Sibling Competition Research Articles

Associations between adolescent siblings' relationship quality and similarity and differences in values.

Theoretically framed by a self-determination perspective on value acquisition and taking into account research into sibling similarity and differences, the current study examined links between sibling relationship quality and adolescents' intrinsic values (benevolence and universalism) and extrinsic values (power, achievement, and materialism). Positive sibling experiences were expected to be positively linked to intrinsic values and negatively linked to extrinsic values, and negative experiences were hypothesized to fuel extrinsic values and thwart intrinsic values. Using a sample of 205 adolescent sibling pairs (older children M = 17.61 years, younger children M = 14.63 years) and multilevel modeling, we assessed within- and between-dyads differences to identify whether the sibling relationship functions as a correlate of values in a similar or different way for 2 siblings. Although siblings were not very similar in their values, sibling competition predicted sibling similarity in higher levels of extrinsic values and lower levels of intrinsic values. Implications for value acquisition research that so far has focused almost solely on parents and failed to acknowledge other processes within families are discussed.

Evolution of parental care driven by mutual reinforcement of parental food provisioning and sibling competition

In mammals, altricial birds and some invertebrates, parents care for their offspring by providing them with food and protection until independence. Although parental food provisioning is often essential for offspring survival and growth, very little is known about the conditions favouring the evolutionary innovation of this key component of care. Here, we develop a mathematical model for the evolution of parental food provisioning. We find that this evolutionary innovation is favoured when the efficiency of parental food provisioning is high relative to the efficiency of offspring self-feeding and/or parental guarding. We also explore the coevolution between food provisioning and other components of parental care, as well as offspring behaviour. We find that the evolution of food provisioning prompts evolutionary changes in other components of care by allowing parents to choose safer nest sites, and that it promotes the evolution of sibling competition, which in turn further drives the evolution of parental food provisioning. This mutual reinforcement of parental care and sibling competition suggests that evolution of parental food provisioning should show a unidirectional trend from no parental food provisioning to full parental food provisioning.

Sudden weather deterioration but not brood size affects baseline corticosterone levels in nestling Alpine swifts

While evidence is accumulating that stress-induced glucocorticoid responses help organisms to quickly adjust their physiology and behaviour to life-threatening environmental perturbations, the function and the ecological factors inducing variation in baseline glucocorticoid levels remain poorly understood. In this study we investigated the effects of brood size by experimentally manipulating the number of nestlings per brood and the effect of weather condition on baseline corticosterone levels of nestling Alpine swifts ( Apus melba). We also examined the potential negative consequences of an elevation of baseline corticosterone on nestling immunity by correlating corticosterone levels with ectoparasite intensity and the antibody production towards a vaccine. Although nestlings reared in enlarged broods were in poorer condition than nestlings reared in reduced broods, they showed similar baseline corticosterone levels. In contrast, nestling baseline corticosterone levels were higher immediately after cold and rainy episodes with strong winds. Neither nestling infestation rate by ectoparastic flies nor nestling antibody production against a vaccine was correlated with baseline corticosterone levels. Thus, our results suggest that altricial Alpine swift nestlings can quickly modulate baseline corticosterone levels in response to unpredictable variations in meteorological perturbation but not to brood size which may be associated with the degree of sibling competition. Apparently, short-term elevations of baseline corticosterone have no negative effects on nestling immunocompetence.

Kin competition, natal dispersal and the moulding of senescence by natural selection

Most theoretical models for the evolution of senescence have assumed a very large, well mixed population. Here, we investigate how limited dispersal and kin competition might influence the evolution of ageing by deriving indicators of the force of selection, similar to Hamilton (Hamilton 1966 J. Theor. Biol. 12, 12-45). Our analytical model describes how the strength of selection on survival and fecundity changes with age in a patchy population, where adults are territorial and a fraction of juveniles disperse between territories. Both parent-offspring competition and sib competition then affect selection on age-specific life-history traits. Kin competition reduces the strength of selection on survival. Mutations increasing mortality in some age classes can even be favoured by selection, but only when fecundity deteriorates rapidly with age. Population structure arising from limited dispersal however selects for a broader distribution of reproduction over the lifetime, potentially slowing down reproductive senescence. The antagonistic effects of limited dispersal on age schedules of fecundity and mortality cast doubts on the generality of conditions allowing the evolution of 'suicide genes' that increase mortality rates without other direct pleiotropic effects. More generally, our model illustrates how limited dispersal and social interactions can indirectly produce patterns of antagonistic pleiotropy affecting vital rates at different ages.

