Chemical and vibratory signals used in alarm communication in the termite Reticulitermes flavipes (Rhinotermitidae)

Termites have evolved diverse defence strategies to protect themselves against predators, including a complex alarm communication system based on vibroacoustic and/or chemical signals. In reaction to alarm signals, workers and other vulnerable castes flee away while soldiers, the specialized colony defenders, actively move toward the alarm source. In this study, we investigated the nature of alarm communication in the pest Reticulitermes flavipes. We found that workers and soldiers of R. flavipes respond to various danger stimuli using both vibroacoustic and chemical alarm signals. Among the danger stimuli, the blow of air triggered the strongest response, followed by crushed soldier head and light flash. The crushed soldier heads, which implied the alarm pheromone release, had the longest-lasting effect on the group behaviour, while the responses to other stimuli decreased quickly. We also found evidence of a positive feedback, as the release of alarm pheromones increased the vibratory communication among workers and soldiers. Our study demonstrates that alarm modalities are differentially expressed between castes, and that the response varies according to the nature of stimuli

Complex alarm strategy in the most basal termite species

Studying basal taxa often allows shedding a light on the evolution of advanced representatives. The most basal termite species, Mastotermes darwiniensis, possesses unique morphological and behavioural traits, of which many remain scarcely studied. For these reasons, we conducted a comprehensive study of the alarm communication in this species and compared its components to behavioural modes described in other termites. In M. darwiniensis, the alarm is communicated by substrate-borne vibrations resulting from vertical vibratory movements. Another similar behaviour consists in longitudinal movements, by which the alarm is delivered to other termites in contact with alerted individual. Both these two behavioural modes could be used in synergy to create complex movements. M. darwiniensis also uses chemical alarm signals produced by labial gland secretion, in contrast to Neoisoptera in which this function is fulfilled exclusively by the frontal gland secretion. Moreover, we demonstrated in M. darwiniensis the presence of a positive feedback mechanism thought to occur exclusively in the crown group Termitidae. This positive feedback consists in both oscillatory movements of alerted individuals in response to alarm signals and release of alarm pheromone by excited soldiers. Our results confirm that M. darwiniensis is a remarkable example of mosaic evolution, as it combines many primitive and advanced features, and its alarm communication clearly belongs to the latter category

The oral gland, a new exocrine organ of termites

Termites have a rich set of exocrine glands. These glands are located all over the body, appearing in the head, thorax, legs and abdomen. Here, we describe the oral gland, a new gland formed by no more than a few tens of Class I secretory cells. The gland is divided into two secretory regions located just behind the mouth, on the dorsal and ventral side of the pharynx, respectively. The dominant secretory organelle is a smooth endoplasmic reticulum. Secretion release is under direct control of axons located within basal invaginations of the secretory cells. The secretion is released through a modified porous cuticle located at the mouth opening. We confirmed the presence of the oral gland in workers and soldiers of several wood- and soil-feeding species of Rhinotermitidae and Termitidae, suggesting a broader distribution of the oral gland among termites. The oral gland is the smallest exocrine gland described in termites so far. We hypothesise that the oily secretion can either ease the passage of food or serve as a primer pheromone

White-gutted soldiers: simplification of the digestive tube for a non-particulate diet in higher Old World termites (Isoptera: Termitidae)

Previous observations have noted that in some species of higher termites the soldier caste lacks pigmented particles in its gut and, instead, is fed worker saliva that imparts a whitish coloration to the abdomen. In order to investigate the occurrence of this trait more thoroughly, we surveyed a broad diversity of termite specimens and taxonomic descriptions from the Old World subfamilies Apicotermitinae, Cubitermitinae, Foraminitermitinae, Macrotermitinae, and Termitinae. We identified 38 genera that have this “white-gutted” soldier (WGS) trait. No termite soldiers from the New World were found to possess a WGS caste. Externally, the WGS is characterized by a uniformly pale abdomen, hyaline gut, and proportionally smaller body-to-head volume ratio compared with their “dark-gutted” soldier (DGS) counterparts found in most termitid genera. The WGS is a fully formed soldier that, unlike soldiers in other higher termite taxa, has a small, narrow, and decompartmentalized digestive tube that lacks particulate food contents. The presumed saliva-nourished WGS have various forms of simplified gut morphologies that have evolved at least six times within the higher termites

Substrate evaporation drives collective construction in termites

AbstractTermites build complex nests which are an impressive example of self-organization. We know that the coordinated actions involved in the construction of these nests by multiple individuals are primarily mediated by signals and cues embedded in the structure of the nest itself. However, to date there is still no scientific consensus about the nature of the stimuli that guide termite construction, and how they are sensed by termites. In order to address these questions, we studied the early building behavior ofCoptotermes formosanustermites in artificial arenas, decorated with topographic cues to stimulate construction. Pellet collections were evenly distributed across the experimental setup, compatible with a collection mechanism that is not affected by local topography, but only by the distribution of termite occupancy (termites pick pellets at the positions where they are). Conversely, pellet depositions were concentrated at locations of high surface curvature and at the boundaries between different types of substrate. The single feature shared by all pellet deposition regions was that they correspond to local maxima in the evaporation flux. We can show analytically and we confirm experimentally that evaporation flux is directly proportional to the local curvature of nest surfaces. Taken together, our results indicate that surface curvature is sufficient to organize termite building activity, and that termites likely sense curvature indirectly through substrate evaporation. Our findings reconcile the apparently discordant results of previous studies.</jats:p

Breaking the cipher: ant eavesdropping on the variational trail pheromone of its termite prey

Predators may eavesdrop on their prey using innate signals of varying nature. In regards to social prey, most of the prey signals are derived from social communication and may therefore be highly complex. The most efficient predators select signals that provide the highest benefits. Here, we showed the use of eusocial prey signals by the termite-raiding ant Odontoponera transversaO. transversa selected the trail pheromone of termites as kairomone in several species of fungus-growing termites (Termitidae: Macrotermitinae: Odontotermes yunnanensis, Macrotermes yunnanensis, Ancistrotermes dimorphus). The most commonly predated termite, O. yunnanensis, was able to regulate the trail pheromone component ratios during its foraging activity. The ratio of the two trail pheromone compounds was correlated with the number of termites in the foraging party. (3Z)-Dodec-3-en-1-ol (DOE) was the dominant trail pheromone component in the initial foraging stages when fewer termites were present. Once a trail was established, (3Z,6Z)-dodeca-3,6-dien-1-ol (DDE) became the major recruitment component in the trail pheromone and enabled mass recruitment of nest-mates to the food source. Although the ants could perceive both components, they revealed stronger behavioural responses to the recruitment component, DDE, than to the common major component, DOE. In other words, the ants use the trail pheromone information as an indication of suitable prey abundance, and regulate their behavioural responses based on the changing trail pheromone component. The eavesdropping behaviour in ants therefore leads to an arms race between predator and prey where the species specific production of trail pheromones in termites is targeted by predatory ant species