Groups Of Social Insects Research Articles

Nest-based information transfer and foraging activation in the common wasp (Vespula vulgaris)

The apparent absence of intra-nest signals and communication about food resources (recruitment) among social wasps does not rule out the possibility of information transfer and coordinated foraging among nestmates. In the present study, we tested the hypothesis that the common wasp (Vespula vulgaris) shows nest-based information transfer and foraging activation: an increase in the probability of an individual leaving the nest as a result of information about resources received from successful foragers. We controlled for the possibility of local enhancement, chemical trails at the food source and climatic variation. We found evidence that food choice and discovery of resources in the field by naive foragers were assisted by information previously or simultaneously provided by experienced nestmates. This information was related to chemical cues associated with the food and possibly to its location. Our observations suggest piloting between common wasp foragers. At the trained nest, there was a change in foraging effort at the colony level when known resources were available. Reactivated, experienced foragers were the main group responsible for the increase in foraging traffic rate observed at the colony level. To our knowledge, this is the first study clearly demonstrating nest-based information transfer about food resources in V. vulgaris and one of the few providing evidence of foraging activation in social wasps. Our data are consistent with the possibility of recruitment in this group of social insects.

Social interactions affecting caste development through physiological actions in termites.

A colony of social insects is not only an aggregation of individuals but also a functional unit. To achieve adaptive social behavior in fluctuating environmental conditions, in addition to coordination of physiological status in each individual, the whole colony is coordinated by interactions among colony members. The study on the regulation of social-insect colonies is termed “social physiology.” Termites, a major group of social insects, exhibit many interesting phenomena related to social physiology, such as mechanisms of caste regulation in a colony. In their colonies, there are different types of individuals, i.e., castes, which show distinctive phenotypes specialized in specific colony tasks. Termite castes comprise reproductives, soldiers and workers, and the caste composition can be altered depending on circumstances. For the regulation of caste compositions, interactions among individuals, i.e., social interactions, are thought to be important. In this article, we review previous studies on the adaptive meanings and those on the proximate mechanisms of the caste regulation in termites, and try to understand those comprehensively in terms of social physiology. Firstly, we summarize classical studies on the social interactions. Secondly, previous studies on the pheromone substances that mediate the caste regulatory mechanisms are overviewed. Then, we discuss the roles of a physiological factor, juvenile hormone (JH) in the regulation of caste differentiation. Finally, we introduce the achievements of molecular studies on the animal sociality (i.e., sociogenomics) in terms of social physiology. By comparing the proximate mechanisms of social physiology in termites with those in hymenopterans, we try to get insights into the general principles of social physiology in social animals.

Geographical patterns of genetic divergence in the widespread Mesoamerican bumble bee Bombus ephippiatus (Hymenoptera: Apidae)

Bumble bees (Bombus Latreille) are an important group of social insects, well recognized throughout northern temperate regions as important pollinators of wild and agricultural plants. Little is known about the biology of this group in southern portions of the Americas, especially in Mesoamerica, a region of geological and ecological complexity from Mexico through Central America. One ubiquitous Mesoamerican species, Bombus ephippiatus, is enigmatic. Like many other Bombus, this species is homogeneous in body structure yet exhibits striking intraspecific color pattern polymorphism across its range, leading to uncertainty about its genealogical boundaries. It has been grouped taxonomically with B. wilmattae, a species narrowly restricted to southern Mexico and northern Guatamala. Furthermore, the relationships between these two taxa and a third species, B. impatiens, found only in America north of Mexico, have been controversial. Our phylogenetic analysis of DNA sequences from mitochondrial COI and nuclear PEPCK and CAD resolves the phylogeny of these three taxa as (B. impatiens, (B. ephippiatus, B. wilmattae)). Additional data from eight nuclear microsatellite markers reveal complex patterns of genetic divergence and isolation among populations of B. ephippiatus across its extensive geographic range, providing evidence for multiple independent evolutionary lineages. These lineages correspond not only to geographic and habitat variation across their range, but also to distinct color pattern groups present in the species. Knowledge of the phylogeny and genetic divergence of the B. ephippiatus group will provide a framework for understanding evolutionary and ecological origins of color pattern polymorphism in bumble bees, as well as providing insight into geographical factors enhancing speciation in Mesoamerica.

