Alternative strategies for a father with a small brood: mate, cannibalise or care

Brood cycling, the alternation of periods of courtship and mating activity ("mating phase") with days of full brood care ("parental phase"), and total filial cannibalism, the consumption of all current offspring by a parent, have been interpreted as a response to the conflicting demands faced by parental males during the breeding season. Under the assumption that these two behaviours result from trade-offs among mating, caring and feeding, the strategy adopted by the male should depend on egg number and age. This hypothesis was tested in the scissors-tail sergeant (Abudefduf sexfasciatus), a common tropical damselfish. Observations of undisturbed males and a brood reduction experiment confirmed all theoretical predictions. The shift from the mating to parental phase occurred when an appropriate number of eggs had been obtained. When the brood was reduced on the first day of the parental phase, males often reverted to the mating phase, but never did so if the manipulation occurred on the second or third day. Only broods that were smaller than average were consumed by the parental male. Artificial reduction of the brood increased the incidence of total filial cannibalism, but the probability of cannibalism decreased with brood age. Electronic Publication     

Effects of foundress number on brood raids and queen survival in the fire ant Solenopsis invicta

In several ant species, colonies are founded by small groups of queens (pleometrosis), which coexist until the first workers eclose, after which all but one queen is killed. It has been hypothesized that, by producing a larger cohort of workers, cooperating queens may increase colony success during brood raids, a form of competition in which brood and workers from losing nests are absorbed into winning colonies. To test whether this benefit is sufficient to favor pleometrosis, newly mated queens of the fire ant Solenopsis invicta were assembled in groups of one, two, three, or four, reared in the laboratory until the first workers eclosed, then planted in the field in replicated assemblages. The proportion of colonies engaging in brood raids increased with average foundress number per nest and with colony density but was unaffected by variance in foundress number among interacting colonies. Within mixed assemblages of single-queen and multiple-queen colonies, queen number had no effect on the likelihood of engaging in raids or the probability of nest survival through the brood raiding period. However, following nearly 30% of raids, queens moved to new nests and displaced the resident queens. When queen relocation and subsequent mortality were accounted for, it was found that the survival of queens from four-queen groups was substantially higher than that of solitary queens. By contrast, the survival of queens from two-queen colonies was no greater than that of solitary queens. These results show that the competitive advantages of multiple-queen colonies are sufficient to counterbalance the increased mortality of queens within groups only when the number of foundresses is greater than two and when colonies are founded at high density. When colonies lose brood raids, the workers appear to abandon their mothers to join surviving colonies. However, in laboratory experiments, queens attempting to enter foreign nests were significantly more likely to displace the resident queen if their own daughters were present within the invaded nest. Thus, workers may be able to bias the probability that their mother rejoins them and displaces competing queens.     

Eggshell strength of an obligate brood parasite: a test of the puncture resistance hypothesis

Eggs of several brood parasites have thicker and stronger shells than expected for their size. The present study evaluated the puncture resistance hypothesis for the occurrence of thick-shelled eggs in common cuckoos Cuculus canorus by investigating costs of cuckoo egg ejection in four Acrocephalus warblers—the great reed warbler A. arundinaceus, reed warbler A. scirpaceus, marsh warbler A. palustris and sedge warbler A. schoenobaenus. The three latter species all suffered ejection costs, while ejection was not costly in the larger great reed warbler. The occurrence of ejection costs was negatively related to host bill size. In the marsh warbler, we compared ejection costs in naturally parasitized nests and two experimental treatments, in which broods were parasitized artificially with great reed warbler and conspecific eggs. Hosts damaged their own eggs significantly more often when ejecting the thick-shelled cuckoo eggs than when ejecting the similarly sized but thinner-shelled great reed warbler eggs, providing some support for the puncture resistance hypothesis. Ejection of conspecific eggs did not involve any costs. Furthermore, contrary to predictions derived from the laying damage hypothesis, there was no evidence that egg damage was associated with cuckoo egg laying. Hosts damaging their own eggs during ejection were more likely to subsequently desert their clutches than those that did not. The frequency of clutches smeared with the contents of the ejected egg were positively related to the hypothesized difficulty of foreign egg puncturing. Potential advantages of thicker shells in common cuckoo eggs are discussed.     

Extra-pair paternity, offspring mortality and offspring sex ratio in the socially monogamous coal tit (Parus ater)

Females of many socially monogamous bird species commonly engage in extra-pair copulations. Assuming that extra-pair males are more attractive than the females’ social partners and that attractiveness has a heritable component, sex allocation theory predicts facultative overproduction of sons among extra-pair offspring (EPO) as sons benefit more than daughters from inheriting their father’s attractiveness traits. Here, we present a large-scale, three-year study on sex ratio variation in a passerine bird, the coal tit (Parus ater). Molecular sexing in combination with paternity analysis revealed no evidence for a male-bias in EPO sex ratios compared to their within-pair maternal half-siblings. Our main conclusion, therefore, is that facultative sex allocation to EPO is absent in the coal tit, in accordance with findings in several other species. Either there is no net selection for a deviation from random sex ratio variation (e.g. because extra-pair mating may serve goals different from striving for ‘attractiveness genes’) or evolutionary constraints preclude the evolution of precise maternal sex ratio adjustment. It is interesting to note that, however, we found broods without EPO as well as broods without mortality to be relatively female-biased compared to broods with EPO and mortality, respectively. We were unable to identify any environmental or parental variable to co-vary with brood sex ratios. There was no significant repeatability of sex ratios in consecutive broods of individual females that would hint at some idiosyncratic maternal sex ratio adjustment. Further research is needed to resolve the biological significance of the correlation between brood sex ratios and extra-pair paternity and mortality incidence, respectively.     