Egg size and yolk steroids vary across the laying order in cockatiel clutches: A strategy for reinforcing brood hierarchies?

When a female bird begins incubation before clutch completion, the nestlings hatch sequentially, and a size hierarchy forms within the brood. This size hierarchy may be minimized or exacerbated through differential allocation of resources to eggs across the laying order. In this study, we characterize intra-clutch variation in cockatiel clutches by measuring egg mass, yolk mass, and concentrations of yolk testosterone, androstenedione, and corticosterone. Cockatiels are a long-lived member of the Psittaciformes. Because asynchronous hatching may reduce sibling competition and allow for extended development periods in long-lived birds, we predicted that female cockatiels would allocate maternal resources in a way that would reinforce the brood size hierarchy. Significant within-clutch differences in egg size and steroid concentrations were observed. Eggs at the end of the laying sequence were smaller and had significantly smaller yolks than eggs early in the laying order. Fifth-laid eggs, as well as first-laid eggs, contained significantly lower concentrations of testosterone than eggs in other positions of the laying sequence. No differences in yolk androstenedione concentration were observed. Yolk corticosterone concentrations increased linearly with laying order. Together, these patterns might reinforce the brood size hierarchy created by asynchronous hatching.

Maternal effects on offspring size and packaging constraints in the whelk

Life-history theory predicts an optimal offspring size, irrespective of reproductive effort; however, in some species offspring size correlates positively with maternal size. We examine hypotheses for why this latter situation should occur in the whelk Buccinum undatum. The trade-offs between aspects of reproduction in whelks are complicated due to the provision of protective egg capsules. Many eggs are placed within each capsule but c. 99% of these eggs are consumed by the remaining developing young. Large maternal size results in more eggs, larger eggs, more eggs consumed per hatchling, more capsules, larger capsules, more eggs per capsule, a larger number of hatchlings per capsule and larger hatchlings. Increased intra-capsule and post-hatch sibling competition may decrease the marginal value for additional young and select for larger young, however, our data do not support this explanation. Instead, packaging constraints within each capsule limit the size of hatchlings but this constraint is relaxed for medium to large females because they produce large capsules. Small females appear to produce young below optimum size because of the space constraint thus explaining the correlation between maternal size and offspring size.

Sibling Competition and Conspicuousness of Nestling Gapes in Altricial Birds: A Comparative Study

BackgroundNestlings of altricial birds capture parents' attention through conspicuous visual displays, including exposure of their gape coloration which informs parents about their level of need, competitive ability or health; information that parents use for deciding food allocation among their offspring. Thus, because nestlings compete with nest mates for parental care, nestling conspicuousness is expected to increase with level of sibling competition along bird phylogeny.Methodology/Principal FindingsWe test this prediction by jointly using information of brood reduction, clutch size and duration of nestling period as proxies for intensity of sibling competition, and visual models that assess detectability of nestlings by adult birds. As predicted, we found a positive association between nestling conspicuousness and intensity of brood reduction, while clutch size and duration of nestling period did not enter in the best models. Level of brood reduction was positively related with the achromatic component of nestling conspicuousness and body mass was negatively related with the chromatic component.ConclusionsThese associations are in agreement with the hypothesis that sibling competition for parental attention has driven the evolution of visual nestling conspicuousness in a context of parent-offspring communication in altricial birds.

