Winter resource wealth drives delayed dispersal and family-group living in western bluebirds

Delayed dispersal, where offspring remain with parents beyond the usual period of dependence, is the typical route leading to formation of kin-based cooperative societies. The prevailing explanations for why offspring stay home are variation in resource wealth, in which offspring of wealthy parents benefit disproportionately by staying home, and nepotism, where the tendency for parents to be less aggressive and share food with offspring makes home a superior place to wait to breed. These hypotheses are not strict alternatives, as only wealthy parents have sufficient resources to share. In western bluebirds, Sialia mexicana, sons usually delay dispersal until after winter, gaining feeding advantages through maternal nepotism in a familial winter group. Experimentally reducing resource wealth (mistletoe) by half on winter territories caused sons to disperse in summer, even though their parents remained on the territory during the winter. Only 8% of sons remained with their parents on mistletoe-removal territories compared to 50% of sons on control territories (t(9,10)=3.33, p<0.005). This study is the first to demonstrate that experimentally reducing wealth of a natural food resource reduces delayed dispersal, facilitating nepotism and family-group living. The results clarify the roles of year-round residency, resource limitation and relative wealth outside the breeding season in facilitating the formation of kin-based cooperative societies.     

Begging scrambles with unequal chicks: interactions between need and competitive ability

When offspring compete for the attentions of provisioning parents, empirical and theoretical work has generally concluded that chicks honestly signal their "need" for resources and that parents control allocation. Here, we develop models to show that when allocation of food resources is determined by competitive begging scrambles between sibs, the offspring's ESS begging levels, shares of food and personal fitness gained will be determined by an interaction between their competitive abilities and their true needs. Many of the predictions of this scramble competition model are qualitatively very similar to models of honest signalling of need, where parents, not offspring, control the allocation of food. Consequently it will be difficult to distinguish between the two mechanisms of food allocation based on empirical observations of the responses of chicks to feeding by parents.     

Patterns of reproductive skew in the polygynandrous acorn woodpecker

We compared observed levels of reproductive skew in the cooperatively breeding acorn woodpecker (Melanerpes formicivorus) with those predicted by two alternative transactional models. "Concession" models predict the degree to which parentage is shared assuming that a single dominant is in complete control of reproduction. Alternatively, "restraint" models predict reproductive sharing assuming that the dominant controls only whether subordinates remain in the group but does not control its share of reproduction. Reproductive skew is high among males: on average, the most successful male sires more than three times as many offspring as the next most successful male. Females share parentage equally and have lower constraints on dispersal and lower survival rates compared with males, which is consistent with predictions from the concessions model. Also as predicted by the concessions model, yearly variation in opportunities for dispersal before the breeding season correlates positively with skew. However, in contrast to concessions but consistent with the restraint model, skew decreases with relatedness. Thus, neither model consistently predicts patterns of reproductive skew in this species. We suggest that models of reproductive skew will need to include competitive interactions among potential breeders and mate choice before they will adequately predict patterns of reproductive partitioning in most vertebrate societies.     

To share or not to share: clonal integration in a submerged macrophyte in response to light stress

The ability of clonal plant species to share resources has been studied in many experiments. The submerged macrophyte Potamogeton perfoliatus produces interconnected ramets within short time intervals and hence may or may not share resources with ramets growing in less favourable microhabitats. From a genet point of view, sharing with ramets growing under less favourable conditions might not be an optimal strategy when photosynthates could be used to establish other ramets growing under more favourable conditions. To analyse the plasticity in clonal integration of P. perfoliatus, we set up a factorial aquaria experiment with unshaded or shaded recipient ramets (offspring), which were connected to or separated from donor ramets (parents). Increased biomass production of offspring in parent–offspring systems compared with severed offspring in both light and shade showed that ramets share resources through clonal integration. The relative translocation to the first- and second-offspring generation was influenced by habitat quality: If first-offspring ramets grew in a shaded microhabitat, second-offspring ramets clearly profited. This may be at least partially because of the fact that resources are shifted from first-offspring to second-offspring ramets, indicating controlled senescence of the first-offspring. This complex sharing behaviour might be relevant when plants produce ramets within a dense patch of macrophytes, where support of a shaded ramet might not pay off.     