Provisional rejection of three alternative hypotheses on the maintenance of a size dichotomy in males of Dawson’s burrowing bee,Amegilla dawsoni (Apidae,Apinae, Anthophorini)

I examine three alternative hypotheses on the male size dimorphism of Dawson’s burrowing bees (Amegilla dawsoni) in which there are large (major) and small (minor) males. One possibility is that minor males are simply the incidental byproduct of environmental conditions that prevent females from provisioning brood cells optimally. This hypothesis is not supported by the finding that males of intermediate size are consistently rare in populations sampled across years and in different regions, nor can it easily account for the absence of a size dichotomy in females. A second possibility is that minors represent a “best of a bad job” response of those females that are small or otherwise disadvantaged. However, presumptive male siblings sometimes include both majors and minors, a result not predicted from this hypothesis. A third explanation is that female brood provisioning strategy results in the production of minors and majors with equal fitness benefit to fitness cost ratios. However, although it is true that minor males weigh on average about half what a major weighs, and so represent approximately half the provisioning expense of a major, minor males on average appear to secure far fewer than half the number of matings of majors. If the estimate of mating success of minors is accurate, the net gain to females from producing a minor son is unlikely to equal that derived from a major son. Therefore the third hypothesis must also be tentatively rejected, although with caution given the uncertainties in estimating the relative costs and benefits of producing major and minor sons. Received: 12 January 1996/Accepted after revision: 27 April 1996     

Patterns of brood division and an absence of behavioral plasticity in a neotropical passerine

Studies of parental behavior in various habitats provide an opportunity to gain insight into how different environments may mold strategies of parental care. Brood division by parents has been hypothesized to occur facultatively within and among species. Brood division occurs when each parent cares for specific offspring within a brood. We studied brood division in a neotropical passerine, the western slaty antshrike (Thamnophilus atrinucha). Our results present a unique picture of a highly specialized example of avian brood division. Division was a fixed behavioral pattern in the population studied: all broods divided by fledging and remained divided during the entire post-fledging period. Brood division before fledging, a previously unreported phenomenon, occurred in 40% of nests observed. Parents that preferentially fed a certain offspring (defined as their focal offspring) in the nest fed the same individual after fledging. Each parent fed only its focal offspring in broods of one and two. The male parent cared for the heavier offspring and the first offspring to leave the nest. Siblings were segregated spatially during the time of highest predation risk. These observations suggest that a consistently high risk of predation on offspring has favored initial spatial segregation and inflexibility of brood division behavior in this species. Factors other than predation risk alone may explain the observed patterns of long-term, perfect brood division. Because high predation is common and relatively predictable in the tropics, selection for fixed brood division may be stronger in tropical birds than in the temperate zone.     

The effect of helpers on artificially increased brood size in sociable weavers (Philetairus socius)

In cooperatively breeding birds, the presence of helpers is expected to increase the reproductive success of the breeding pair. However, some studies fail to find this effect. A positive effect of helpers may be restricted to cases in which a breeding pair has a poor likelihood of raising the entire brood on its own, as would be the case under stressful environmental conditions or with enlarged brood sizes. We conducted brood size manipulations in a cooperative breeder, the sociable weaver, Philetairus socius, to investigate the relationship between the difficulty of raising nestlings and the effort and impact of helpers. Overall, sociable weavers did not work harder to raise the enlarged broods. However, the presence of helpers significantly increased the feeding rates at enlarged nests, but not controls. This was insufficient to prevent generalised brood reduction in enlarged broods, whether attended by pairs alone or with helpers. Nonetheless, the presence of helpers was associated with decreased nestling mortality and an increase in the numbers of young fledged. Our results suggest that groups are better able to respond to the needs of enlarged broods than pairs alone and that the presence of helpers has a beneficial effect on overall reproductive success.Communicated by J. Dickinson     

Moorhens have an internal representation of their own eggs

How do birds recognize their own eggs? Do they have a stored template for their own egg characteristics, or do they use another mechanism? Intraspecific brood parasitism is considered to be an additional reproductive tactic where females can increase their own reproductive success. Because of the costs involved in rearing young that are not their own, it will pay females to detect and reject the eggs of a parasite, although it is not known how they do this. Here, we show experimentally that moorhens will cease laying in a nest when their first egg is replaced with another hen’s egg but not when it is replaced with their own egg taken from an earlier clutch. This provides good evidence that birds have an internal representation of their own eggs and use this in decisions about whether to reject foreign eggs.