The evolution of sex-specific grandparental harm

Recent empirical studies indicate that grandparents favour some categories of grandchildren over others. Here, we expand the previous theoretical foundation for this finding and show that grandchild-harming phenotypes are predicted to evolve by 'sexually antagonistic zygotic drive (SA-zygotic drive) of the sex chromosomes'. We use the logic of Hamilton's rule to develop a new 'no-cost-to-self nepotism rule' that greatly simplifies the determination of the invasion criteria for mutations that cause grandparents to harm grandchildren. We use this theory to generate predictions that distinguish SA-zygotic drive from theory based solely on paternity assurance. The major diagnostic prediction is that grandmothers, and to a lesser degree grandfathers, will evolve grandson-harming phenotypes that reduce the level of sib competition experienced by their more closely related granddaughters, especially in their sons' families. This prediction is supported by data from recent studies showing (i) grandmothers invest more in granddaughters than grandsons, and counterintuitively, (ii) paternal grandmothers reduce the survival of their grandsons. We conclude that SA-zygotic drive is plausibly operating in humans via sexually antagonistic grandparental care.

Egg size and asymmetric sibling rivalry in red-winged blackbirds

How big to make an egg is a life history decision that in birds is made coincident with a series of other similar decisions (how many eggs to have, whether to fortify them with maternally derived hormones or immune system boosters, whether to hatch the eggs synchronously or asynchronously). Though within-population variation in egg size in birds has been well studied, its adaptive significance, if any, is unclear. Here we examine within-population variation in egg size in relation to asymmetric sibling rivalry in a 17-year study of red-winged blackbirds (Agelaius phoeniceus), an altricial songbird. Egg mass showed a twofold range of variation, with roughly 80% of the variation occurring across clutches. By commencing incubation before the clutch is complete, mothers create advantaged core and disadvantaged marginal elements within their brood. Previous work on this system has shown that sibling competition is asymmetric, and that core offspring enjoy priority access to food, and as a consequence show higher growth and lower mortality than marginal offspring. Here we examine the effect of initial egg size on nestling growth and survival in relation to these competitive asymmetries. Egg mass was strongly linked to hatchling mass, and remained significantly related to the mass of both core and marginal nestlings; the effect of egg size was stronger for core offspring early in the nestling period, but the disparity between core and marginal nestlings narrowed as they approached fledging age, and slower growing marginals fell victim to brood reduction. The effect of egg mass on survival differed dramatically between core and marginal nestlings. Egg mass was significantly related to the survival of marginal but not core nestlings: below average egg mass was associated primarily with very early mortality. Asymmetric sibling competition is clearly a strong determinant of the consequences of egg size variation.

Developmental plasticity varied with sex and position in hatching hierarchy in nestlings of the asynchronous European roller, Coracias garrulus

Allocation rules between ornamental and other functional traits of birds may differ among individuals and vary with environmental conditions. We supplemented roller (Coracias garrulus) nestlings with methionine in a between-nest design to investigate the way in which the sex and position in the hatching hierarchy affect the allocation of resources among growth, immunity, and plumage coloration. Methionine induces the production of lymphocytes at expense of growth; thus, we used it to manipulate growth and immunity, which are two traits likely to compromise plumage coloration. We predicted that late-hatched chicks within a brood (juniors) compared to early-hatched chicks (seniors) should allocate more to traits directly providing fitness than to ornamental traits because juniors are more affected than seniors by sibling competition. The methionine treatment effectively enhanced the production of lymphocytes in experimental broods. This appeared to be at the expense of plumage coloration in junior nestlings because, in supplemented nests, junior males showed a trend to display less greenish bellies than junior males from control nests. However, juniors from supplemented nests maintained wing growth as in control juniors. The plumage coloration of seniors was unaffected by the methionine supplementation, although they paid the costs of lymphocyte production at a level of growth that was reduced compared to senior nestlings in control nests. Hence, sex, and hatching order affected resource allocation among growth, immunity, and plumage coloration of roller nestlings. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99, 500–511.

