The degree of genetic heterogeneity among the individuals in an animal society depends on the society's genetic structure. Genetic heterogeneity, in turn, means that group members will differ in their reproductive objectives and conflicts over reproduction may arise. The resolution of these conflicts may be reflected in the way that reproduction is partitioned between potential reproductives. We used 5 microsatellite loci to investigate genetic structure and reproductive skew in 17 nests of the Malaysian hover wasp, Parischnogaster alternata. Parischnogaster alternata colonies are small (1–10 females), and all adult colony members are capable of mating and producing offspring. We found that colonies tended to consist of closely related individuals and that at any one time the production of both female and male offspring was nearly always monopolized by a single dominant female, despite considerable variation between nests in parameters predicted to affect skew. Subordinate females that remained in their natal colonies obtained indirect fitness benefits by helping to raise offspring to which they were related. Subordinate females also appeared to be positioned within an age-based queue for inheritance of the dominant egg-laying position. We suggest that the high skew in P. alternata may result from strong ecological constraints on solitary nesting, high relatedness, and a relatively high probability that subordinates will eventually inherit the position of dominance.
We explore the effects of group size on the direct reproductive success of subordinate helpers in eusocial animals where only a single, dominant individual reproduces at one time. Helpers can reproduce directly if they inherit dominance, but when dominance is age based, an individual born into a larger group has a longer wait to inherit. We show that this disincentive to help can potentially be offset by increased productivity, increased life span, and insurance-based benefits for helpers if they survive to inherit dominance in larger groups. We analyze a field experiment in which group size was manipulated in the hover wasp Liostenogaster flavolineata. Productivity increased linearly with group size, larger groups were less likely to fail, and dominants in larger groups may have lived longer. Combined with the probability of inheriting dominance, these effects led overall to a negative correlation between group size and expected direct fitness, mainly because group size decreased during our study period, so that helpers could not expect to inherit as large a group as they started queuing in. Our analysis suggests that the relationship between group size and productivity plays a central role in determining the fitness consequences of helping.
Recent field experiments suggest that cooperative breeding in vertebrates can be driven by a shortage of breeding territories. We did analogous experiments on facultatively eusocial hover wasps (Stenogastrinae: Liostenogaster flavolineata). We provided nesting opportunities by removing residents from 39 nests within a large aggregation (1995), and by glueing 20 nests obtained from a distant site into a second aggregation (1996). We prevented nest–less floaters from competing for these opportunities in 1995 but not in 1996. In both years, helpers in unmanipulated groups were given opportunities to nest independently without having to incur nest–building costs and with a reduced wait before potential helpers emerged. Helpers visited the nests we provided, but adopted only a small proportion (5% of 111 vacancies created in 1995). Others were adopted by floaters, but a significant proportion of nests were never adopted (nine out of 20 in 1995, seven out of 20 in 1996). Helpers that visited nests did not originate from particular kinds of social group. Nests containing older brood were more likely to be adopted, and adopting females rarely destroyed older brood. A general feature of social insect but not vertebrate life–histories, the long period of offspring dependency relative to the short life expectancy of adult carers, may be a key factor constraining independent nesting.
Recent explanations for the evolution of eusociality, focusing more on costs and benefits than relatedness, are largely untested. We validate one such model by showing that helpers in foundress groups of the paper wasp Polistes dominulus benefit from an insurance–based mechanism known as Assured Fitness Returns (AFRs). Experimental helper removals left remaining group members with more offspring than they would normally rear. Reduced groups succeeded in preserving the dead helpers' investment by rearing these extra offspring, even when helper removals occurred long before worker emergence. While helpers clearly gained from AFRs, offspring of lone foundresses failed after foundress death, so that AFRs represent a true advantage for helpers. Smaller, less valuable offspring were probably sacrificed to feed larger offspring, but reduced groups did not preferentially attract joiners or increase their foraging effort to compensate for their smaller workforce. We failed to detect a second insurance–based advantage, Survivorship Insurance, in which larger groups are less likely to fail than smaller groups. We suggest that through their use of small offspring as a food store to cope with temporary shortages, wasps may be less susceptible than vertebrates to offspring failure following the death of group members.
'%3e%3cpath fill='%23012169' d='M0 0v30h60V0z'/%3e%3cpath stroke='%23fff' stroke-width='6' d='m0 0 60 30m0-30L0 30'/%3e%3cpath stroke='%23C8102E' stroke-width='4' d='m0 0 60 30m0-30L0 30' clip-path='url(%23b)'/%3e%3cpath stroke='%23fff' stroke-width='10' d='M30 0v30M0 15h60'/%3e%3cpath stroke='%23C8102E' stroke-width='6' d='M30 0v30M0 15h60'/%3e%3c/g%3e%3c/svg%3e)
'/%3e%3cuse xlink:href='%23a' transform='rotate(72 0 0)'/%3e%3cuse xlink:href='%23a' transform='rotate(-72 0 0)'/%3e%3cuse xlink:href='%23a' transform='scale(-1 1) rotate(72)'/%3e%3c/g%3e%3c/defs%3e%3cpath fill='%23012169' d='M0 0h1200v600H0z'/%3e%3cpath stroke='%23FFF' stroke-width='60' d='m0 0 600 300M0 300 600 0' clip-path='url(%23b)'/%3e%3cpath stroke='%23C8102E' stroke-width='40' d='m0 0 600 300M0 300 600 0' clip-path='url(%23c)'/%3e%3cpath stroke='%23FFF' stroke-width='100' d='M300 0v300M0 150h600' clip-path='url(%23b)'/%3e%3cpath stroke='%23C8102E' stroke-width='60' d='M300 0v300M0 150h600' clip-path='url(%23b)'/%3e%3cuse xlink:href='%23d' fill='%23FFF' transform='matrix(45.4 0 0 45.4 900 120)'/%3e%3cuse xlink:href='%23d' fill='%23C8102E' transform='matrix(30 0 0 30 900 120)'/%3e%3cg transform='rotate(82 900 240)'%3e%3cuse xlink:href='%23d' fill='%23FFF' transform='rotate(-82 519.022 -457.666) scale(40.4)'/%3e%3cuse xlink:href='%23d' fill='%23C8102E' transform='rotate(-82 519.022 -457.666) scale(25)'/%3e%3c/g%3e%3cg transform='rotate(82 900 240)'%3e%3cuse xlink:href='%23d' fill='%23FFF' transform='rotate(-82 668.57 -327.666) scale(45.4)'/%3e%3cuse xlink:href='%23d' fill='%23C8102E' transform='rotate(-82 668.57 -327.666) scale(30)'/%3e%3c/g%3e%3cuse xlink:href='%23d' fill='%23FFF' transform='matrix(50.4 0 0 50.4 900 480)'/%3e%3cuse xlink:href='%23d' fill='%23C8102E' transform='matrix(35 0 0 35 900 480)'/%3e%3c/svg%3e)
