Juvenile hormone affects age polyethism, ovarian status, and cuticular hydrocarbon profile in workers of a Polybia occidentalis wasp

Division of labor is one of the most striking features in the evolution of eusociality. Juvenile hormone (JH) mediates reproductive status and aggression among nestmates in primitively eusocial Hymenoptera (species without morphologically distinct castes). In highly social species it has apparently lost its gonadotropic role and primarily regulates the division of labor in the worker caste. Polybia occidentalis, a Neotropical swarm-founding wasp, is an ideal model to understand how JH levels mirror social context and reproductive opportunities because of the absence of a clear morphological caste dimorphism. In this study, we tested the hypothesis that JH influences division of labor, ovary activation and cuticular hydrocarbon profiles of workers. Our observations confirmed that JH analog (methoprene) and precocene affected the cuticular chemical profile associated with the age polyethism. Also, methoprene and precocene-I influenced differently ovarian activation of treated females (individuals treated with methoprene expressed more activated ovaries while precocene treatment did not have significant effect). These results suggest that different hormonal levels induce a differential expression in the cuticular chemical profiles associated with the workers’ age polyethism, which may be essential for keeping the social cohesion among workers throughout their lives in the colony. Furthermore, JH is likely to play a gonadotropic role in P. occidentalis. JH has apparently undergone certain modifications in social Hymenoptera, presenting multifaceted functions in different species.

Temporal division of labor in an aphid social system

Temporal division of labor, or age polyethism, in which altruistic caste individuals change their tasks with aging, is widely found in bees and ants (Hymenoptera) and also in other social insects. Here we report the discovery of elaborate age polyethism in a social aphid (Hemiptera). Tuberaphis styraci is a gall-forming aphid in which monomorphic first instar nymphs differentiate into normal nymphs and soldiers upon second instar molt. Soldiers neither grow nor reproduce but perform gall cleaning and colony defense. Using an artificial diet rearing system, we collected age-defined groups of soldiers and monitored their social behaviors. We observed that young soldiers tend to clean whereas old soldiers preferentially attack, thereby verifying age-dependent task switching from housekeeping to defense. Strategic sampling, age estimation and behavioral observation of soldiers from natural galls revealed that (1) young cleaning soldiers tend to inhabit upper gall regions with adult insects, (2) old attacking soldiers tend to be distributed in lower gall regions, particularly around the gall openings, and (3) the gall structure is linked to intra-nest movement, aging and task switching of soldiers in an adaptive manner. These results highlight an evolutionary parallelism comparable to the sophisticated temporal division of labor observed in honeybee colonies.

Age Polyethism in Atta sexdens (Linnaeus) (Hymenoptera: Formicidae)

This study aimed to verify age polyethism occurrence in medium-sized (cephalic capsule = 2.3 ± 0.21 mm) and small-sized (cc = 1.4 ± 0.10 mm) workers from Atta sexdens (Linnaeus) colonies. Four laboratory colonies were used, and they were maintained at 25 ± 2 °C, with 75 ± 3% relative humidity and a 12-hour photoperiod. Workers from these colonies were marked after their emergence and observed throughout their lifetime to determine which tasks they performed. The number of ants performing each activity was analyzed using linear mixed-effect models (LME), considering the temporal effect and the block design (colonies). We found that fungal garden maintenance tasks were frequent for both sizes, but their occurrence decreased significantly from the ninth week. The foraging activity occurred gradually in both sizes, with stabilization in the number of workers from the fourth week onwards and declined in the last three weeks of lifespan. Waste management tasks occurred throughout life but were more frequent during the first two weeks of life, in both medium and small workers. Therefore, age polyethism may be related to all activities; however, foraging tasks presented a distinct pattern compared to tasks in the fungus garden and refuse dump, where younger ants were more frequently observed.