Cooperation and the evolution of anisogamy

In the lights of the concept of cooperation wholes, I discuss why the differentiation of sperm and ova can occur with a mathematical model. Most of Parker's explanations for anisogamy are not completely proper, because it is proved that sperm competition is neither sufficient nor necessary for anisogamy and cooperation to deal with fertilization risks is the real key to understand the evolution of anisogamy. According to the computer simulation results, the transport of gametes between different individuals, risks of the transport, the consequent inequality of sperm and eggs and competition among different individuals were the main causes of gamete differentiation. But these factors have different roles and effects. The transport risk is the main reason for individuals of different mating types to cooperate and differentiate into sperm and egg producers. The transported gametes have an advantage to evolve into sperm to seek for a larger gamete number over the fixed gametes, because they suffer more risks as they can encounter the same fixed gamete and less sibling competition as they can be dispersed better. Gamete competition among different individuals just causes the transported gametes to become as small as possible if they have already become smaller beyond a critical state. In the final discussion, I further put the evolution of anisogamy into a broader background of levels of selection and of the evolution of cooperation, the most important existential mode of matters that makes life as life.

Are acoustical parameters of begging call elements of thin-billed prions related to chick condition?

Chicks of burrowing petrels use begging calls to advertise their hunger levels when parents arrived at the nest. In a previous study, adult thin-billed prions Pachyptila belcheri responded to higher begging call rates of their single chick by regurgitating larger meals. We tested whether acoustic parameters of begging call elements may also be involved in signalling. To describe variation in begging, we determined begging session parameters, namely the duration, number of calls and the mean and maximum rate of calling. We then digitised calls and carried out a semi-automatic extraction of six acoustic parameters of call elements, including mean and maximum acoustic frequency, the length of call elements and the location of the maximum frequency and amplitude within calls. Chicks showed strong individual differences in all parameters. While the session parameters were correlated with body condition and with the meal size the chick received, none of the acoustic parameters were related to body condition and provisioning. A cross-fostering experiment showed the same pattern, as only session parameters changed related to an experimentally altered body condition, while acoustical cues appear to play no role in signalling hunger levels. We suggest that this may be explained by the absence of sibling competition in these birds. As parents do not need to decide which chick to feed, immediate information on condition at the time of adult arrival may not be required.

Group provisioning limits sharing conflict among nestlings in joint-nesting Taiwan yuhinas

Offspring often compete over limited available resources. Such sibling competition may be detrimental to parents both because it entails wasted expenditure and because it allows stronger offspring to obtain a disproportionate share of resources. We studied nestling conflict over food and its resolution in a joint-nesting species of bird, the Taiwan yuhina (Yuhina brunneiceps). We show that adult yuhinas coordinate their feeding visits, and that this coordination limits competition among nestlings, leading to a 'fairer' division of resources. Transponder identification and video-recording systems were used to observe adult feeding and nestling begging behaviours. We found that: (i) yuhinas feed nestlings more often in large parties than in small parties; (ii) feeding events occurred non-randomly in bouts of very short intervals; and (iii) food distribution among nestlings was more evenly distributed, and fewer nestlings begged, during large-party feeding bouts compared with small-party feeding bouts. To our knowledge, this is the first study in a cooperative breeding species showing that adults can influence food allocation and competition among nestlings by coordinating their feeding visits. Our results confirm the hypothesis that the monopolizability of food affects the intensity of sibling competition, and highlight the importance of understanding the temporal strategies of food delivery.

Effect of sibling competition and male carotenoid supply on offspring condition and oxidative stress

