The physiology and behavior of ectothermic organisms is strongly influenced by temperature. For ground nesting species like the primitively eusocial halictid bee, Lasioglossum malachurum, soil temperature might influence the life cycle as well as the complexity of the social group since the number of broods that can be fitted into the flight season might increase with increasing temperature. Our study population of L. malachurum at Wuerzburg exhibits a remarkable variability with respect to the number of broods and the pattern of sexual production. Broods are separated by activity pauses during which the larvae develop. In this study we investigate the influence of soil temperature on the pattern of nesting activity (duration of broods and pauses) and on the number of broods in L. malachurum. We observed a total of 1138 nests in 13 aggregations near Wuerzburg. As expected, soil temperature shortened the duration of the pauses, resulting in an overall shortening of the nesting cycle. This is most probably due to a physiological effect of soil temperature on the development of the larvae. With regard to the nesting strategies, we hypothesized that a shortening of the nesting cycle within the limited flight season should enhance the success of a strategy with more worker broods. In fact, patches with higher soil temperature showed more broods. However, this effect was rather weak, suggesting that other factors might have a stronger impact on the variability in nesting strategy within our study population of L. malachurum.
Foundress queens of social Hymenoptera require considerable amounts of energy for survival, solitary nest founding, provisioning of the first brood, and egg production. Energy reserves in insects mostly consist of fat. We investigated how hibernation and the subsequent flight season, especially the solitary nest founding phase, influenced the abdominal fat content of gynes in the primitively eusocial sweat bee, Lasioglossum malachurum (Hymenoptera, Halictidae). In our study population, sexuals are produced in both the second and the third broods. Emerging gynes of the third brood had significantly more fat than those of the second brood, whereas there was no such difference in males. As expected, fat reserves in samples of female sexuals caught at emergence, after hibernation, during solitary nest founding, and at the end of the social phase of the nest cycle indicate a severe decrease of reserves that was highest during the 7 weeks of the solitary founding phase. Thus, the amount of fat reserves of foundress queens seems to be crucial, particularly for nest founding. However, investment of energy reserves in the solitary nest founding phase has probably to be balanced with the subsequent social phase in a way that maximizes the queen's fitness. Possible consequences for the complexity and progress of the nest cycle are discussed.
The classical model of colony dynamics developed by Macevicz and Oster predicts that optimal colony fitness in annual eusocial insects is achieved by a bangbang strategy of reproduction: exclusive production of workers (ergonomic phase) followed by exclusive production of sexuals (reproductive phase). We propose an alternative model that assumes colony development in discrete broods and a limited overall investment potential of the queen. Based on the costs for producing eggs, workers, and sexuals and efficiency of individuals we predict the optimal number of workers and sexuals in the colony for each brood of the colony cycle that maximizes overall colony fitness. To link our model assumptions to the real world we chose model parameters according to field data of the halictid bee Lasioglossum malachurum. However, our model is representative of a large number of species with an annual life cycle and with discrete broods. Our model shows that the optimal partitioning of resources, i.e. the optimal workers/sexuals ratio depends on rearing cost for sexuals as well as productivity of workers but not on the queens total investment, egg cost, or rearing cost for workers. In complete accordance to Macevicz and Oster we predict a bang-bang reproduction strategy despite the differences in the basic assumptions. Potential deviations from this strategy and transitions from social to solitary breeding are discussed in the framework of our model.
BackgroundAccording to the classical model of Macevicz and Oster, annual eusocial insects should show a clear dichotomous "bang-bang" strategy of resource allocation; colony fitness is maximised when a period of pure colony growth (exclusive production of workers) is followed by a single reproductive period characterised by the exclusive production of sexuals. However, in several species graded investment strategies with a simultaneous production of workers and sexuals have been observed. Such deviations from the "bang-bang" strategy are usually interpreted as an adaptive (bet-hedging) response to environmental fluctuations such as variation in season length or food availability.To generate predictions about the optimal investment pattern of insect colonies in fluctuating environments, we slightly modified Macevicz and Oster's classical model of annual colony dynamics and used a dynamic programming approach nested into a recurrence procedure for the solution of the stochastic optimal control problem.Results1) The optimal switching time between pure colony growth and the exclusive production of sexuals decreases with increasing environmental variance. 2) Yet, for reasonable levels of environmental fluctuations no deviation from the typical bang-bang strategy is predicted. 3) Model calculations for the halictid bee Lasioglossum malachurum reveal that bet-hedging is not likely to be the reason for the graded allocation into sexuals versus workers observed in this species. 4) When environmental variance reaches a critical level our model predicts an abrupt change from dichotomous behaviour to graded allocation strategies, but the transition between colony growth and production of sexuals is not necessarily monotonic. Both, the critical level of environmental variance as well as the characteristic pattern of resource allocation strongly depend on the type of function used to describe environmental fluctuations.ConclusionUp to now bet-hedging as an evolutionary response to variation in season length has been the main argument to explain field observations of graded resource allocation in annual eusocial insect species. However, our model shows that the effect of moderate fluctuations of environmental conditions does not select for deviation from the classical bang-bang strategy and that the evolution of graded allocation strategies can be triggered only by extreme fluctuations. Detailed quantitative observations on resource allocation in eusocial insects are needed to analyse the relevance of alternative explanations, e.g. logistic colony growth or reproductive conflict between queen and workers, for the evolution of graded allocation strategies.
Background: Social insects show considerable variability not only in social organisation but also in the temporal pattern of nest cycles. In annual eusocial sweat bees, nest cycles typically consist of a sequence of distinct phases of activity (queen or workers collect food, construct, and provision brood cells) and inactivity (nest is closed). Since the flight season is limited to the time of the year with sufficiently high temperatures and resource availability, every break reduces the potential for foraging and, thus, the productivity of a colony. This apparent waste of time has not gained much attention.
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