Incipient Colonies Research Articles

How inquilinism shaped breeding systems in a termite host-inquiline relationship.

Social insects have developed a broad diversity of nesting and foraging strategies. One of these, inquilinism, occurs when one species (the inquiline) inhabits the nest built and occupied by another species (the host). Obligatory inquilines must overcome strong constraints upon colony foundation and development, due to limited availability of host colonies. To reveal how inquilinism shapes reproductive strategies in a termite host-inquiline dyad, we carried out a microsatellite marker study on Inquilinitermes inquilinus and its host Constrictotermes cavifrons. The proportion of simple, extended and mixed families was recorded in both species, as well as the presence of neotenics, parthenogenesis and multiple foundations. Most host colonies (95%) were simple families and all were monodomous. By contrast, the inquiline showed a higher proportion of extended (30%) and mixed (5%) families, and frequent neotenics (in 25% of the nests). This results from the simultaneous foundation in host nests of numerous incipient colonies, which, as they grow, may compete, fight, or merge. We also documented the use of parthenogenesis by female-female pairs. In conclusion, the classical monogamous colony pattern of the host species suggests uneventful development of simple foundations dispersed in the environment, in accordance with the wide distribution of their resources. By contrast, the multiple reproductive patterns displayed by the inquiline species reveal strong constraints on foundation sites: founders first concentrate into host nests, then must attempt to outcompete or absorb the neighbouring foundations to gain full control of the resources provided by the host nest.

Shaping the tripartite symbiosis: are termite microbiome functions directed by the environmentally acquired fungal cultivar?

Microbiome assembly critically impacts the ability of hosts to access beneficial symbiont functions. Fungus-farming termites have co-evolved with a fungal cultivar as a primary food source and complex gut microbiomes, which collectively perform complementary degradation of plant biomass. A large subset of the bacterial community residing within termite guts are inherited (vertically transmitted) from parental colonies, while the fungal symbiont is, in most termite species, acquired from the environment (horizontally transmitted). It has remained unknown how the gut microbiota sustains incipient colonies prior to the acquisition of the fungal cultivar, and how, if at all, bacterial contributions are modulated by fungus garden establishment. Here, we test the latter by determining the composition and predicted functions of the gut microbiome using metabarcoding and shotgun metagenomics, respectively. We focus our functional predictions on bacterial carbohydrate-active enzyme and nitrogen cycling genes and verify compositional patterns of the former through enzyme activity assays. Our findings reveal that the vast majority of microbial functions are encoded in the inherited microbiome, and that the establishment of fungal gardens incurs only minor modulations of predicted bacterial capacities for carbohydrate and nitrogen metabolism. While we cannot rule out that other symbiont functions are gained post-fungus garden establishment, our findings suggest that fungus-farming termite hosts are equipped with a near-complete set of gut microbiome functions at the earliest stages of colony life. This inherited, incipient bacterial microbiome likely contributes to the high extent of functional specificity and coevolution observed between termite hosts, gut microbiomes, and the fungal cultivar.

Innate and learned components of egg recognition in the ant Camponotus floridanus.

Insect societies discriminate against foreigners to avoid exploitation. In ants, helper workers only accept individuals with the familiar chemical cues of their colony. Similarly, unfamiliar eggs may get rejected at their first appearance in the nest. We investigated egg acceptance mechanisms by introducing different types of foreign eggs into worker groups of the ant Camponotus floridanus. Workers from established colonies familiar with queen-laid eggs always accepted eggs from highly fecund queens, but worker-laid eggs only after exposure for several weeks. Workers naive to eggs only rejected worker-laid eggs once they had prior exposure to eggs laid by highly fecund queens, suggesting that prior exposure to such eggs is necessary for discrimination. The general acceptance of eggs from highly fecund queens, irrespective of previous worker egg exposure, suggests an innate response to the queen pheromone these eggs carry. Workers learned to accept queen-laid eggs from different species, indicating high flexibility in learning egg-recognition cues. In incipient colonies with queen-laid eggs that carry a weak queen pheromone, worker-laid eggs were more likely to get accepted than queen-laid eggs from a different species, suggesting that the similarity of egg-recognition cues between the two types of C. floridanus eggs increases acceptance.

