Annagiri Sumana

Choosing an appropriate index to construct dominance hierarchies in animal societies: a comparison of three indices

A plethora of indices have been proposed and used to construct dominance hierarchies in a variety of vertebrate and invertebrate societies, although the rationale for choosing a particular index for a particular species is seldom explained. In this study, we analysed and compared three such indices, viz Clutton-Brock et al.s index (CBI), originally developed for red deer, Cervus elaphus, Davids score (DS) originally proposed by the statistician H. A. David and the frequency-based index of dominance (FDI) developed and routinely used by our group for the primitively eusocial wasps Ropalidia marginata and Ropalidia cyathiformis. Dominance ranks attributed by all three indices were strongly and positively correlated for both natural data sets from the wasp colonies and for artificial data sets generated for the purpose. However, the indices differed in their ability to yield unique (untied) ranks in the natural data sets. This appears to be caused by the presence of noninteracting individuals and reversals in the direction of dominance in some of the pairs in the natural data sets. This was confirmed by creating additional artificial data sets with noninteracting individuals and with reversals. Based on the criterion of yielding the largest proportion of unique ranks, we found that FDI is best suited for societies such as the wasps belonging to Ropalidia, DS is best suited for societies with reversals and CBI remains a suitable index for societies such as red deer in which multiple interactions are uncommon

Diploid males and their triploid offspring in the paper wasp Polistes dominulus.

Although the hymenopteran sex-determining mechanism generally results in haploid males and diploid females, diploid males can be produced via homozygosity at the sex-determining locus. Diploid males have low fitness because they are effectively sterile or produce presumably sterile triploid offspring. Previously, triploid females were observed in three species of North American Polistes paper wasps, and this was interpreted as indirect evidence of diploid males. Here we report what is, to our knowledge, the first direct evidence: four of five early male-producing Polistes dominulus nests from three populations contained diploid males. Because haploid males were also found, however, the adaptive value of early males cannot be ignored. Using genetic and morphological data from triploid females, we also present evidence that both diploid males and triploid females remain undetected throughout the colony cycle. Consequently, diploid male production may result in a delayed fitness cost for two generations. This phenomenon is particularly relevant for introduced populations with few alleles at the sex-determining locus, but cannot be ignored in native populations without supporting genetic data. Future research using paper wasp populations to test theories of social evolution should explicitly consider the potential impacts of diploid males.

The function of dart behavior in the paper wasp, Polistes fuscatus

Dominance behavior in Polistes wasps is a composite trait consisting of various discrete behaviors such as darts, lunges, bites, and mounts. The majority of these behaviors are considered ‘aggressive’, and these aggressive behaviors are considered to form a continuum from mild (e.g., darts) to severe (e.g., falling fights). In this paper we focus on darts, the most common of the dominance behaviors, and investigate their function in un-manipulated post-emergent colonies of the primitively eusocial wasp P. fuscatus. Here we show that darts are correlated with the more severe dominance behaviors, and that dominance ranks do not change with the addition or exclusion of darts. We find no correlation, however, between receiving darts and receiving more severe dominance behaviors. This result suggests that darts are not indicative of aggressive reinforcement of dominance, but rather may serve a different function. Our data suggest that the function of darts is to regulate activity on nests. Both foundresses and workers dart inactive workers significantly more often than by chance, and workers respond to a foundress’s (but not a worker’s) dart by becoming less inactive. We also found that active workers who receive a dart from either a foundress or worker respond mostly by switching from one activity to another. Thus, our data suggest that darts are not aggressive behaviors, that foundresses use this signal to initiate activity, and that foundresses and workers both use the signal to regulate worker activity.

Key relocation leaders in an Indian queenless ant.

