Social Organisation And Cooperation In Genetically Mixed Colonies Of The Primitively Eusocial Wasp, Ropalidia Marginata
Altruism in its extreme form is seen in social insects where most individuals give up their own reproduction and work to rear the offspring of their queen. The origin and evolution of such sterile worker castes remains a major unsolved problem in evolutionary biology. Primitively eusocial polistine wasps are an attractive model system for investigating this phenomenon. Ropalidia marginata (Lep.) (Hymenoptera: Vespidae) is one such tropical primitively eusocial wasp, in which new nests are initiated either by a single foundress or by a group of female wasps. Worker behaviour in Ropalidia marginata cannot be satisfactorily explained by the haplodiploidy hypothesis due to the existence of polyandry and serial polygyny which reduce intra-colony genetic relatedness to levels lower than the value expected between a solitary foundress and her offspring. Besides, wasps appear to move frequently between newly initiated nests, perhaps further reducing intracolony genetic relatedness. To study social organization and examine the possibility of kin recognition and task specialization under conditions of low intra-colony relatedness, genetically mixed colonies were created by introducing alien one-day old wasps onto recipient nests. As a first step I have tried to determine the factors that influence the acceptance of foreign wasps onto established colonies. I have introduced wasps between 1 to 20 days of age from donor colonies located at least 10 km away onto 12 different recipient colonies, observed these wasps for a period of 10 hours and later dissected them to examine their ovarian condition. Observations were carried out in the blind i.e. the observer was unaware of the identity of the wasps. Wasps upto 6 days of age were accepted by the alien nests. Older wasps may have been rejected because their relatively better ovarian condition may have been perceived as a reproductive threat to the recipient nest. Alternatively, younger wasps may have been accepted because they may be more easily moulded to the desired roles or due to some other correlate of age per se independent of ovarian condition. Although ovarian condition appeared to influence the probability of acceptance, it was not statistically significant in the presence of age in multiple regression models, making a favourable case for the 'ease of moulding hypothesis' or 'age per se hypothesis' over the 'reproductive threat hypothesis'. In any case these findings gave me a method to create genetically mixed colonies. On 12 different nests Ropalidia marginata, I similarly introduced one-day old wasps and thus created genetically mixed colonies. Such an introduction simulates the eclosion of distantly related individuals which is quite common on nests of R. marginata due to the presence of serial polygyny. About 7 such wasps were introduced per colony and the introductions were so arranged as to matched with natural eclosions on the recipient nest. After 7 days following the last introduction, colonies were observed for 20 hours each. Alien wasps became well integrated and performed most of the behaviours and tasks shown by the natal wasps. There was no evidence of kin recognition or task specialization between natal and introduced wasps. The introduced wasps also sometimes became replacement queens. In an attempt to test the costs in terms of brood rearing efficiency, of living in such genetically variable groups, I created kin and non-kin pairs of wasps in plastic containers. They were provided with ad libitum food, water and building material. The nests initiated were monitored till an adult offspring eclosed. There were no detectable differences in either the productivities or the developmental periods of immature stages in the kin and nonkin pairs suggesting that there is no apparent cost of living with unrelated or distantly related individuals. To compare the extent of cooperation between the two wasps in kin and non-kin pairs, I conducted behavioural observations on 12 pairs each of kin and nonkin wasps. I found no difference in the rates at which the non-egg layers brought food and pulp, fed larvae and built the nest in the kin and nonkin pairs suggesting that cooperative nest building and brood rearing was common to the kin as well as non-kin pairs. The results reported here strengthen the idea that factors other than genetic relatedness must play a prominent role in the maintenance of worker behaviour in Ropalidia marginata.