Inter-clonal competition over queen succession imposes a cost of parthenogenesis on termite colonies.

In social insect colonies, selfish behaviour due to intracolonial conflict among members can result in colony-level costs despite close relatedness. In certain termite species, queens use asexual reproduction for within-colony queen succession but rely on sexual reproduction for worker and alate production, resulting in multiple half-clones of a single primary queen competing for personal reproduction. Our study demonstrates that competition over asexual queen succession among different clone types leads to the overproduction of parthenogenetic offspring, resulting in the production of dysfunctional parthenogenetic alates. By genotyping the queens of 23 field colonies of Reticulitermes speratus, we found that clone variation in the queen population reduces as colonies develop. Field sampling of alates and primary reproductives of incipient colonies showed that overproduced parthenogenetic offspring develop into alates that have significantly smaller body sizes and much lower survivorship than sexually produced alates. Our results indicate that while the production of earlier and more parthenogenetic eggs is advantageous for winning the competition for personal reproduction, it comes at a great cost to the colony. Thus, this study highlights the evolutionary interplay between individual-level and colony-level selection on parthenogenesis by queens.

Hybrids of two destructive subterranean termites established in the field, revealing a potential for gene flow between species.

Hybridization between invasive pest species may lead to significant genetic and economic impacts that require close monitoring. The two most invasive and destructive termite species worldwide, Coptotermes formosanus Shiraki and Coptotermes gestroi (Wasmann), have the potential for hybridization in the field. A three-year field survey conducted during the dispersal flight season of Coptotermes in Taiwan identified alates with atypical morphology, which were confirmed as hybrids of the two Coptotermes species using microsatellite and mitochondrial analyses. Out of 27,601 alates collected over three years, 4.4% were confirmed as hybrid alates, and some advanced hybrids (>F1 generations) were identified. The hybrid alates had a dispersal flight season that overlapped with the two parental species 13 out of 15 times. Most of the hybrid alates were females, implying that mating opportunities beyond F1 may primarily be possible through female hybrids. However, the incipient colony growth results from all potential mating combinations suggest that only backcross colonies with hybrid males could sometimes lead to brood development. The observed asymmetrical viability and fertility of hybrid alates may critically reduce the probability of advanced-hybrid colonies being established in the field.

A comparison of tunnel geometry between the Formosan subterranean termite and the Asian subterranean termite (Blattodea: Rhinotermitidae).

Stake surveys and in-ground monitoring stations have been widely used to study field populations of many subterranean termite species, but thus far they have never been intercepted by the invasive Asian subterranean termite, Coptotermes gestroi (Wasmann), in southeastern Florida. To investigate the reasons for the inability of C. gestroi to intercept these in-ground monitoring devices, we compared its tunnel geometry with that of the Formosan subterranean termite, Coptotermes formosanus Shiraki. Two-year-old incipient colonies of both species confined in simulated structural infestations were connected to planar arenas containing four wooden discs. Coptotermes formosanus colonies constructed more abundant and more complex tunnel networks and intercepted more wooden discs than C. gestroi. C. formosanus propagated shorter primary tunnels and longer secondary tunnels with more branching frequency than C. gestroi, and probably adopted an area searching strategy to search for food in an area before moving on to other areas. In comparison, C. gestroi used a distance searching strategy by constructing linear and long primary tunnels to search for food at distance. Because tunnels of C. gestroi were less abundant and they tend to travel straight for some distance, they may have bypassed survey stakes or in-ground monitoring stations that are sparsely distributed in soil. The tunnel geometry of C. gestroi may explain why none of these in-ground monitoring devices has been intercepted by this species in southeastern Florida. © 2023 Society of Chemical Industry.

Us, them, and the others: Testing for discrimination amongst outgroups in a single-piece nesting termite, Zootermopsis angusticollis.