Division of labor is a central feature in social insects, wherein, simple individuals come together in groups to perform tasks that could be quite complex. It is generally believed that individuals who perform a specific task are themselves simple, interchangeable units. However, the variances in the performances of these individual insects need to be explored in greater detail. In this study, individual specialization in the context of colony relocation was examined in the Indian ponerine ant Diacamma indicum. One ant termed the maximum tandem leader (Max TL) was found to have a key role. Max TL performed 24% of the adult relocation in the colony and recruited more tandem leaders than other leaders thereby contributing to the organization of the relocation. The Max TLs role in the relocation process was further examined by comparing control relocations with experiments in which the Max TL was removed during the relocation process. Even though all the colonies relocated successfully, the relocation dynamics was significantly altered in the absence of the Max TL. We find that a single individual, the Max TL, takes up roles of a performer, organizer and catalyst during the colony relocation process, which challenges the norm that all workers are equal.

Workers of the primitively eusocial wasp Ropalidia marginata do not perceive their queen across a wire mesh partition

Queens of primitively eusocial wasps generally have active and behaviourally dominant queens who use physical aggression to suppress worker reproduction. Although a Ropalidia marginata queen is strikingly docile and behaviourally non-dominant, she is completely successful in maintaining reproductive monopoly. R. marginata queens must achieve such reproductive monopoly by some means other than overt physical aggression. Upon loss or removal of the queen, one of the workers (referred to as the potential queen) becomes extremely aggressive and will eventually go on to become the next queen of the colony, if the original queen is not returned. The fact that potential queens are not discernible in the presence of the queen but become obvious within minutes after removal of the queen raises the question of how workers in general and the potential queens in particular, perceive the presence or absence of their queens. Here, we have conducted experiments in which we separate half of the workers from their queen by a wire mesh screen and study their behavioural response to such separation. We demonstrate that the presence of the queen is not perceived across the wire mesh screen, which suggests that if the queen uses a pheromone to signal her presence, then that pheromone is not very volatile.

Characterization of recruitment through tandem running in an Indian queenless ant Diacamma indicum

Tandem running is a primitive recruitment method employed by many ant genera. This study characterizes this behaviour during the recruitment of colony mates to a new nest in an Indian ant Diacamma indicum. Tandem leaders who have knowledge of the new nest lead a single follower at a time, to the destination by maintaining physical contact. In order to characterize tandem running, we captured and analysed 621 invitations, 217 paths and 226 termination events. Remarkably, not a single colony member was lost. While invitations were stereotypic in behaviour, termination was not. Analysis of speed revealed that the average transport speed was 4.2 cm s−1. Coupled adult-brood transport was slower than other transports but was more efficient than individual trips. Comparing tandem running with other popular recruitment methods in ants allows us to postulate that even though tandem running is primitive it is probably just another means to achieve the same end.

Social insect colony as a biological regulatory system: modelling information flow in dominance networks

Social insects provide an excellent platform to investigate flow of information in regulatory systems since their successful social organization is essentially achieved by effective information transfer through complex connectivity patterns among the colony members. Network representation of such behavioural interactions offers a powerful tool for structural as well as dynamical analysis of the underlying regulatory systems. In this paper, we focus on the dominance interaction networks in the tropical social wasp Ropalidia marginata—a species where behavioural observations indicate that such interactions are principally responsible for the transfer of information between individuals about their colony needs, resulting in a regulation of their own activities. Our research reveals that the dominance networks of R. marginata are structurally similar to a class of naturally evolved information processing networks, a fact confirmed also by the predominance of a specific substructure—the ‘feed-forward loop’—a key functional component in many other information transfer networks. The dynamical analysis through Boolean modelling confirms that the networks are sufficiently stable under small fluctuations and yet capable of more efficient information transfer compared to their randomized counterparts. Our results suggest the involvement of a common structural design principle in different biological regulatory systems and a possible similarity with respect to the effect of selection on the organization levels of such systems. The findings are also consistent with the hypothesis that dominance behaviour has been shaped by natural selection to co-opt the information transfer process in such social insect species, in addition to its primal function of mediation of reproductive competition in the colony.