Epigenetics in Social Insects: A New Direction for Understanding the Evolution of Castes

Epigenetic modifications to DNA, such as DNA methylation, can expand a genome's regulatory flexibility, and thus may contribute to the evolution of phenotypic plasticity. Recent work has demonstrated the importance of DNA methylation in alternative queen and worker “castes” in social insects, particularly honeybees. Social insects are an excellent system for addressing questions about epigenetics and evolution because: (1) they have dramatic caste polyphenisms that appear to be tied to differential methylation, (2) DNA methylation is widespread in various groups of social insects, and (3) there are intriguing connections between the social environment and DNA methylation in many species, from insects to mammals. In this article, we review research on honeybees, and, when available, other social insects, on DNA methylation and queen and worker caste differences. We outline a conceptual framework for the effects of methylation on caste determination in honeybees that may help guide studies of epigenetic regulation in other polyphenic taxa. Finally, we suggest future paths of study for social insect epigenetic research, including the importance of comparative studies of DNA methylation on a broader range of species, and highlight some key unanswered mechanistic questions about how DNA methylation affects gene regulation.

Descriptive attributes used in the characterization of stingless bees (Apidae: Meliponini) in rural populations of the Atlantic forest (Misiones-Argentina)

BackgroundHuman beings employ a combination of morphological, sensorial, utilitarian, cultural and ecological characters when they identify and classify organisms. Ethnotaxonomy has provided a store of information about the characters cultures employ when they identify and classify a vast diversity of taxonomic groups. Nevertheless, some more research is needed to provide a comparison of the characters employed in the description of taxons, and an analysis of the extent to which those descriptors are represented. Stingless bees constitute a diverse group of social insects that have been widely studied from an ethnobiological perspective due to their utilitarian and cultural importance. The objective of this study is to identify the elements local people consider when characterizing stingless bees, and how important these elements are in the study of local classifications.MethodsThe methodology used involves semi-structured interviews and trips with the informants to rural areas. Locally known ethnospecies are characterized, descriptive traits and salient criteria used in those characterizations are identified, and the frequency of reference of descriptive traits and salient criteria are estimated. Besides, the descriptive traits used for each ethnospecies are compared, and the contribution of the characterizations as a heuristic strategy in the study of folk classification systems is analyzed.ResultsThe use of 19 biological descriptors (grouped according to 4 salient criteria) and of comparisons among ethnospecies was found. Results suggest the existence of group and specific descriptors. Researchers identified which ethnospecies are considered similar, how less important traits contribute to descriptions, the relation between specific descriptors and ethnospecies, the presence of cognitive prototypes, and the most relevant salient properties from the emic perspective.ConclusionsThe estimated importance of attributes descriptors allowed us to identify the spectrum of salient properties relevant from the emic perspective to characterize the stingless bees. In this sense, the analysis proposed here is useful to study folk taxonomy in culturally heterogeneous groups or multicultural regions, where the linguistic elements usually employed cannot be applied.ResumenAntecedentesLos seres humanos, al identificar y clasificar a los organismos emplean una combinación de características morfológicas, sensoriales, utilitarias, culturales y ecológicas. Entre los aportes generados desde la etnotaxonomía, se ha obtenido información sobre los caracteres utilizados para identificar y clasificar una gran diversidad de grupos taxonómicos. Sin embargo, aún faltan trabajos donde se comparen los caracteres utilizados en las descripciones de taxones y se analice en qué medida estos descriptores se encuentran representados. Las abejas sin aguijón conforman un diverso grupo de insectos sociales que han sido estudiadas desde la perspectiva etnobiológica, dada su importancia utilitaria y cultural. Los interrogantes que guían este trabajo son ¿Qué elementos tienen en cuenta los pobladores para caracterizar a las abejas sin aguijón? y ¿Qué importancia revisten los mismos en el estudio de las clasificaciones locales?MétodosSe realizaron entrevistas semiestructuradas y recorridos en áreas rurales con los informantes. Se caracterizan a las etnoespecies conocidas localmente; se identifican los atributos descriptores y los criterios emergentes utilizados para dichas caracterizaciones; y se estima la frecuencia de citas de los atributos descriptores y criterios emergentes. Por otra parte, se comparan los atributos descriptores empleados para cada etnoespecie y se analiza el aporte de las caracterizaciones como estrategia heurística en el estudio de los sistemas de clasificación folclóricos.ResultadosSe halló el empleo de 19 descriptores biológicos (que fueron agrupados en 4 criterios emergentes) y de comparaciones entre etnoespecies. Los resultados sugieren la existencia de descriptores de grupo y descriptores específicos. Se identificaron cuales etnoespecies son consideradas similares, cómo contribuyen a las descripciones los atributos de menor peso, la relación entre los descriptores específicos y las etnoespecies, la presencia de prototipos cognitivos, así como las propiedades emergentes más relevantes desde la perspectiva émica.ConclusionesLa importancia estimada de los atributos descriptores nos ha permitido identificar el espectro de propiedades emergentes que son relevantes desde la perspectiva emic para caracterizar las abejas sin aguijón. En este sentido el análisis aquí propuesto es de utilidad para estudiar taxonomías folclóricas en grupos heterogéneos culturalmente o en regiones pluriculturales, donde los elementos lingüísticos usualmente empleados no son aplicables.