Recognition of group members is an important adaptation in social organisms because it allows help to be directed toward kin or individuals that are likely to reciprocate, and harm to be directed toward members of competing groups. Evidence in a wide range of animals shows that responses to outgroups vary with context, suggesting that cues to group membership also depend on the social or environmental context. In termites, intergroup encounters are frequent and their outcomes highly variable, ranging from destruction of a colony to colony fusion. As well as genetic factors, nestmate recognition in social insects commonly relies on cues that are mediated by environmental factors such as food source. However, single-piece nesting termite colonies share nesting material and food source with rival colonies (their wood substrate serves as both). In principle, the shared environment of single-piece nesting termite colonies could constrain their ability to identify non-nestmates, contributing to some of the variation seen in encounters, but this has not been investigated. In this study, we raised incipient colonies of a single-piece nesting termite, Zootermopsis angusticollis, on two different wood types and conducted behavioral assays to test whether nestmate discrimination can be constrained by common environmental conditions. We found that non-nestmates elicited higher rates of identity checking and defense behavior compared to nestmates, but there was no effect of wood type on the strength of behavioral responses to non-nestmates. We also found that one key cooperative behavior (allogrooming) was performed equally toward both nestmates and non-nestmates. These findings offer no support for the hypothesis that common wood type constrains the nestmate recognition system of single piece nesting termites. We suggest that where groups encounter each other frequently in a common environment, selection will favor discrimination based on genetic and/or higher resolution environmentally mediated cues.

Exuviae Recycling Can Enhance Queen Oviposition and Colony Growth in Subterranean Termites (Blattodea: Rhinotermitidae: Coptotermes).

Wood-feeding termites have a nitrogen-poor diet and have therefore evolved nitrogen conservation strategies. However, termite workers molt periodically, and throughout the lifetime of a colony, millions of exuviae, a nitrogen-rich resource, are produced by the colony. In Coptotermes Wasmann, workers foraging at remote feeding sites must return to the central part of the nest to molt, where the queen, king, eggs, and larvae are located. It was hypothesized that this molting-site fidelity is an efficient way to recycle nitrogen for reproduction and colony growth, as nestmates involved in exuviae consumption can directly transfer such resources to individuals engaged in reproduction (the queen) or growth (larvae). This study investigates whether incipient colonies of C. gestroi (Wasmann) can gain additional biomass when they are fed supplementary exuviae. Incipient colonies were reared in nitrogen-poor or nitrogen-rich conditions, and 0, 1, 5, or 10 exuviae were added to 3-month-old colonies. After 6.5 months, colonies reared in nitrogen-poor environments gained significantly more biomass when exuviae were added than colonies with no added exuviae. However, the addition of exuviae had no effect on colony growth for colonies reared in nitrogen-rich environments. In a second experiment, queens from colonies in which exuviae were effectively removed laid fewer eggs than queens from colonies in which exuviae were not removed. Therefore, consumption of exuviae from molting individuals by nestmates is an important part of the nitrogen recycling strategy in Coptotermes colonies, as it facilitates queen oviposition and colony growth, especially when such colonies have limited access to nitrogen-rich soils.

Post-embryonic development of the female reproductive system of the Neotropical termite Silvestritermes euamignathus (Isoptera: Termitidae: Syntermitinae)

In termites, the maturation of the female reproductive system is progressive following post-embryonic development. In Silvestritermes euamignathus, the imaginal line is characterized by five nymphal instars that develop into the imago, but it can deviate in some instars to neotenic reproductives. In order to understand the rate of development of the female reproductive system throughout post-embryonic development, we analyzed the morphology of the reproductive system of nymphs, characterizing the stage of development and comparing it with neotenics and primary queens. We also followed embryonic development and compared eggs from neotenics and primary queens from incipient and mature colonies. Our results showed that gonadal development follows the post-embryonic instars, and previtellogenic oocytes are present in third-instar nymphs and are retained for the next two successive nymphal instars. The full maturation of the ovaries with vitellogenic oocytes requires molting to either imagos or neotenics. Eggs from neotenics follow the same embryonic development and each stage presents similar volume when compared to those of mature primary queens. Eggs of primary queens from incipient colonies are greater in volume than those of mature primary queens and of neotenics, suggesting an investment in egg quality rather than quantity during colony foundation.

