A comparative social network analysis of wasp colonies and classrooms: Linking network structure to functioning

A major question in current network science is how to understand the relationship between structure and functioning of real networks. Here we present a comparative network analysis of 48 wasp and 36 human social networks. We have compared the centralisation and small world character of these interaction networks and have studied how these properties change over time. We compared the interaction networks of (1) two congeneric wasp species (Ropalidia marginata and Ropalidia cyathiformis), (2) the queen-right (with the queen) and queen-less (without the queen) networks of wasps, (3) the four network types obtained by combining (1) and (2) above, and (4) wasp networks with the social networks of children in 36 classrooms. We have found perfect (100%) centralisation in a queen-less wasp colony and nearly perfect centralisation in several other queen-less wasp colonies. Note that the perfectly centralised interaction network is quite unique in the literature of real-world networks. Differences between the interaction networks of the two wasp species are smaller than differences between the networks describing their different colony conditions. Also, the differences between different colony conditions are larger than the differences between wasp and children networks. For example, the structure of queen-right R. marginata colonies is more similar to children social networks than to that of their queen-less colonies. We conclude that network architecture depends more on the functioning of the particular community than on taxonomic differences (either between two wasp species or between wasps and humans).

Predatory and trophobiont-tending ants respond differently to fig and fig wasp volatiles

The interaction between figs and their pollinating or parasitic fig wasps is mediated largely by chemical communication. These fig wasps are often preyed upon by predatory ants. In this study, we found that predatory ants (Oecophylla smaragdina) patrolling Ficus racemosa trees were attracted to the odour from fig syconia at different developmental phases, as well as to the odours of fig wasps, whereas other predatory ants (Technomyrmex albipes) responded only to odours of syconia from which fig wasps were dispersing and to fig wasp odour. However, trophobiont-tending ants (Myrmicaria brunnea) patrolling the same trees and exposed to the same volatiles were unresponsive to fig or fig wasp odours. The predatory ants demonstrated a concentration-dependent response towards volatiles from figs receptive to pollinators and those from which wasps were dispersing while the trophobiont-tending ants were unresponsive to such odours at all concentrations. Naive predatory ants failed to respond to the volatiles to which the experienced predatory ants responded, indicating that the response to fig-related odours is learned. We suggest that predatory ants could use fig-associated volatiles to enhance their probability of wasp encounter and can eavesdrop on signals meant for pollinators. (C) 2009 The Association for the Study of Animal Behaviour.

Polymorphic microsatellite loci for primitively eusocial wasp Ropalidia marginata

We report here development and characterization of 48 novel microsatellite markers for Ropalidia marginata, a tropical, primitively eusocial polistine wasp from peninsular India. Thirty-two microsatellites showed polymorphism in a wild population of R. marginata (N = 38) collected from Bangalore, India. These markers will facilitate answering some interesting questions in ecology and evolutionary biology of this wasp, such as population structure, serial polygyny, intra-colony genetic relatedness and the pattern of queen succession.

Behavior of the indian social wasp ropalidia-cyathiformis on a nest of separate combs (hymenoptera, vespidae)

Observations were made on a nest of Ropalidia cyathiformis consisting of three combs. The number of eggs, larvae, pupae and adults were monitored at about 3-day intervals for a 2-month period. The behaviour of the adults was observed with special reference to the proportion of time spent on each of the three combs, the proportion of time spent away from the nest site and the frequencies of dominance interactions and egg laying. The adults moved freely between the three combs suggesting that all of them and all the three combs belonged to one nest. However, most of the adults preferred combs 2 and 3 over comb 1. Of the 10 animals chosen for a detailed analysis of behaviour, seven spent varying periods of time away from the nest site and oRen brought back food or building material. Five of the 10 animals laid at least one egg each but two adults monopolized most of the egg-laying. The animals showed a variety of dominance interactions on the basis of which they have been arranged in a dominance hierarchy. The dominant individuals laid most of the eggs and spent little or no time foraging, while the subordinate individuals spent more time foraging and laid few eggs or none. It is argued that R. cyathiformis is different from R. marginata, the only other Indian social wasp whose behaviour has been studied, in being at a more primitive stage of social organization.

Middle aged wasps mate through most of the year, without regard to body size, ovarian development and nestmateship: a laboratory study of the primitively eusocial wasp Ropalidia marginata

We studied the mating behaviour of the primi-tively eusocial wasp Ropalidia marginata and the factors that may influence sperm transfer. By introducing a male and a female R. marginata into ventilated transparent plastic boxes, we were able to observe mating behaviour, and it involved mounting and short or long conjugation of the wasps. Dissection of female wasps after the observation indicated that long conjugation is a good behavioural predictor of sperm transfer. This finding makes it possible to obtain mated females without dissecting them every time. We tested the effect of age, season, relatedness, body size and female's ovarian status on mating. Under laboratory conditions, mating success declined rapidly below and above the ages 5-20 days. Within this age range mating success was significantly low in December compared to other months tested. There was no nestmate discrimination, and there was no influence of male and female body size or of the ovarian state of the female on the probability of sperm transfer.