Nestmate recognition in the stingless bee Frieseomelitta varia (Hymenoptera, Apidae, Meliponini): sources of chemical signals

Social insects use cuticular lipids for nestmate recognition. These lipids are chiefly hydrocarbons that can be endogenously produced or acquired from the environment. Although these compounds are already described as coming from different sources for different groups of social insects, nothing is known about the source of cuticular hydrocarbons in stingless bees. We used behavioural recognition tests and cuticle chemical investigation to elucidate the role of endogenous and environmentally based cues for nestmate recognition in the stingless bee Frieseomelitta varia. We found that although newly emerged workers present specific cuticle patterns according to their nest origin, these compounds are not used for nestmate recognition, since newly emerged workers are broadly accepted in different colonies. The cerumen used in nest construction played an important role in recognition behaviour. Twenty minutes of contact with foreign cerumen was sufficient to increase the rejection rates of nestmates and separate the groups of workers according to their chemical profile. On the other hand, tests of feeding on a common diet showed no effect on chemical cuticle pattern or recognition behaviour.

Sexual dimorphism of arrestment and gregariousness in the bed bug (Cimex lectularius) in response to cuticular extracts from nymphal exuviae

Releaser pheromones have direct behavioural effects to arrest, attract or disperse insects, whereas interactions within groups of social insects are often influenced by primer pheromones. The behaviour of insects displaying intermediate levels of sociality is largely unexplored in this context. In the present study, both the gregariousness and arrestment (settling near the odour source) of bed bugs Cimex lectularius L. (Hemiptera: Cimicidae) in response to conspecific exuvial extracts are described. Adult males are arrested on filter papers with extracts derived from exuviae of fifth-instar nymphs. Adult females and nymphs display no significant evidence for such behaviour. Adults of both sexes show no preference for extracts of male versus female fifth-instar exuviae. Arrestment of adult males does not occur on papers treated with fourth-instar exuvial extracts. Because the insects are assayed behaviourally in groups, an index is calculated describing how far bugs are away from being located independently of one another, as a measure of gregariousness. Adult males have lower values for this index (i.e. locations are closer to independence). Adult females, nymph cohorts and mixed age groups all have higher values for this index, which tend to increase over time. Females exhibit a clear increasing dose-dependent relationship for this index. It is concluded that the extracts of fifth-instar nymphal exuvia arrest males on refuges that possess the odour source. However, gregariousness is induced in females, without evidence of a tendency to assemble near the odour source.