Eusociality and the transition from biparental to alloparental care in termites

AbstractIn eusocial organisms, cooperative brood care within a colony represents a situation where the ancestral parental care duties have shifted away from the reproductive parent(s) towards their offspring. The shift to alloparental care was often instrumental in the initial emergence of eusociality, as it ultimately contributed to the establishment of the reproductive division of labour.Remarkably, eusocial taxa such as ants and termites, which still display an ancestral independent colony foundation phase, must go through an obligatory parental care period, as a temporary subsocial family unit. In termites specifically, an incipient colony inherently remains a woodroach family unit until alloparental care is established. Colony foundation success can then be limited by a series of factors that may include environmental, behavioural, symbiotic and physiological constraints.In this study, 450 incipient termite colonies (Coptotermes gestroi) were established to investigate the timing of physiological changes in founders during the transition from biparental to alloparental care. Results showed that the finite initial internal nutritional resources that alates carry during the dispersal flight are a primary limiting factor for successful colony establishment. TheCoptotermesqueen and king must rapidly establish (<150 days) their first cohort of offspring to reach alloparental care or simply run out of resources and die. Alates, therefore, carry just enough internal resources to produce the first few alloparents (< 15 workers) to prime the system towards colony ergonomic growth, with a definitive shift to solely reproductive functions.Eusocial insect primary reproductive traits were optimized for three successive functions within the life cycle of a colony: alate dispersal (sexual reproduction), colony foundation (parental care) and colony growth (increased egg production towards colony maturity). However, results suggest that trade‐offs involving these functions appear to primarily favour dispersal ones (quantity vs. quality of alates), as founder(s) carry minimal resources and have no room for parental care inefficiency and as they then fully rely on their alloparents for further reproductive output.The transition towards alloparental care during colony foundation of eusocial insects may, therefore, reflect on the initial evolutionary transition from ancestral subsociality to eusociality.Read the freePlain Language Summaryfor this article on the Journal blog.

Termite graveyards. Hidden geochemical patches?

Entombment, or the production of graveyards for the disposal of dead bodies, is not only a practice of human societies but is also observed in nature, including among small invertebrates such as termites. While the influence of termites on soil dynamics has largely been studied in comparing the specific properties of their mounds and protective sheeting with those of the surrounding soil, the properties of their graveyards have never been described before. Using incipient colonies of Macrotermes natalensis reared in a controlled environment, we showed that graveyard sheeting was characterized by a much higher C content in comparison with the reference soil and protective sheeting (4.7-fold increase). As a consequence, a slight increase in the C:N ratio was measured from 8 in the reference soil to 10 in graveyard sheeting. No changes in soil particle size fractions were measured. However, lower Fe and Al contents were measured in sheeting, and micrographs obtained from scanning electron microscopy revealed the presence of calcium carbonate, or calcium oxalate crystals, in sheeting, as well as the presence of organic substances and salt crystals covering termite corpses, most likely for controlling the spread of pathogens. The presence of calcium carbonates and/or calcium oxalate was explained by the very high Ca content within termite bodies. Therefore, this study shows that termite graveyards are likely to constitute unexplored patches of nutrients in soil.

Short and long-term costs of inbreeding in the lifelong-partnership in a termite

Social life and lifelong partner commitments are expected to favor thorough partner choice, as an ill-suited partnership may have long-term consequences, adversely affecting the parents and spanning several cohorts of offspring. Here, we used ~1400 termite incipient colonies to estimate the short- and long-term costs of inbreeding upon the survival of the parents over a 15-month period, their productivity, and the resistance of their offspring toward pathogen pressure. We observed that foundation success was not influenced by the relatedness of partners, but by their levels of microbial load. We showed faster growth in inbred colonies with low levels of microbial load, revealing a potential tradeoff between pathogen defense and offspring production. Yet, inbreeding takes its toll later in colony development when offspring from incipient colonies face pathogen pressure. Although the success of a lifetime partnership is initially determined by the partner’s health, the cost of inbreeding in incipient colonies favors outbred colonies reaching maturity.