Comparative life-history traits in a fig wasp community: implications for community structure

1. Whether life-history traits can determine community composition and structure is an important question that has been well explored theoretically, but has received scant empirical attention. Life-history traits of a seven-member community of galler and parasitoid fig wasp species (Chalcidoidea), developing within the inflorescences (syconia) of Ficus racemosa (Moraceae) in India, were determined and used to examine community structure and ecology. 2. Gallers were pro-ovigenic (all eggs are mature upon adult emergence) whereas parasitoids were synovigenic (eggs mature progressively during adult lifespan). Initial egg load was correlated with body size for some species, and there was a trade-off between egg number and egg size across all species. Although all species completed their development and left the syconium concurrently, they differed in their adult and pre-adult lifespans. Providing sucrose solutions increased parasitoid lifespan but had no effect on the longevity of some galler species. While feeding regimes and body size affected longevity in most species, an interaction effect between these variables was detected for only one species. 3. Life-history traits of wasp species exhibited a continuum in relation to their arrival sequence at syconia for oviposition during syconium development, and therefore reflected their ecology. The largest number of eggs, smallest egg sizes, and shortest longevities were characteristic of the earliest-arriving galling wasps at the smallest, immature syconia; the converse characterised the later-arriving parasitoids at the larger, already parasitised syconia. Thus life history is an important correlate of community resource partitioning and can be used to understand community structure. 4. This is the first comprehensive study of life-history traits in a fig wasp community. The comparative approach revealed constraints and flexibility in trait evolution.

Regulation of Reproduction in the Primitively Eusocial Wasp Ropalidia marginata: on the Trail of the Queen Pheromone

Queens and workers are not morphologically differentiated in the primitively eusocial wasp, Ropalidia marginata. Upon removal of the queen, one of the workers becomes extremely aggressive, but immediately drops her aggression if the queen is returned. If the queen is not returned, this hyper-aggressive individual, the potential queen (PQ), will develop her ovaries, lose her hyper-aggression, and become the next colony queen. Because of the non-aggressive nature of the queen, and because the PQ loses her aggression by the time she starts laying eggs, we hypothesized that regulation of worker reproduction in R marginata is mediated by pheromones rather than by physical aggression. Based on the immediate loss of aggression by the PQ upon return of the queen, we developed a bioassay to test whether the queen's Dufour's gland is, at least, one of the sources of the queen pheromone. Macerates of the queen's Dufour's gland, but not that of the worker's Dufour's gland, mimic the queen in making the PQ decrease her aggression. We also correctly distinguished queens and workers of R. marginata nests by a discriminant function analysis based on the chemical composition of their respective Dufour's glands.

Natural history and behaviour of the primitively eusocial wasp Ropalidia marginata (Hymenoptera: Vespidae): a comparison of the two sexes

We report the natural history and behaviour of the primitively eusocial wasp Ropalidia marginata with a special reference to the males. We found that just as nests of this species are found throughout the year, so are the males. Females spend all their life in their nests but males stay in their natal nests only for 1-12 days and leave to lead a nomadic life. Males maintained in the laboratory can live for up to 140 days. Like all eusocial hymenopteran males, R. marginata males also do not perform any colony maintenance activities. We found that males did not forage or feed larvae. Compared with females, males showed fewer dominance and subordinate behaviours and being solicited behaviour and more feeding self and soliciting behaviours. By comparing males with young females, we found similar differences, except that the males showed similar rates of feeding self and higher rates of subordinate behaviour.

Temporal associations in fig-wasp-ant interactions: diel and phenological patterns

In a complex multitrophic plant-animal interaction system in which there are direct and indirect interactions between species, comprehending the dynamics of these multiple partners is very important for an understanding of how the system is structured. We investigated the plant Ficus racemosa L. (Moraceae) and its community of obligatory mutualistic and parasitic fig wasps (Hymenoptera: Chalcidoidea) that develop within the fig inflorescence or syconium, as well as their interaction with opportunistic ants. We focused on temporal resource partitioning among members of the fig wasp community over the development cycle of the fig syconia during which wasp oviposition and development occur and we studied the activity rhythm of the ants associated with this community. We found that the seven members of the wasp community partitioned their oviposition across fig syconium development phenology and showed interspecific variation in activity across the day-night cycle. The wasps presented a distinct sequence in their arrival at fig syconia for oviposition, with the parasitoid wasps following the galling wasps. Although fig wasps are known to be largely diurnal, we documented night oviposition in several fig wasp species for the first time. Ant activity on the fig syconia was correlated with wasp activity and was dependent on whether the ants were predatory or trophobiont-tending species; only numbers of predatory ants increased during peak arrivals of the wasps.