Gene expression changes during caste-specific neuronal development in the damp-wood termite Hodotermopsis sjostedti

BackgroundOne of the key characters of social insects is the division of labor, in which different tasks are allocated to various castes. In termites, one of the representative groups of social insects, morphological differences as well as behavioral differences can be recognized among castes. However, very little is known about the neuronal and molecular bases of caste differentiation and caste-specific behavior. In almost all termite species, soldiers play defensive roles in their colonies, and their morphology and behavior are largely different from workers (or pseudergates). Therefore, we predicted that some genes linked to defensive behavior and/or those required for neuronal changes are differentially expressed between workers and soldiers, or during the soldier differentiation, respectively.ResultsUsing the brain and suboesophageal ganglion (SOG) of the damp-wood termite Hodotermopsis sjostedti, we first screened genes specifically expressed in soldiers or during soldier differentiation by the differential display method, followed by quantitative real-time polymerase chain reaction. No distinctive differences in expression patterns were detected between pseudergates and soldiers. In the course of soldier differentiation, however, five genes were found to be up-regulated in brain and/or SOG: 14-3-3epsilon, fibrillin2, beta-tubulin, ciboulot, and a hypothetical protein containing a SAP motif. Some of these genes are thought to be associated with cytoskeletal structure or motor-associated proteins in neuronal tissues.ConclusionThe identified five genes could be involved in soldier-specific neuronal modifications, resulting in defensive behaviors in termite soldiers. The temporal expression patterns of these genes were consistent with the neuronal changes during soldier differentiation, suggesting that molecular machineries, in which the identified factors would participate, play important roles in behavioral differentiation of termite soldiers.

Scent of a queen—cuticular hydrocarbons specific for female reproductives in lower termites

In social insects, it is assumed that signals of the queen inform nestmates about her reproductive status. Thus, workers forego their own reproduction if the queen signals high fertility. In hemimetabolous termites, little is known about reproductive inhibition, but evidence exists for a royal-pair control. Workers of lower termites exhibit a high developmental flexibility and are potentially able to become reproductives, but the presence of a fertile reproductive restrains them from reaching sexual maturity. The nature of this control, however, remains unknown. Here, we report on qualitative differences in cuticular hydrocarbon profiles between queens and workers of the basal drywood termite Cryptotermes secundus. Queens were characterized by a shift to long-chained and branched hydrocarbons. Most remarkably, similar chemical patterns are regarded as fertility cues of reproductives in social Hymenoptera. This might suggest that both groups of social insects convergently evolved similar chemical signatures. The present study provides deeper insights into how termites might have socially exploited these signatures from sexual communication in their cockroach-like ancestor.

Life history and development ‐ a framework for understanding developmental plasticity in lower termites

Termites (Isoptera) are the phylogenetically oldest social insects, but in scientific research they have always stood in the shadow of the social Hymenoptera. Both groups of social insects evolved complex societies independently and hence, their different ancestry provided them with different life-history preadaptations for social evolution. Termites, the 'social cockroaches', have a hemimetabolous mode of development and both sexes are diploid, while the social Hymenoptera belong to the holometabolous insects and have a haplodiploid mode of sex determination. Despite this apparent disparity it is interesting to ask whether termites and social Hymenoptera share common principles in their individual and social ontogenies and how these are related to the evolution of their respective social life histories. Such a comparison has, however, been much hampered by the developmental complexity of the termite caste system, as well as by an idiosyncratic terminology, which makes it difficult for non-termitologists to access the literature. Here, we provide a conceptual guide to termite terminology based on the highly flexible caste system of the "lower termites". We summarise what is known about ultimate causes and underlying proximate mechanisms in the evolution and maintenance of termite sociality, and we try to embed the results and their discussion into general evolutionary theory and developmental biology. Finally, we speculate about fundamental factors that might have facilitated the unique evolution of complex societies in a diploid hemimetabolous insect taxon. This review also aims at a better integration of termites into general discussions on evolutionary and developmental biology, and it shows that the ecology of termites and their astounding phenotypic plasticity have a large yet still little explored potential to provide insights into elementary evo-devo questions.