Worker task organization in incipient bumble bee nests

Bumble bees (Bombus: Apidae) are a long-standing model system for understanding animal behaviour, ecology and evolution. However, how workers in this system are organized to perform fundamental tasks related to brood feeding and food collection remains unclear. Bumble bees undergo dramatically different life stages, across which the social environment, and therefore task organization, changes over time. Queens initiate nests solitarily, and when the first cohort of workers emerge, they help the queen carry out brood-feeding and food collection tasks, whereas the queen transitions to primarily egg laying. Although task organization has been studied in mature colonies, few studies to date have explored how these tasks are organized in young, incipient nests. Here, we explored how food-related tasks, including brood feeding and food collection, are organized by workers in incipient colonies. We found that food-related tasks were nested, in that the majority of workers fed brood, a subset also collected nectar and a smaller subset also collected pollen. These patterns suggest that brood feeding is a task shared by most workers, and that the distinction between pollen collecting versus nonpollen collecting might be the most important axis of division of labour in bumble bee nests, at least at the early nesting stage.

The lose-to-win strategy of the weak: intraspecific parasitism via egg abduction in a termite.

Predation by larger conspecifics poses a major threat to small juveniles in many animal species. However, in social insects, raids perpetrated by large colonies may provide smaller colonies with opportunities for parasitization. Herein, in the termite Reticulitermes speratus, we demonstrate that small incipient colonies parasitize large mature colonies through egg abduction when attacked by raiding conspecifics. We observed that the eggs of incipient colonies were brought into raiding colonies while their parents were killed during the attack. In this species, unmated females found new colonies with female-female (FF) cooperation, in addition to the typical monogamous colony foundation. Interestingly, the abducted eggs of FF pairs developed into nymphs (reproductive caste) in the raiding colonies, whereas the eggs of male-female (MF) pairs developed into workers (non-reproductive caste). Parthenogenetic eggs are known to be developmentally predisposed to becoming female reproductives owing to genomic imprinting in termites. This study demonstrates that the plundering of small colonies by larger conspecific colonies not only results in the extinction of the weaker colonies, but also serves as a strategy that incipient colonies use to obtain the reproductive position in large colonies by stealth. The results elucidate the diversity and complexity of inter-colonial interactions in social insects.

Queen Egg Laying and Egg Hatching Abilities are Hindered in Subterranean Termite Colonies When Exposed to a Chitin Synthesis Inhibitor Bait Formulation.

Subterranean termite control methods using chitin synthesis inhibitors (CSIs) aim at eliminating colonies that feed upon a bait formulation. Several benzoylurea active ingredient formulations are currently commercially available as alternative termite management strategies to liquid termiticides. Individual workers need to molt on a regular basis and CSIs interfere with such molting process, allowing sufficient time for the acquisition of a colony-wide lethal dose prior to widespread mortality. As workers progressively die, the colony eventually collapses, leaving only soldiers and primary reproductives that starve to death. One common observation is that young workers often die early owing to their relatively short molting cycle. However, the absence of brood in dying colonies raises questions about the potential fate of eggs laid by the queen. This study aims to determine if CSI baits also terminate the ability of a colony to produce a new cohort of workers by disabling the ongoing brood development. Incipient termite colonies were used to test the impact of noviflumuron on the queen's ability to lay eggs and on the eggs' ability to hatch. Our results showed that queens in colonies exposed to CSI not only initially laid less eggs than the control queens, but eggs also did not develop and were progressively cannibalized, eventually leading to colony establishment failure. This result implies that queens of mature colonies exposed to CSI would lose the ability to lay viable eggs as the colony collapses, leading to an absence of worker replacement, aiding in colony elimination.

The early life of a leaf-cutter ant colony constrains symbiont vertical transmission and favors horizontal transmission.