Sex allocation within broods: the intrabrood sharing-out hypothesis

Selection is expected to cause parents to adjust the sex oftheir offspring when the environment is predictable during development,and it is expected to affect each sex differently. When severaloffspring compete for limited resources, the environmental conditionsacting on the brood are not a good predictor of the conditionsaffecting individual offspring. There is evidence for some speciesthat, regardless of any bias in brood sex ratio, the sex ofindividual offspring within a brood may be related to its positionin the hatching/birth/weight rank, in ways that might correlatewith the expected share of available resources. Here I proposethat parents may be selected to adjust offspring sex withinthe brood, provided that some depreciable environmental qualityis unequally distributed among siblings in a predictable manner.I call this the "intrabrood sharing-out" hypothesis and presenta graphical model to derive predictions about the relationshipbetween offspring sex and positions within the brood. The modelconsiders that sibling competition not only produces differencesin the mean share of resources among siblings, but it also increasesthe predictability of the share obtained by high-ranking sibsand decreases the predictability of the share for low-rankingones. Consequently, parents should be selected to deal withsuch a distribution by promoting the conditions to make it morepredictable and then adaptively adjust the sex of particularsiblings, especially in high-ranking positions within the brood,rather than to modify the sex ratio of the brood as a whole.     

Begging and sibling competition: how should offspring respond to their rivals?

Godfray's influential model of competitive begging predicted that offspring should respond to each other's behavior, displaying more intensely when competing with needier rivals. Empirical tests of this prediction have, however, yielded equivocal results. Here, I develop a series of evolutionarily stable strategy models of begging as a signal of need, which show that this prediction holds only for competitive aspects of display that influence the division of food among the brood. No such response is expected for cooperative begging (which influences the total level of provisioning by the parent), and the models even predict the opposite trend under some circumstances (where the indirect costs of extracting additional resources from the parent are high). These contrasting sets of predictions may help to explain the varying empirical results obtained by studies of sibling interaction. Cooperative (as opposed to competitive) begging is likely to be of greater significance in cases where dominant young can gain direct control of allocation (or enjoy some competitive advantage). Dominants are then predicted to parasitize the efforts of their weaker rivals and reduce their own investment in cooperative signaling while continuing to claim a disproportionately large share of the resources provided by the parent.     

Parental nepotism enhances survival of retained offspring in the Siberian jay

The evolutionary payoff accruing to parents from breeding offspringcould be an incentive for prolonged investments in the offspring.Enhanced survival for offspring as a result of such a prolongedparental investment would increase the value of remaining inthe natal territory for the offspring. Here we show that first-yearsurvival in Siberian jays is higher in the company of theirparents. Two observations point to that the enhanced survivalof retained offspring is due to nepotistic parents rather thanto the quality of a shared habitat. First, winter survivalis higher only for those retained offspring whose parents havesurvived too ; this precludes the possibility that the linkbetween timing of dispersal and survival should reflect a higher phenotypic quality of retained offspring in general. Second,there is no support for the more parsimonious explanation thatthis link between the survival of parents and retained offspringreflects habitat quality of a shared territory. We could, withhigh statistical power, reject the possibility of a correlationbetween the survival of parental birds and unrelated immigrantsto the territory. Such a correlation would have been expectedif survival reflected habitat quality and not kinship. Our data instead suggest a direct fitness gain to retained offspringin enhanced survival through parental nepotism (parental facilitation).The behavior of parents in allowing retained offspring accessto food that is denied to immigrants is one proximate mechanismmediating a benefit of delayed dispersal.     

Fine-tuned modulation of competitive behaviour according to kinship in barn swallow nestlings

Kin selection theory predicts that, in species where progeny members compete for limiting parental care, individual offspring should be more prone to monopolize parental resources as their genetic relatedness to brood competitors decreases. Mixed parentage among broodmates may arise as a consequence, for example, of extra-pair fertilization or brood parasitism events. In this experimental study of barn swallows (Hirundo rustica), we reciprocally partially cross-fostered hatchlings between broods and compared the behaviour of pairs of related and unrelated broodmates in a competitive context, both under normal food provisioning regime and after mild food deprivation. We found that scramble competition for food mediated by visual and vocal solicitation displays (begging) is inversely related to relatedness among competitors, independent of their level of satiation. Nestlings may modulate their competitive behaviour according to vocal cues that vary with their origin and allow kin recognition. We also uncover direct fitness costs to both parents and offspring arising from mixed parentage in a brood, in terms of increased parental workload and reduced survival of the nestlings. Such previously neglected costs may select for reduced frequency of extra-pair fertilizations and brood parasitism in species with extensive parental care.     

Overproduction in the Lesser Black‐backed Gull – can marginal chicks overcome the initial handicap of hatching asynchrony?

