profile and preventing conflict escalation with their hosts. By exploring the behavioural dynamics of the encounter between both cohabitants, we find empirical evidence for a lack of aggressiveness by inquilines towards their hosts. Such a nonaggressive behaviour, somewhat uncommon among termites, is characterised by evasive manoeuvres that include reversing direction, bypassing and a defensive mechanism using defecation to repel the host. The behavioural adaptations we describe may play an important role in the stability of cohabitations between host and inquiline termite species: by preventing conflict escalation, inquilines may improve considerably their chances of establishing a stable cohabitation with their hosts. We investigate the behavioral dynamics of the encounter between an obligate inquiline termite and its host termite species. Hosts are aggressive toward inquilines. When threatened or even physically injured by hosts, inquilines react with low aggressiveness, exhibiting a repertoire of evasive manoeuvres that prevent conflict escalation. While reducing costs of conflict, the nonthreatening behavior exhibited by inquilines may facilitate the stability of cohabitations with hosts in the long term.
Journal Article
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.
Journal Article
Smells Like Home: Chemically Mediated Co-Habitation of Two Termite Species in a Single Nest
Termite nests often are referred to as the most elaborate constructions of animals. However, some termite species do not build a nest at all and instead found colonies inside the nests of other termites. Since these so-called inquilines do not need to be in direct contact with the host population, the two colonies usually live in separate parts of the nest. Adaptations of both the inquiline and its host are likely to occur to maintain the spatial exclusion and reduce the costs of potential conflicts. Among them, mutual avoidance, based on chemical cues, is expected. We investigated chemical aspects of cohabitation between
Constrictotermes cavifrons
(Nasutitermitinae) and its obligatory inquiline
Inquilinitermes inquilinus
(Termitinae). Inquiline soldiers produce in their frontal glands a blend of wax esters, consisting of the C
12
alcohols (3
Z
)-dodec enol, (3
Z
,6
Z
)-dodecadienol, and dodecanol, esterified with different fatty acids. The C
12
alcohols appear to be cleaved gradually from the wax esters, and they occur in the frontal gland, in soldier headspace, and in the walls of the inquiline part of the nest. Electrophysiological experiments revealed that (3
Z
)-dodecenol and (3
Z
,6
Z
)-dodecadienol are perceived by workers of both species. Bioassays indicated that inquiline soldier heads, as well as the two synthetic compounds, are attractive to conspecific workers and elicit an arresting behavior, while host soldiers and workers avoid these chemicals at biologically relevant amounts. These observations support the hypothesis that chemically mediated spatial separation of the host and the inquiline is an element of a conflict-avoidance strategy in these species.
Journal Article
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.
Journal Article
Termitarium volume as a determinant of invasion by obligatory termitophiles and inquilines in the nests of Constrictotermes cyphergaster (Termitidae, Nasutitermitinae)
A range of organisms can be found inside termite nests where the degree of association can vary from facultative to obligatory dependence. Studies of the dynamics of nest invasion are still unresolved, so how and when cohabitants enter termite nests remain open questions. This study analyzed one specific aspect of the dynamics of termite nest invasion by obligatory termitophiles and inquilines, i.e., whether cohabitants were more likely to invade a nest when it reached a critical nest size. We collected 36 Constrictotermes cyphergaster nests of different sizes and sampled their cohabitant fauna. Our results indicated that the invasion of C. cyphergaster nests by obligatory termitophiles and inquilines was dependent on nest size. There appeared to be a critical nest size above which nests were more prone to invasion. Above this size, there was a significantly higher likelihood of finding obligatory cohabitants. Termitophile species were observed in nests ≥2.2 L, whereas inquiline species were only occur in nests ≥13.6 L. This may indicate that the obligatory cohabitants studied here did not occupy C. cyphergaster nests at random and that they were dependent on features that made these nests suitable for cohabitation, which are linked to colony development.
Journal Article
Top‐down network analysis characterizes hidden termite–termite interactions
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. Many species–species interactions are difficult to observe directly. Here we propose that comparing the network topology of these obfuscated communities to the network topology of well‐studied, detail‐rich communities will provide insight into the structure of the obfuscated communities. We examine termite mound cohabitation as a case study; the analysis raises the hypothesis that the interactions contained therein may be overall mutualistic.
Journal Article
Diagram: Termite Mound Interior
two illustrations of the interior of a termite mound by Adam Weiskind
Web Resource