The effect of age on non-reproductive division of labour in the tropical primitively eusocial wasp, Ropalidia cyathiformis

Division of labour among workers (non-reproductive division of labour), a characteristic feature of eusocial insects enables the efficient functioning of their colonies. In many advanced insect societies division of labour is based on age (age polyethism). Primitively eusocial insects however are believed to have a weak age polyethism. Here we investigated the role of age in non-reproductive division of labour in the tropical primitively eusocial wasp, Ropalidia cyathiformis and compared it with that in Ropalidia marginata, a congeneric species that exhibits relatively strong age polyethism. Age had a significant effect on the first performance of the four tasks studied; tasks were initiated in the sequence feed larva, build, bring food and bring building material. We measured task performance as the absolute frequency of tasks performed (FTP) and the probability of performing a task relative to other tasks (PTP) and age as absolute age in days since eclosion as well as relative age compared to nestmates. FTP varied significantly with both absolute and relative age, although absolute age explained more variance. PTP varied significantly with absolute age but not always with relative age. This is contrary to R. marginata, where more variation is explained by relative age than by absolute age. There was no trade-off between intranidal and extranidal tasks in R. cyathiformis unlike in R. marginata where the frequency of intranidal tasks decreased and that of extranidal tasks increased with age. We conclude that age polyethism is weak and less flexible in R. cyathiformis compared to that in R. marginata.

The role and significance of stingless bees (Hymenoptera: Apiformes: Meliponini) in the natural environment

This article refers to the biology and ecology of stingless bees (Meliponini), living in tropical and subtropical areas. Similar to honey bees (Apis mellifera), stingless bees (Meliponini) belong to the category of proper social insects and are at the highest level of social development. This group of insects comprises about 500 species and they are the most common bees pollinating the native plants in many tropical areas. Families of stingless bees are usually quite numerous, reaching up to 100,000 individuals. They are characterised by polymorphism, age polyethism and perennialism. This article presents the structural complexity of natural nesting of these tropical insects and their ability to settle in artificial nest traps. The main significance of stingless bees for humans is their role in the natural environment as pollinators, which is an essential factor influencing biodiversity.

Changing of the guard: mixed specialization and flexibility in nest defense (Tetragonisca angustula)

Task allocation is a central challenge of collective behavior in a variety of group-living species, and this is particularly the case for the allocation of social insect workers for group defense. In social insects, both benefits and considerable costs are associated with the production of specialized soldiers. We asked whether colonies mitigate costs of production of specialized soldiers by simultaneously employing behavioral flexibility in nonspecialist workers that can augment defense capabilities at short time scales. We studied colonies of the stingless bee Tetragonisca angustula, a species that has 2 discrete nest-guarding tasks typically performed by majors: hovering guarding and standing guarding. Majors showed age polyethism across nest-guarding tasks, first hovering and then changing to the task of standing guarding after 1 week. Colonies were also able to reassign minors to guarding tasks when majors were experimentally removed. Replacement guards persisted in nest defense tasks until colonies produced enough majors to return to their initial state. Tetragonisca angustula colonies thus employed a coordinated set of specialization strategies in nest defense: morphologically specialized soldiers, age polyethism among soldiers within specific guarding tasks, and rapid flexible reallocation of nonspecialists to guarding during soldier loss. This mixed strategy achieves the benefits of a highly specialized defensive force while maintaining the potential for rapid reinforcement when soldiers are lost or colonies face unexpectedly intense attack.

Interactions between leaf-cutter ants and fungus garden: Effects of division of labor, age polyethism, and egg cannibalism

Division of labor (DOL), age polyethism, and egg cannibalism all play roles in shaping colony-level population dynamics in social insect colonies. The ways in which these mechanisms interact with one another to shape population dynamics is not currently understood. In this study, we examine how these mechanisms influence population dynamics in colonies of fungus-gardening leaf-cutter ants by developing and studying two sets of models: (1) We study age polyethism contribution to the dynamics of this multi-species interaction model which incorporates mechanisms of DOL; (2) We explore effects of egg cannibalism in colony dynamics and understand how to model such social conflict behavior realistically using different functional responses. Our results suggest that: (a) Age polyethism is important to keep stable population dynamics. (b) Large maturation rate and mortality rate of inside workers induce colony death. (c) Small enough egg cannibalism rate benefits adult worker ant’s growth and (or) development, large proportion of ants performing a given task can promote colony survival, and too large egg cannibalism rate can lead to colony’s death. (d) Increasing energy invested on brood care and (or) the conversion rate between fungus and ants could induce oscillatory dynamics in models with cannibalism.