In response to unpredictability of both food availability and core offspring failure, parents of many avian species initially produce more offspring than they commonly rear (overproduction). When parental investment is insufficient to raise the whole brood the handicap of hatching last means ‘marginal’ chicks are less likely to survive if brood reduction occurs. Conversely, if marginal offspring are required as replacements for failed ‘core’ chicks, or parental investment is sufficient to rear the whole brood, the handicap imposed on marginal chicks must be reversible if overproduction is to be a viable strategy. I investigated the ability of marginal offspring to overcome the handicap imposed by hatching asynchrony using a combination of a field experiment, designed to manipulate both the amount of total competition and the relative competitive ability of chicks within a brood, and data on the growth and survival of unmanipulated, three‐chick broods from three consecutive years. The results indicate that, even when resources are abundant, marginal offspring do not begin to overcome the competitive handicap imposed by hatching asynchrony until the period of growth when energetic requirements reach their peak, and subsequent survival to fledging is almost assured. This is apparently a consequence of parents controlling allocation of early parental investment, so that any brood reduction ‘decisions’ can be left as late as possible. Marginal chicks initially channel resources into maintaining mass, relative to skeletal size, as a buffer against starvation. However this also means competitiveness is reduced, so if conditions are poor marginal chicks are rapidly out‐competed, lose condition and die. Conversely, when food availability is good marginal offspring devote more resources to skeletal growth and quickly close the gap on their core siblings, meaning the handicap is reversible. The benefits of overproduction and hatching asynchrony as reproductive strategies to maximise success in Lesser Black‐backed Gulls are discussed in relation to the reproductive alternatives.     

Experimentally increased food resources in the natal territory promote offspring philopatry and helping in cooperatively breeding carrion crows

Kin-based societies, where families represent the basic social unit, occur in a relatively small number of vertebrate species. In the majority of avian kin societies, families form when offspring prolong their association with the parents on the natal territory. Therefore, the key to understanding the evolution of families in birds is to understand natal philopatry (i.e. the tendency to remain on the natal territory). It has been shown that, within populations, the strength of the association between parents and offspring (i.e. family stability) increases when offspring dispersal is constrained by external environmental factors, but it is unclear whether and how family wealth influences juvenile dispersal decisions. Here, we show that young carrion crows (Corvus corone corone) from territories that were food-supplemented year-round were more philopatric and more likely to help at their family's nest than the unfed ones. The results suggest that offspring philopatry and helping behaviour are influenced by the quality of 'home' and that the availability of food resources positively affects the cohesion of the family.     

Effect of dispersal and nutrient availability on the competitive ability of toxin-producing yeast

The ecological role of interference competition through toxin production is not well understood. In particular, it is unclear under what conditions the benefits of toxic killing outweigh the metabolic costs involved. A killer advantage has been suggested to rely on local competitive interactions where the benefits of killing accrue to the toxin producer preferentially, but this notion has little empirical support. In addition, contrasting predictions exist about the effect of resource abundance on the benefits of toxin production; this benefit should either be highest when resources are abundant and metabolic costs are relatively low or when resources are scarce and toxic killing is a 'last resort strategy' to obtain nutrients. Here, we test these predictions for one aspect of competitive ability, that is, the ability of toxin producers to invade a population of sensitive non-producers from a low initial frequency. We use competition experiments between isogenic K1 toxin-producing and non-producing strains of Saccharomyces cerevisiae, where we manipulate dispersal under two extreme nutrient conditions: one environment with and the other without replenishment of nutrients. We find that toxin production is beneficial when dispersal is limited under both nutrient conditions, but only when resources are abundant these outweigh its cost and allow invasion of the producer.     

The effects of ecological and genetic neighbourhood size on the evolution of two competing species

Summary Individual-based simulations were conducted to examine the effect of a small ecological neighbourhood (an area in which ecological processes such as density-dependent factors operate) and the genetic neighbourhood size (the size of an area from which the parents may be assumed to be drawn at random) on the coevolution of two competing species. For the simulations, individuals of two consumer species compete for two types of food organisms. Different genotypes (one locus and two alleles) have different efficiencies of food acquisition for different food types. Individual consumer organisms search for food within their home ranges and reproduce depending on the amount of food eaten. The dispersal distance of the offspring follows a normal distribution with a zero mean and _d standard deviation. Simulations were conducted by varying the home range size, mating area (area from where individuals choose their mates), standard deviation of dispersal distance, food generation time, the reproductive rates of food populations and the sizes and number of independent food populations. Food organisms reproduce either within one population or independently within 16 spatially divided populations. For all the simulations, competitive exclusion was the most frequent outcome and character displacement was the least frequent outcome. Through a 200-generation simulation, the two consumer species could co-exist longer and maintain a polymorphic resource use longer when the home range and mating size were small in 16 spatially divided populations than when random mating and homogeneous interaction occurred within a community (perfect mixing population). For perfect mixing populations, the frequency of character displacement increased as the food generation time became short and the reproductive rates of food decreased. It follows from the results that the sizes of the genetic and ecological neighbourhoods and the mode of resource dynamics can affect the evolution of two competing species.     