Seasonal variation in the onset of egg laying in a primitively eusocial wasp: implications for the evolution of sociality

When freshly eclosed females of the primitively eusocial wasp, Rapalidia marginata are isolated into individual cages, only about half of them build nests and lay eggs and those that do so take a long and variable amount of time (Mean +/- SD = 66 +/- 37 days) before they lay their first egg. Part of the reason for this delay is because, when kept in isolation, no wasp begins to lay eggs during a period of approximately 82 days from mid - October to early January. Wasps maintained at a constant temperature of 26 +/- 1-degrees-C however initiate egg laying throughout the year, suggesting that the low temperatures during mid - October to early January may be at least one factor that makes this period unfavourable for wasps maintained at room temperature. Egg laying continues more or less normally throughout October-January however, in all natural and laboratory colonies studied. Natural colonies of R. marginata are initiated throughout the year and often by groups of females. Huddling together is a striking feature of the wasps especially on cold mornings. We therefore suggest that the isolated animals in our experiment are unable to lay eggs during the coldest part of the year because of their inability to huddle together, share metabolic heat and perform "co-operative thermoregulation". Such "co-operative thermoregulation" may thus be another factor that facilitates the evolution of socialitly.

The role of larval nutrition in pre-imaginal biasing of caste in the primitively eusocial wasp Ropalidia marginata (Hymenoptera: Vespidae)

1. When freshly eclosed females of the primitively eusocial wasp Ropalidia marginata (Lep.) are isolated, only about 50% of them build nests and lay eggs thereby suggesting a pre-imaginal biasing of caste. 2. Wasps that lay eggs take a very variable amount of time after eclosion to start doing so. 3. Females eclosing from nests where larvae are fed at a relatively higher rate are more likely to become egg-layers and are likely to take less time after eclosion to begin to lay eggs. 4. Thus, both forms of pre-imaginal biasing of caste, namely, differences in egg laying capacity and differences in the time taken to attain reproductive maturity, appear to be influenced by larval nutrition.

On reconfirming the evidence for pre-imaginal caste bias in a primitively eusocial wasp

Caste is usually thought to be determined entirely in the adult stage in most primitively eusocial wasps and bees. A pre-imaginal caste bias has however been recently discovered in the primitively eusocial waspRopalidia marginata. This study also suggested that reigning queens and possibly other adults may influence the production of new queens and implied at least partial support to the parental manipulation or sub-fertility hypothesis for the evolution of insect sociality. The interest of these results prompted an attempt at their reconfirmation. Complete reconfirmation has now been obtained using data from an independent experiment and two additional methods of data analysis. We therefore conclude that caste is at least partly determined prior to eclosion in the primitively eusocial waspRopalidia marginata which lacks morphological differentiation between egg-layers and non-egg-layers.

Can Dufour's gland compounds honestly signal fertility in the primitively eusocial wasp Ropalidia marginata?

Unlike queens of typical primitively eusocial species, Ropalidia marginata queens are docile and non-interactive, and hence cannot be using dominance to maintain their status. It appears that the queen maintains reproductive monopoly through a pheromone, of which the Dufour's gland is at least one source. Here, we reconfirm earlier results showing that queens and workers can be correctly classified on a discriminant function using the compositions of their respective Dufour's glands, and also demonstrate consistent queen-worker differences based on categories of compounds and on single compounds also in some cases. Since the queen pheromone is expected to be an honest signal of the fecundity of a queen, we investigate the correlation of Dufour's gland compounds with ovarian activation of queens. Our study shows that Dufour's gland compounds in R. marginata correlate with the state of ovarian activation of queens, suggesting that such compounds may portray the fecundity of a queen, and may indeed function as honest signals of fertility.

Science as a hobby: how and why I came to study the social life of an Indian primitively eusocial wasp

For the most part, my research career has involved prying into the life of the locally abundant primitively eusocial paper wasp, Ropalidia marginata, with the aim of understanding the origin and evolution of social life in insects. My interest in this wasp species began as a hobby, but I was privileged to soon convert my hobby into my profession. Here I describe how this conversion came about, what it meant to pursue my hobby as a full-time activity, describe some examples from my research and end with some reflections about the process of doing modern science, especially in India.