Early developmental conditions have major implications for an individual’s fitness. In species where offspring are born simultaneously, the level of sibling competition for food access is intense. In birds, high sibling competition may subject nestlings to decreased growth rate as a result of limited food and increased levels of oxidative stress through high metabolic activity induced by begging behaviors. We manipulated the level of sibling competition in a natural population of great tits and assessed the consequences for nestling body condition and resistance to oxidative stress. In a full factorial design, we both augmented brood size to increase sibling competition and supplemented the male parents with physiological doses of carotenoids thereby doubling the natural carotenoid intake, aiming at increasing the males’ investment in current reproduction and thereby decreasing sibling competition. Nestling body mass was reduced by the brood enlargement and enhanced by the carotenoid supplementation of fathers. Nestling resistance to oxidative stress, measured as total antioxidant defenses in whole blood, was not influenced by the treatments. Because nestlings experience high metabolic activities, an absence of an effect of sibling competition on free radicals production seems unlikely. Nestling body mass decreased and resistance to oxidative stress tended to increase with initial brood size, and hence these correlational effects suggest a trade-off between morphological growth and development of the antioxidant system. However, the result of the experimental treatment did not support this trade-off hypothesis. Alternatively, it suggests that nestling developed compensatory mechanisms that were not detected by our antioxidant capacity measure. Key words: antioxidant capacity, body condition, brood size manipulation, carotenoid supplementation, Parus major, sibling competition. [Behav Ecol]

Reaching independence: food supply, parent quality, and offspring phenotypic characters in kestrels

The duration of the postfledging dependence period (PFDP) is supposed to be modulated by the parent‐offspring conflict: Offspring should extract the highest levels of parental investment, although parents may respond by setting fixed limits to the level of investment or by raising the costs of attempts to extract additional investment. In this context, longer PFDPs are expected in nests tended by higher quality parents and in dominant siblings. We explored these hypotheses with a combination of experimental and correlative results. First, we food supplemented offspring during the PFDP to study whether food supply during fledging, an indicator of parental quality, has an effect on the PFDP duration in the Eurasian kestrel. We found that the PFDP was longer in food-supplemented nests. Second, we measured the duration of the PFDP over 3 years under different environmental conditions to explore whether the quality of parents and nestling phenotype were correlated with the duration of PFDP. Correlative results suggest that fledglings raised by higher quality parents and in the year with poorer food conditions showed longer PFDP. Furthermore, male fledglings showing grayer coloration in the rump (an index of competitive capacity) have longer PFDPs than browner males. Overall, our results suggest that parent, rather than offspring characteristics, can modulate the PFDP duration mediated by food conditions, although more colored nestlings stay in the nest territory for longer periods. Key words: environment conditions, Eurasian kestrel, Falco tinnunculus, melanic coloration, parent‐offspring conflict, sibling competition. [Behav Ecol]

Early inbreeding depression in the sexually polymorphic plant Dianthus sylvestris (Caryophyllaceae): Effects of selfing and biparental inbreeding among sex morphs

Predominantly outcrossing plant species are expected to accumulate recessive deleterious mutations, which can be purged when in a homozygous state following selfing. Individuals may vary in their genetic load because of different selfing histories, which could lead to differences in inbreeding depression among families. Lineage-dependent inbreeding depression can appear in gynodioecious species if obligatory outcrossed females are more likely to produce female offspring and if partially selfing hermaphrodites are more likely to produce hermaphrodites. We investigated inbreeding depression at the zygote, seed, and germination stages in the gynomonoecious-gynodioecious Dianthus sylvestris, including pure-sexed plants and a mixed morph. We performed hand-pollinations on 56 plants, belonging to the three morphs, each receiving 2-3 cross treatments (out-, sib- and self-pollination) on multiple flowers. Effects of cross treatments varied among stages and influenced seed provisioning, with sibling competition mainly occurring within outcrossed fruits. We found significant inbreeding depression for seed mass and germination and cumulative early inbreeding depression varied greatly among families. Among sex morphs, we found that females and hermaphrodites differed in biparental inbreeding depression, whereas uniparental was similar for all. Significant inbreeding depression levels may play a role in female maintenance in this species, and individual variation in association with sex-lineages proclivity is discussed.

