The Biology Of Two Sexes : A Study Of The Primitively Eusocial Wasp Ropalidia Marginata
A striking feature of hymenopterans societies is the absence of male workers. Foraging, nest building, brood care and all other activities required for the functioning of the colony are carried out by the females. These behaviours of the females and the implied cooperation and altruism have led social insect researchers to focus almost exclusively on the female members of hymenopterans societies. As a consequence, males have remained relatively neglected. The imbalance in the attention paid by researchers to females and males has been even more striking in the case of the extensively studied primitively eusocial wasp Rosalinda marginata. I have therefore focussed most of my attention to male R. marginata but of course, wherever possible and appropriate, I have compared the males with the females. Since almost nothing was known about R. marginata males, I began by obtaining basic information on the natural history and behaviour of the males. Male R. marginata are smaller in size and are different in the shape of the head as compared to the females. By conducting a three year survey, I found that just as nests of this species are found throughout the year, so are the males. The presence of males throughout the year has implications for the evolution of sociality by making available the opportunity to mate and found new nests, for females eclosing at any time of the year. Unlike the females who spend all their life on their nests, males stay on their natal nests only for one to twelve days (mean ± sd: 6.0 ± 2.6, N = 55) and leave their natal nest once and for all, to lead a nomadic life. It is difficult to determine how long males live after leaving their natal nests. However, I maintained males in the laboratory with ad libitum food and found that under these conditions they can live up to 140 days (mean physiological life span ± SD: 61.3 ± 28.0, N = 106). Like all eusocial hymenopterans males, R. marginata males also do not take part in any colony maintenance activities. They however, occasionally perform the following behaviours: solicit food, antennae nest, antennae another wasp, feed self, snatch food (from females), fan wings, body jerk and wing jerk, dominance and subordinate behaviours. Females of course perform all of these behaviours and many more. But there are no behaviours which are restricted to the males. Borrowing methods used by ecologists to measure species richness and diversity, I have computed the behavioral richness and diversity of male and female R. marginata. As expected, female behaviour is richer and more diverse compared to males. Comparing what the males did during their short stay on the nest and what females did during their long stay on the nest, I found that males did not forage or feed larvae (although I recorded one male feeding larvae thrice on one occasion when there seemed to be excess food on the nest). Males showed, dominance and subordinate behaviours and being solicited behaviour, significantly less often than females. On the other hand, males showed higher frequencies of feeding self and soliciting behaviour. However, these comparisons may not be fair because what males do in the first few days of their life is being compared with what females do throughout their life. Hence I truncated the female data at six days to make it comparable to the average age of the males on the nest. Even after doing so I found similar differences except that the males show similar rates of feed self and higher rates of subordinate behaviour compared to the young females. As mentioned in the beginning, absence of male workers is a striking feature of social Hymenoptera. I therefore naturally turned my attention to the possible reasons for this. As there has been much speculation on the ultimate, evolutionary explanations for why males do not work, I decided to investigate possible proximate explanations. To make my goal experimentally tractable, I decided to focus on the behaviour of feeding larvae as an example of work. In spite of the fact that R. marginata has been studied for over two decades, male R. marginata had never been seen to feed larvae, before my study. I advanced three hypotheses for why males do not feed larvae. (1) Males are incapable of feeding larvae. (2) Males do not feed larvae because they have insufficient access to food to satisfy themselves and to feed the larvae. (3) Males do not feed larvae because females perform the same task very efficiently. In a series of experiments designed to test these hypotheses, I showed that males are indeed capable of feeding larvae; they do so at small frequencies when given access to additional food and do so at impressive frequencies when deprived of females, provided with excess food and confronted with hungry larvae. Nevertheless, their ‘feed larva’ behaviour is less sophisticated and relatively inefficient compared to females. Thus I have shown male wasps can work, given an opportunity. In addition to being very satisfying, this result negates the preadaptation hypothesis which argues that male social Hymenoptera do not work because their solitary ancestors did not. In spite of what I have shown above, mating remains the main roleof the males. Therefore, I next turned my attention to a study of mating behaviour. It is well known that mating never takes place on the nest of R. marginata. Although some sporadic attempts had been made before, mating behaviour had never been observed in laboratory conditions before my study. In a series of trial-and-error pilot experiments, varying the age, cage size, number of wasps per cage, period of isolation from other wasps, lighting conditions etc., and with considerable help from enthusiastic volunteers among my lab-mates, I succeeded in observing mating behaviour in the laboratory. In the final, standardized experimental set up, we introduced a single male and a single female wasp, both isolated from their nests and other wasps for at least five days, into an aerated transparent plastic box and made observations for one hour. Using such an experimental set up, we first made a detailed qualitative description of mating behaviour. All behavioral interactions were initiated by the males. Males often attempted to mount the females but sometimes the females flew away, making the attempt to mount unsuccessful. On other occasions males successfully mounted the females, which involved climbing on the female and drumming and rubbing his antennae and abdomen on the corresponding parts of the female body. On some occasions mounting led to interlocking of the abdominal tips, a process we refer to as ‘conjugation’. Sometimes the conjugation lasted less than five seconds and during which the male remained on the back of the female; this was referred to as short conjugation (SC). But at other times the conjugation lasted for more than 20 seconds, and the male flipped on its back; this was Referred to as long conjugation (LC). I dissected all females, involved ineither LC or SC or both. Of the 47 pairs, 21 pairs mated successfully as judged by the transfer of sperm into the spermatheca. With the goal of developing a strategy to obtain live mated females for any future experiments, we attempted to determine the behavioral correlate/s of successful sperm transfer. It turns out that SC is inadequate for sperm transfer and LC is a good predictor of sperm transfer. Five to twenty days of age is optimal for mating for both males and females. There was a significant reduction in the probability of mating when one or both partners were younger or older than 5-20 days. We did not find any evidence for nestmate discrimination in the context of mating. This is not surprising because mating takes place away from the nest and active nestmate discrimination may therefore be unnecessary to avoid inbreeding. Under these experimental conditions neither body size of males and females nor ovarian conditions of the females appeared to influence mating success. I hope that my studies will enhance attention of future researchers to the males and will also facilitate experiments requiring mated wasps and permit the study of mate choice and other reproductive behaviours in this otherwise well- studied species.