Colonial organisms host a large diversity of symbionts (collectively, parasites, mutualists, and commensals) that use vertical transmission (from parent colony to offspring colony) and/or horizontal transmission to disperse between host colonies. The early life of some colonies, characterized by the dispersal and establishment of solitary individuals, may constrain vertical transmission and favor horizontal transmission between large established colonies. We explore this possibility with the miniature cockroach Attaphila fungicola, a symbiont of leaf‐cutter ants and the mutualist fungal gardens they cultivate. The early life of a leaf‐cutter colony is characterized by the dispersal of a female alate (winged “queen”) carrying a fungal pellet, and the subsequent establishment of a foundress (workerless “queen”) raising her incipient fungal garden and colony. Roaches hitchhike on female alates during leaf‐cutter nuptial flights, which strongly suggests that roaches are vertically transmitted to foundresses and their incipient colonies; however, weak compatibility between roaches and incipient gardens may constrain roach vertical transmission. Reciprocally, opportunities for horizontal transmission between large established colonies with abundant fungal gardens may weaken selection against roach‐induced harm (virulence) of incipient gardens. We use a laboratory experiment, behavioral observations, field surveys, and a transmission model to estimate the effect roaches have on the survivorship of incipient gardens and the frequency of roach vertical transmission. Contrary to traditional assumptions, our results indicate that roaches harm incipient gardens and predominantly use horizontal transmission between established leaf‐cutter colonies. Ultimately, “costs of generalism” associated with infecting disparate stages of a host's lifecycle (e.g., incipient vs. established colonies) may constrain the vertical transmission of roaches and a broad range of symbionts.

Care-giver identity impacts offspring development and performance in an annually social bumble bee

BackgroundThe developmental fates of offspring have the potential to be influenced by the identity of their care-givers and by the nature of the care that they receive. In animals that exhibit both parental and alloparental care, such as the annually eusocial insects, the influence of care-giver identity can be directly assessed to yield mechanistic and evolutionary insights into the origins and elaboration of brood care. Here, we performed a comparative investigation of maternal and worker brood care in bumble bees, a pollinator group where mothers (queens) rear the first offspring in the nest, and then daughters (workers) assume this role upon their emergence. Specifically, we compared the effects of queen and worker brood care on offspring development and also offspring performance, for a set of traits related to sensory biology, learning, and stress resistance.ResultsWe found that queen-reared workers were smaller-bodied than worker-reared offspring, suggesting that bumble bee queens influence body size determination in their offspring. We also found that queen-reared workers were more resistant to starvation, which might be beneficial for early nesting success. These maternal influences could not be explained by feeding rate, given that we detected a similar offspring feeding frequency in both queens and workers.ConclusionBumble bee queens have a unique influence on the development of the first offspring in the nest, which they rear, relative to worker-reared workers. We propose that bumble bee brood care has been shaped by a suite of evolutionary and ecological factors, which might include a maternal influence on traits that promote survival of incipient colonies.

Dynamic protozoan abundance of Coptotermes kings and queens during the transition from biparental to alloparental care

An incipient termite colony initially functions as a biparental family unit. The emergence of the first workers initiates the transition from biparental to alloparental care within a colony. During this transition, the number of protozoa harbored by Reticulitermes speratus (Kolbe) kings and queens is dynamic. In Coptotermes gestroi (Wasmann), the transition to alloparental care is completed by 150 days after colony foundation. In the present study, we quantified the protozoan abundance of C. gestroi and Coptotermes formosanus Shiraki kings and queens at ten time points during the transition from biparental to alloparental care (0–400 days post colony foundation). The protozoan abundance of C. gestroi and C. formosanus kings and queens peaked on either day 35 or 60 during the biparental care period, and then progressively decreased during the remainder of the study. By day 400 the protozoan abundance of kings and queens was either similar to or less than the abundance observed within unmated dealates. Both C. gestroi and C. formosanus kings and queens still harbored protozoa at day 150 even though the transition to alloparental care was completed by this time. On days 250 and 400 for C. formosanus, and on day 400 for C. gestroi, the protozoan abundance of kings was greater than the abundance of queens. These results indicate that alloparental care can become set within a colony prior to the loss of protozoa within the royal pair, and that queens lose their protozoa earlier than kings during incipient colony development.