Competition for space can drive the evolution of dormancy in a temporally invariant environment

I present a model for the evolution of a seed bank in the absence of externally driven environmental variation. I use Evolutionarily Stable Strategy (ESS) analyses of both analytic and simulation models to assess the conditions under which a dormant genotype can invade and resist invasion. In my models, plant seeds compete through lottery for discrete safe sites holding one individual each. Analyzing the conditions under which a dormant genotype can invade when rare and resist invasion once established, I conclude that dormancy can be an ESS when some fraction of seeds is retained locally, seed bank survival is high, and mortality in the seed bank is low. The advantage of dormancy stems from the ability of dormant seeds to recapture a lost site and the fact that a plant’s offspring are more likely to win the lottery in its own site than in any new site. The advantage of dormancy does not depend on individual fecundity or on low relatedness with the offspring of kin, making this mechanism distinct from earlier models of sib competition.

A polyandrous female moth discriminates against previous mates to gain genetic diversity

Polyandry is common in many animal species. The hypotheses proposed to explain the evolution of polyandry generally fall into two categories: to gain material benefits and to obtain genetic benefits. Here we report our findings on a polyandrous moth, Ephestia kuehniella (Lepidoptera: Pyralidae). The present study does not support the hypothesis that females gain material benefits from multiple copulations in terms of sperm replenishment, fecundity and longevity, because females that copulated once and females that copulated more than once with the same or different males had similar fertility, fecundity and longevity. Females used three strategies to gain possible genetic benefits: discriminating against previous mates probably via chemical cues, giving 60% chance of paternity to second males, and adjusting their oviposition patterns depending on whether they encountered new or previous mates after the first copulation. For the third strategy, females that encountered new mates after the first copulation saved eggs until the next day to facilitate fertilization of their eggs with the sperm of their new mates. Our findings do not support the hypothesis that polyandry increases offspring viability, because polyandry did not affect egg hatching success, offspring survival or weight. Ephestia kuehniella showed limited ability to disperse, and females laid all their eggs locally, mostly within 2 days, suggesting that offspring live together and that sib copulations are common in this species. Therefore, polyandry is probably a mechanism for reducing sib competition and inbreeding costs in offspring.

Male feeding rate and extrapair paternity in the facultatively polygynous spotless starling

Male birds often face a loss of paternity in the broods they attend because of promiscuous behaviour by their mates. Parental investment theory predicts that caring for the offspring of rival males incurs costs, selecting against males that care for unrelated offspring. We examined the relationship between male parental care and paternity in the facultatively polygynous spotless starling, Sturnus unicolor, during three consecutive breeding seasons. We found no evidence that male spotless starlings reduce their feeding rates to their nestlings in relation to their loss of paternity. Unexpectedly, cuckolded males provisioned nestlings more than noncuckolded males in relation to the presence of extrapair offspring. Although this result deserves further investigation, we argue that spotless starling males do not reduce their feeding rate when they lose paternity mostly because variance in male feeding rate is not very costly in a species in which polygamy is related to male fitness, the occurrence of extrapair offspring is unpredictable and reliable cues to assess paternity are unlikely. Several hypotheses are presented to interpret an increase in feeding rates of cuckolded males over non-cuckolded males in terms of parental quality with feeding rates either envisaged as a form of mating effort or related to kin selection and depending on begging level, sibling competition and genetic parentage within broods.

A Natural Experiment: Heterospecific Cross-fostering of House Wrens (Troglodytes aedon) by Tree Swallows (Tachycineta bicolor)

Abstract We report a natural cross-fostering experiment of two House Wren (Troglodytes aedon) eggs and nestlings by Tree Swallows (Tachycineta bicolor). We document an increase in duration of the House Wren incubation period by 1–2 d, which closely corresponds to the mean incubation period of their hosts. We also documented nestling growth rates of both the host Tree Swallows and the alloparented House Wrens. The host parents apparently fed the House Wren nestlings; however, the wrens exhibited slow growth compared with conspecific nestlings in neighboring nests. The wren nestlings eventually died at 6 and 13 d of age. We hypothesize that the lower incubation temperatures, and later, interspecific sibling competition that the House Wren nestlings were exposed to in the Tree Swallow nest may have prolonged the incubation period and slowed the growth rates of the House Wren nestlings.