Evolutionary ecology of social and sexual systems: crustaceans as model organisms

Understanding of animal social and sexual evolution has seen a renaissance in recent years with discoveries of frequent infidelity in apparently monogamous species, the importance of sperm competition, active female mate choice, and eusocial behavior in animals outside the traditional social insect groups. Each of these findings has raised new questions, and suggested new answers, about the evolution of behavioral interactions among animals. This volume synthesizes recent research on the sexual and social biology of the Crustacea, one of the dominant invertebrate groups on earth. Its staggering diversity includes ecologically important inhabitants of nearly every environment from deep-sea trenches, through headwater streams, to desert soils. The wide range of crustacean phenotypes and environments is accompanied by a comparable diversity of behavioral and social systems, including the elaborate courtship and wildly exaggerated morphologies of fiddler crabs, the mysterious queuing behavior of migrating spiny lobsters, and even eusociality in coral-reef shrimps. This diversity makes crustaceans particularly valuable for exploring the comparative evolution of sexual and social systems. Despite exciting recent advances, however, general recognition of the value of Crustacea as models has lagged behind that of the better studied insects and vertebrates. This book synthesizes the state of the field in crustacean behavior and sociobiology and places it in a conceptually based, comparative framework that will be valuable to active researchers and students in animal behavior, ecology, and evolutionary biology. It brings together a group of internationally recognized and rising experts in fields related to crustacean behavioral ecology, ranging from physiology and functional morphology, through mating and social behavior, to ecology and phylogeny. Each chapter makes connections to other, non-crustacean taxa, and the volume closes with a summary section that synthesizes the contributions, discusses anthropogenic impacts, highlights unanswered questions, and provides a vision for profitable future research.

Castes and asynchronous colony cycle in Polybia bistriata (Fabricius) (Hymenoptera: Vespidae)

Neotropical swarm-founding wasps, the Epiponini, are an outstanding group of social insects whose societies are polygynic and complex nest builders. Caste dimorphism in these wasps ranges from incipient to highly distinct. Morphometric analyses of nine body parts, ovarian status, relative age and development of the 5th gastral sternite gland (Richards' gland) of Polybia bistriata Fabricius were undertaken in order to estimate caste differentiation in nine colonies. Morphological and physiological data were used in multivariate analyses in order to evaluate the level of discrimination between inseminated and non-inseminated females. Clear physiological differences were found: queens had highly developed ovaries and they were inseminated, and workers had totally undeveloped ovaries or they had few developed oocytes (only in two colonies), but in both cases insemination was not detected. ANOVA and discriminant function analysis detected slight, but significant differences between castes. In relation to colony cycle, colonies were considered to be in the following stages: one in pre-emergence, four in worker production and four in sexuals production. Richards' gland analyses indicated that in small colonies (<100 females) queens had a less developed gland than in medium (100-200 females), and large colonies (>200 females). Taking the whole data, it was possible to conclude that caste differences were slight, but more evident in some phases of the colony cycle, a phenomenon previously described for other epiponines.

Social complexity in bees is not sufficient to explain lack of reversions to solitary living over long time scales

BackgroundThe major lineages of eusocial insects, the ants, termites, stingless bees, honeybees and vespid wasps, all have ancient origins (≥ 65 mya) with no reversions to solitary behaviour. This has prompted the notion of a 'point of no return' whereby the evolutionary elaboration and integration of behavioural, genetic and morphological traits over a very long period of time leads to a situation where reversion to solitary living is no longer an evolutionary option.ResultsWe show that in another group of social insects, the allodapine bees, there was a single origin of sociality > 40 mya. We also provide data on the biology of a key allodapine species, Halterapis nigrinervis, showing that it is truly social. H. nigrinervis was thought to be the only allodapine that was not social, and our findings therefore indicate that there have been no losses of sociality among extant allodapine clades. Allodapine colony sizes rarely exceed 10 females per nest and all females in virtually all species are capable of nesting and reproducing independently, so these bees clearly do not fit the 'point of no return' concept.ConclusionWe argue that allodapine sociality has been maintained by ecological constraints and the benefits of alloparental care, as opposed to behavioural, genetic or morphological constraints to independent living. Allodapine brood are highly vulnerable to predation because they are progressively reared in an open nest (not in sealed brood cells), which provides potentially large benefits for alloparental care and incentives for reproductives to tolerate potential alloparents. We argue that similar vulnerabilities may also help explain the lack of reversions to solitary living in other taxa with ancient social origins.

