Eusocial insects frequently face death of colony members as a consequence of living in large groups where the success of the colony is not dependent on the fate of the individual. Whereas death of conspecifics commonly triggers aversion in many group-living species due to risk of pathogens, eusocial insects perform cooperative corpse management. The causes and social context of the death, as well as feeding and nesting ecology of the species, influence the way that corpses are treated. The corpse itself releases cues that dictate the colony's response. As a result, social insects exhibit behavioural responses that promote disease resistance, colony defence and nutrient recycling. Corpse management represents a unique adaption that enhances colony success, and is another factor that has enabled eusocial insects to be so successful. In this review, we summarize the causes of death, the sensory detection of death and corpse management strategies of social insects. In addition, we provide insights into the evolution of behavioural response to the dead and the ecological relevance of corpse management. This article is part of the theme issue ‘Evolutionary thanatology: impacts of the dead on the living in humans and other animals’.

Reproductive conflicts are common in insect societies where helping castes retain reproductive potential. One of the mechanisms regulating these conflicts is policing, a coercive behaviour that reduces direct reproduction by other individuals. In eusocial Hymenoptera (ants, bees and wasps), workers or the queen act aggressively towards fertile workers, or destroy their eggs. In many termite species (order Blattodea), upon the death of the primary queen and king, workers and nymphs can differentiate into neotenic reproductives and inherit the breeding position. During this process, competition among neotenics is inevitable, but how this conflict is resolved remains unclear. Here, we report a policing behaviour that regulates reproductive division of labour in the eastern subterranean termite, Reticulitermes flavipes . Our results demonstrate that the policing behaviour is a cooperative effort performed sequentially by successful neotenics and workers. A neotenic reproductive initiates the attack of the fellow neotenic by biting and displays alarm behaviour. Workers are then recruited to cannibalize the injured neotenic. Furthermore, the initiation of policing is age-dependent, with older reproductives attacking younger ones, thereby inheriting the reproductive position. This study provides empirical evidence of policing behaviour in termites, which represents a convergent trait shared between eusocial Hymenoptera and Blattodea.

Antennae are the primary sensory organs in insects, where a variety of sensilla are distributed for the perception of the chemical environment. In eusocial insects, colony function is maintained by a division of labor between reproductive and non-reproductive castes, and chemosensation is essential for regulating their specialized social activities. Several social species in Hymenoptera display caste-specific characteristics in antennal morphology and diversity of sensilla, reflecting their differential tasks. In termites, however, little is known about how the division of labor is associated with chemosensory morphology among castes. Using light and scanning electron microscopy, we performed antennal morphometry and characterized the organization of sensilla in reproductive (female and male alates) and non-reproductive (worker and soldier) castes in the Formosan subterranean termite, Coptotermes formosanus Shiraki. Here, we show that the antennal sensilla in alates are twice as abundant as in workers and soldiers, along with the greater number of antennal segments and antennal length in alates. However, all castes exhibit the same types of antennal sensilla, including basiconicum, campaniformium, capitulum, chaeticum I, chaeticum II, chaeticum III, marginal, trichodeum I, and trichodeum I. The quantitative composition of sensilla diverges between reproductive and non-reproductive castes, but not between female and male alates or between worker and soldier castes. The sensilla display spatial-specific distribution, with basiconicum exclusively and capitulum predominantly found on the ventral side of antennae. In addition, the abundance of chemosensilla increases toward the distal end of antennae in each caste. This research provides morphological signatures of chemosensation and their implications for the division of labor, and suggests future neurophysiological and molecular studies to address the mechanisms of chemical communication in termites.

Sexual attraction and perception are crucial for mating and reproductive success. In Drosophila melanogaster , the male-specific isoform of Fruitless (Fru), Fru ^M , is a known master neuro-regulator of innate courtship behavior to control the perception of sex pheromones in sensory neurons. Here, we show that the non–sex-specific Fru isoform (Fru ^COM ) is necessary for pheromone biosynthesis in hepatocyte-like oenocytes for sexual attraction. Loss of Fru ^COM in oenocytes resulted in adults with reduced levels of cuticular hydrocarbons (CHCs), including sex pheromones, and show altered sexual attraction and reduced cuticular hydrophobicity. We further identify Hepatocyte nuclear factor 4 ( Hnf4 ) as a key target of Fru ^COM in directing fatty acid conversion to hydrocarbons. Fru or Hnf4 depletion in oenocytes disrupts lipid homeostasis, resulting in a sex-dimorphic CHC profile that differs from doublesex- and transformer- dependent CHC dimorphism. Thus, Fru couples pheromone perception and production in separate organs to regulate chemosensory communications and ensure efficient mating behavior.