Cuticular hydrocarbon profile for queen recognition in the termite Reticulitermes speratus

Chemical communication underlies the sophisticated colony organization of social insects. In these insects, cuticular hydrocarbons (CHCs) play central roles in nestmate, task, and caste recognition, which contribute to maintenance of the social and reproductive division of labor. Queen-specific CHCs reflect queen fertility status and function as a queen recognition pheromone, triggering aggregation responses around the queens. However, there are only a few studies about the royal recognition mechanism in termites, and particularly, no study has reported about queen-specific CHCs in the species using asexual queen succession (AQS) system, in which the primary queen is replaced by neotenic queens produced parthenogenetically. In this study, we identified the CHC pheromone for neotenic queen recognition in the AQS termite species Reticulitermes speratus. Gas chromatography-mass spectrometry analyses revealed that the relative amount of n-pentacosane was disproportionately greater in the CHC profiles of queens than in the CHC profiles of kings, soldiers, and workers. Furthermore, we investigated the cuticular chemicals of the queen aggregate workers; bioassays demonstrated that n-pentacosane shows a worker arrestant activity in the presence of workers’ cuticular extract. These results suggest that R. speratus workers identify whether each individual is a neotenic queen by recognizing the relatively higher ratio of n-pentacosane in the conspecific CHC background. Moreover, they suggest that termites have evolved queen recognition behavior, independently of social hymenopterans.

Queen succession conflict in the paper wasp Polistes dominula is mitigated by age-based convention

Reproduction in cooperative animal groups is often dominated by one or a few individuals, with the remaining group members relegated to nonreproductive helping roles. This reproductive skew can evolve if helpers receive fitness benefits such as potential future inheritance of the breeding position, but the mechanisms by which inheritance is determined are not well resolved. Polistes paper wasps form highly reproductively skewed groups and inheritance of the breeding position is likely to play a key role in the maintenance of this social structure, making them excellent models for the processes by which simple societies are maintained. Reproductive succession is thought to be determined via an age-based convention in some Polistes species, but there is also evidence for contest-based succession systems in which the replacement queen uses physical aggression to overpower and thereby subordinate her nestmates. Here, we provide evidence that queen succession in colonies of the European paper wasp Polistes dominula is determined via convention rather than contest, with little disruption to the colony’s social functioning. We use queen removal experiments and fine-scale behavioral analyses to confirm that age is a strong predictor of succession, and that behavioral responses to queen removal are restricted to the oldest individuals rather than being experienced equally across the group. We provide the most comprehensive and detailed experimental analysis on the dynamics of breeder succession in a cooperatively breeding invertebrate to date, thereby shedding light on the mechanisms by which animal societies are able to maintain cohesion in the face of within-group conflict.

Facultative asexual reproduction and genetic diversity of populations in the humivorous termite Cavitermes tuberosus

Termite colonies are typically founded by a pair of sexually reproducing dispersers, which can sometimes be replaced by some of their offspring. Some Reticulitermes and Embiratermes species routinely practice asexual queen succession (AQS): the queen is replaced by neotenic daughters produced by parthenogenesis, which mate with the primary king. Here, to cast light on the evolution of AQS, we investigated another candidate species, Cavitermes tuberosus (Termitinae). Of 95 nests, 39 contained a primary queen and 28 contained neotenic females (2–667 individuals), usually with the primary king. Microsatellite analyses confirmed that colonies were initiated by single pairs after large dispersal flights. More than 80% of the neotenic females were of exclusively maternal origin and completely homozygous, suggesting automictic parthenogenesis with gamete duplication. Conversely, workers, soldiers, and most alates and primary reproductives were produced sexually. AQS often occurs late, after colonies have reached maturity, whereas early AQS in other species may boost the young colony's growth rate. We suggest additional benefits of AQS in C. tuberosus , related with a smaller size, lesser stability and higher mobility of colonies. Our data add to the phylogenetical dispersion and diversity of modalities of AQS in termites, supporting a multiple evolutionary origin of this process.

Physiological reason for ceasing growth of unfertilized eggs produced by unmated queens in the subterranean termite Reticulitermes chinensis

In Reticulitermes chinensis, a close relative of R. speratus with asexual queen succession, unfertilized eggs can be produced but are not incubated. To explain this phenomenon, we analysed the physiological differences between unfertilized eggs/unmated queens and fertilized eggs/mated queens. Fertilized eggs consumed significantly larger quantities of five amino acids (Cys, Met, Ile, Leu and Tyr), Ca, protein and cholesterol during incubation. The higher levels of four trace elements (Na, K, Zn and Fe) in fertilized eggs and their lower levels in mated queens indicated that mated queens might transfer these trace elements to fertilized eggs to complete incubation. The higher levels of Mn, triglycerides and serotonin in mated queens and higher levels of Mn and glucose in fertilized eggs suggested that these substances are very important for normal ovarian and embryonic growth. The different expression of three reproductive genes (vtgl, rabil and JHE1) suggested that they might be involved in the regulation of ovarian and embryonic growth. Overall, changes in these physiological indices may substantially affect ovarian and embryonic growth and prohibit the incubation of unfertilized eggs in R. chinensis.