A test of adaptive hypotheses: Mandibular traits, nest construction materials, and feeding habits in neotropical social wasps (Vespidae, Polistinae)

The success of many behaviors in hymenoptera depends on the shape and structure of their mandibles. Neotropical social wasps exhibit variation in the shape of their mandibles; both the type of material used in the nest construction and the food sources have been proposed as selective forces that explain that variation. These hypotheses were studied using Independent Contrast analyses and combined tests of significance between: nine mandible’s traits, the type of nest material and food source types. Necrophagy and short fiber use are derived conditions in the subfamily that have evolved five times each, while the use of long fibers and live prey are primitive conditions. The mandible structures appear related to the nest fiber type but not to necrophagy. Particularly, species that use long fibers have bigger internal dorsal tooth, longer internal ventral tooth, and shorter and more curved mandibles. Similar results were obtained by including plant hairs as a third nest material type. These findings open related questions for groups of social insects where studies on evolution of mandibles have emphasized food sources.

Shape differences rather than size differences between castes in the Neotropical swarm-founding wasp Metapolybia docilis (Hymenoptera: Vespidae, Epiponini)

BackgroundSwarm-founding epiponine wasps are an intriguing group of social insects in which colonies are polygynic (several queens share reproduction) and differentiation between castes is often not obvious. However, caste differences in some may be more pronounced in later phases of the colony cycle.ResultsUsing morphometric analyses and multivariate statistics, it was found that caste differences in Metapolybia docilis are slight but more distinct in latter stages of the colony cycle.ConclusionsBecause differences in body parts are so slight, it is proposed that such variation may be due to differential growth rates of body parts rather than to queens being larger in size, similar to other previously observed epiponines.

Demographic mechanisms for the evolution of long life in social insects.

One of the most remarkable life history traits of many species of eusocial insects is the extraordinary longevity of their queens, with the recorded life span of some individuals exceeding 30 years. Surprisingly, little research has been conducted on either the proximate or ultimate questions concerning what factors favor the evolution of the long life spans in social insects. The broad objective of this paper is to address four sets of questions: (1) What are the similarities and differences in the longevity patterns among species in the four main groups of social insects - bees, wasps, ants and termites? (2) What is the evolutionary association of longevity and sociality in insects? (3) Are there biodemographic principles that can be derived from studies on longevity in social insects? and (4) What are the general implications for longevity in vertebrates, including humans?

Distributed problem solving in social insects

In a social insect colony, large numbers of individuals all follow the same set of behavioral rules. Without centralized control, these individuals' interactions with each other and with their environment result in the allocation of individuals to various tasks, and in the distribution of foragers among available food sources. We review this highly parallel and distributed form of information processing, discussing its potential sophistication, its actual performance in various groups of social insects, its general strengths and liabilities, and finally, the adaptations that compensate for these liabilities.

Effect of Synthetic ‘Queen Substance’ (9-oxodec-trans-2-enoic acid) on Ovary Development of the House-fly, Musca domestica L.

RECENT investigations1 have shown that when ‘queen substance’ (9-oxodec-trans-2-enoic acid) is distributed throughout the colony of honey bees, it partially inhibits the development of ovaries in workers and influences their behaviour by inhibiting queen-cell construction. Similar effects of crude extracts of queen honey bees have been demonstrated on other groups of social insects, for example, Formica fusca L.2 and Kalotermes flavicollis F.3 This non-specificity for orders of social insects suggests that ‘queen substance’ might have similar inhibitory effects on ovary development even of non-social insects. This hypothesis was tested by examining the effect of synthetic ‘queen substance’ on ovary development in the house-fly, Musca domestica L.