Age-related division of labor occurs in ants at the earliest stages of colony initiation

Mature colonies of many social insect species exhibit division of labor (DOL) where individual workers specialize in doing only a subset of the multiple tasks needed to maintain homeostasis. In newly initiated ant colonies, however, the first workers (called nanitics) are few in number and much smaller in size than those in mature colonies. This limited workforce must perform most of the tasks of mature colonies, but it is unknown if they also exhibit DOL. In this study, we tracked several inside-nest and outside-nest behaviors of nanitics in incipient Pogonomyrmex rugosus colonies. DOL arises in these colonies, whereby the relatively oldest workers (even if only by a few hours) are biased towards foraging, while younger nanitics concentrate on brood care. The addition of new nanitics shifts behavior in the oldest individual away from brood care but does not immediately increases its foraging. Conversely, nanitics left alone due to mortality of nest-mates forage more, but do not reduce brood care. The results suggest P. rugosus nanitics follow an age-related task specialization pattern that is broadly similar to mature colonies. The nanitic life stage may be, however, unique for ants in how fine-grained the DOL is in terms of absolute age differences and flexibility for task switching. The ecological success of ants is thought to be facilitated by workers dividing their labor across tasks. Commonplace in mature colonies is that individuals will perform safer, within-nest tasks such as brood care when young, and shift as they age towards riskier, outside-nest tasks such as foraging. Because many do the same task, the death of any single worker has minimal effect. We show, for the first time, a similar age-related, task specialization pattern also occurs in newly founded colonies. In this earliest life, history stage workers are small and few in number, precluding the massive task redundancy that characterizes mature colonies. Nevertheless, a division of labor correlating with relative age differences arises even when workers differ in age by only a few hours. This supports the hypothesis that a similar developmental pathway for task allocation in worker behavior occurs at all stages in colony life history.

Reproductive Soldier Development Is Controlled by Direct Physical Interactions with Reproductive and Soldier Termites.

Simple SummaryGenerally, in termites, the soldier caste is sterile, and their reproductive organ formation is suppressed. However, in some primitive taxa, neotenic reproductives with soldier-like mandibles, called reproductive soldiers, occasionally appear in the incipient colony. Our first step in understanding the developmental mechanism of this unique caste was to establish efficient conditions to induce reproductive soldiers in Zootermopsis nevadensis. Removing both the king and soldiers from an incipient colony strongly stimulated the formation of male reproductive soldiers, which possessed soldier-like mandibles and developed testes. Similarly, high differentiation rates of male reproductive soldiers were observed after king-and-soldier separation treatment using wire mesh. However, no male reproductive soldiers were produced without direct interaction with the queen. These results suggest that reproductive soldier development might be initiated and controlled by direct physical interactions with both reproductives (the king and queen) and soldiers.In eusocial insects (e.g., ants, bees, and termites), the roles of different castes are assigned to different individuals. These castes possess unique phenotypes that are specialized for specific tasks. The acquisition of sterile individuals with specific roles is considered a requirement for social evolution. In termites, the soldier is a sterile caste. In primitive taxa (family Archotermopsidae and Stolotermitidae), however, secondary reproductives (neotenic reproductives) with their mandibles developed into weapons (so-called reproductive soldiers, also termed as soldier-headed reproductives or soldier neotenics) have been reported. To understand the developmental mechanism of this unique caste, it is necessary to understand the environmental cues and developmental processes of reproductive soldiers under natural conditions. Here, we established efficient conditions to induce reproductive soldiers in Zootermopsis nevadensis. Male reproductive soldiers frequently developed after the removal of both the king and soldiers from an incipient colony. Similarly, high differentiation rates of male reproductive soldiers were observed after king-and-soldier separation treatment using wire mesh. However, no male reproductive soldiers were produced without direct interaction with the queen. These results suggest that male reproductive soldier development is repressed by direct physical interactions with both the king and soldiers and facilitated by direct physical interaction with the queen.