Sex investment ratios in eusocial Hymenoptera support inclusive fitness theory

Inclusive fitness theory predicts that sex investment ratios in eusocial Hymenoptera are a function of the relatedness asymmetry (relative relatedness to females and males) of the individuals controlling sex allocation. In monogynous ants (with one queen per colony), assuming worker control, the theory therefore predicts female-biased sex investment ratios, as found in natural populations. Recently, E.O. Wilson and M.A. Nowak criticized this explanation and presented an alternative hypothesis. The Wilson–Nowak sex ratio hypothesis proposes that, in monogynous ants, there is selection for a 1 : 1 numerical sex ratio to avoid males remaining unmated, which, given queens exceed males in size, results in a female-biased sex investment ratio. The hypothesis also asserts that, contrary to inclusive fitness theory, queens not workers control sex allocation and queen–worker conflict over sex allocation is absent. Here, I argue that the Wilson–Nowak sex ratio hypothesis is flawed because it contradicts Fisher’s sex ratio theory, which shows that selection on sex ratio does not maximize the number of mated offspring and that the sex ratio proposed by the hypothesis is not an equilibrium for the queen. In addition, the hypothesis is not supported by empirical evidence, as it fails to explain ‘split’ (bimodal) sex ratios or data showing queen and worker control and ongoing queen–worker conflict. By contrast, these phenomena match predictions of inclusive fitness theory. Hence, the Wilson–Nowak sex ratio hypothesis fails both as an alternative hypothesis for sex investment ratios in eusocial Hymenoptera and as a critique of inclusive fitness theory

Colony and individual life-history responses to temperature in a social insect pollinator

Pollinating insects are of major ecological and commercial importance, yet they may be facing ecological disruption from a changing climate. Despite this threat, few studies have investigated the life-history responses of pollinators to experimentally controlled changes in temperature, which should be especially informative for species with complex life histories such as eusocial insects. This study uses the key pollinator Bombus terrestris, a eusocial bumble bee with an annual colony cycle, to determine how temperature affects life-history traits at both individual and colony levels. In two laboratory experiments, we reared B. terrestris colonies at either 20°C or 25°C, and measured differences in a set of life-history traits including colony longevity, queen longevity, worker longevity, production of workers, production of sexuals (queen and male production) and growth schedule, as well as effects on thermoregulatory behaviours. Higher rearing temperature had a significant positive effect on colony longevity in one of the two experiments but no significant effects on queen or worker longevity. Higher rearing temperature significantly increased colony size but did not affect the timing of peak colony size. It was also associated with significantly higher queen production but had no effect on the production of workers or males or the timing of male production. Higher temperature colonies exhibited significantly more wing-fanning by workers and significantly less wax canopy construction. Hence an increase in rearing temperature of a few degrees increased colony longevity, colony size and queen production. However, individual longevity was not affected and so may have been buffered by changes in costly thermoregulatory behaviours. We conclude that eusocial insects may show complex phenotypic responses to projected temperature increases under climate change, including effects on productivity and reproduction at the colony level. Such effects should be considered when predicting the impact of climate change on the provision of essential pollination services

Inclusive fitness and the major transitions in evolution

Inclusive fitness theory is the leading framework for explaining the major transitions in evolution, whereby free-living subunits (e.g. cells, organisms) have cooperated to form new, higher-level units (e.g. organisms, eusocial societies). The theory has attracted considerable controversy. From a brief survey of the controversy's present status, I conclude that inclusive fitness theory continues to provide both a concept and a principled modelling tool of value for understanding social evolution, including major transitions. Turning to new developments in the study of major transitions, I describe work defining the point of occurrence of major transitions and, from inclusive fitness theory, the required conditions. I also suggest that it remains important to understand the evolution of individuality that occurs beyond such thresholds

Pacific Island food security: situation, challenges and opportunities

In recent years, there have been large and rapid increases in the prices of basic foods worldwide. Pacific island countries have not been isolated from the global food 'crisis'. This article shows that there are substantial differences in the impacts within and between Pacific island countries. To be effective and not counterproductive, policy and donor responses need to be tailored accordingly. This article recommends a combination of measures directed at mitigating the threats and empowering rural people to take advantage of the opportunities

Queen control of a key life-history event in a eusocial insect

In eusocial insects, inclusive fitness theory predicts potential queen–worker conflict over the timing of events in colony life history. Whether queens or workers control the timing of these events is poorly understood. In the bumble-bee Bombus terrestris, queens exhibit a ‘switch point’ in which they switch from laying diploid eggs yielding females (workers and new queens) to laying haploid eggs yielding males. By rearing foundress queens whose worker offspring were removed as pupae and sexing their eggs using microsatellite genotyping, we found that queens kept in the complete absence of adult workers still exhibit a switch point. Moreover, the timing of their switch points relative to the start of egg-laying did not differ significantly from that of queens allowed to produce normal colonies. The finding that bumble-bee queens can express the switch point in the absence of workers experimentally demonstrates queen control of a key life-history event in eusocial insects. In addition, we found no evidence that workers affect the timing of the switch point either directly or indirectly via providing cues to queens, suggesting that workers do not fully express their interests in queen–worker conflicts over colony life history

Observations of Global and Local Infall in NGC 1333

We report ``infall asymmetry'' in the HCO$^+$ (1--0) and (3--2) lines toward NGC 1333, extended over $\sim 0.39 {\rm pc}^2$, a larger extent than has been reported be fore, for any star-forming region. The infall asymmetry extends over a major portion of the star-forming complex, and is not limited to a single protostar, or to a single dense core, or to a single spectral line. It seems likely that the infall asymmetry represents inward motions, and that these motions are physically associated with the complex. Both blue-asymmetric and red-asymmetric lines are seen, but in both the (3--2) and (1--0) lines of HCO$^+$ the vast majority of the asymmetric lines are blue, indicating inward motions. The (3--2) line, tracing denser gas, has the spectra with the strongest asymmetry and these spectra are associated with the protostars IRAS 4A and 4B, which most likely indicates a warm central source is affecting the line profiles. The (3--2) and (1--0) lines usually have the same sense of asymmetry in common positions, but their profiles differ significantly, and the (1--0) line appears to trace motions on much larger spatial scales than does the (3--2) line. Line profile models fit the spectra well, but do not strongly constrain their parameters. The mass accretion rate of the inward motions is of order 10$^{-4}$ M$_\odot$/yr, similar to the ratio of stellar mass to cluster age.Comment: 28 pages, 11 figures, 1 colour figur

Managing farmed landscapes for pollinating insects

Increasing floral resources and improving habitat conditions can benefit pollinating insect species, wildflowers and crop production