Nepotistic vigilance behavior in Siberian jay parents

Many Siberian jay offspring (up to 50%) postpone independenceand stay with their parents for up to 3 years. Parents offersuch nondispersers a benefit in that they increase their vigilancewhile feeding together with retained offspring. In contrast,parents reduce their vigilance while in company of nonrelatedflock members according to the "many eyes" principle. The preferentialtreatment offered by the parents provides an incentive for offspringto forego dispersal. Given evidence for mortality via surpriseattacks by predators (goshawks), such nepotistic vigilance byparents could have a bearing on offspring survival and therebypromote delayed dispersal.     

When it is costly to have a caring mother: food limitation erases the benefits of parental care in earwigs

The aggregation of parents with offspring is generally associated with different forms of care that improve offspring survival at potential costs to parents. Under poor environments, the limited amount of resources available can increase the level of competition among family members and consequently lead to adaptive changes in parental investment. However, it remains unclear as to what extent such changes modify offspring fitness, particularly when offspring can survive without parents such as in the European earwig, Forficula auricularia. Here, we show that under food restriction, earwig maternal presence decreased offspring survival until adulthood by 43 per cent. This effect was independent of sibling competition and was expressed after separation from the female, indicating lasting detrimental effects. The reduced benefits of maternal presence on offspring survival were not associated with higher investment in future reproduction, suggesting a condition-dependent effect of food restriction on mothers and local mother-offspring competition for food. Overall, these findings demonstrate for the first time a long-term negative effect of maternal presence on offspring survival in a species with maternal care, and highlight the importance of food availability in the early evolution of family life.     

Facultative control of offspring sex in the cooperatively breeding bell miner, Manorina melanophrys

The ability to alter primary sex ratios has the potential toincrease a breeding individual's fitness. This is certainlytrue in those cooperative breeders where one sex is both philopatricand helps raise future offspring of its parents. We examinedthe primary sex ratio variation in the cooperatively breedingbell miner (Manorina melanophrys) in southeastern Australiaover six breeding seasons. Male offspring are the philopatricand helping sex in this system and can increase the reproductiveoutput of their parents. Bell miners aggressively defend theirterritory from all interspecific competitors and by doing soallow food resources to dramatically increase. The increasein phytophagous Psyllidae insects (which secrete a carbohydrate-richcoating that constitutes the major component of bell miner diet)leads to a decrease in tree health, often culminating in deathof the tree. Bell miners then move as groups to new areas withlow psyllid abundance, and the cycle repeats. Using this predictabletemporal variation in food availability, we aimed to determinewhether female breeders adjusted the sex ratio of broods toproduce more of the philopatric sex when food resources werehigh and more of the dispersing sex when food resources werelow. Our results provide clear evidence for such facultativecontrol of sex ratio by female bell miners. Newly founded coloniesare characterized by low food availability and a female-biasedprimary sex ratio, whereas colonies more than 1 year old havean increased food availability and a male-biased primary sexratio. We suggest treating forces associated with resource enhancementand competition as opposing sides of a single general principleand suggest that it is necessary to view both the costs andbenefits of philopatric individuals within a variable environment.     

Spatial ecology of multiple parasitoids of a patchily‐distributed host: implications for species coexistence

1. The coexistence of multiple species sharing similar but spatially fragmented resources (e.g. parasitoids sharing a host species) may depend on their relative competitive and dispersal abilities, or on fine‐scale resource partitioning. Four generalist and one specialist parasitoid species associated with the holly leaf miner, Phytomyza ilicis, in a woodland network of 127 holly trees were investigated. 2. To understand coexistence and persistence of these potential competitors, patterns of occurrence in relation to patch size and isolation, vertical stratum within patches, and incidence and abundance of potential competitors were documented. Field experiments creating empty habitat patches suggested that dispersal rather than local demographic processes determines abundance and incidence. 3. Parasitoids showed species‐specific responses to patch properties, with the incidence of species determined mostly by patch size. Parasitism rates were less clearly related to patch characteristics, but parasitism rates for most species were lower in patches where the numerically dominant parasitoid species, Chrysocharis gemma, was present. No evidence of vertical stratification was found in species composition or abundance within patches, making it unlikely that coexistence is enhanced by fine‐scale resource division. 4. Overall, the patterns detected may be attributed to the distribution of C. gemma and differences in species' ecology other than dispersal ability. The life history of C. gemma may allow it to pre‐emptively exploit a large fraction of the available hosts, avoiding direct competition with other parasitoids. In contrast, direct competition is more likely among the pupal parasitoids Cyrtogaster vulgaris, Chrysocharis pubicornis, and Sphegigaster flavicornis which have a similar biology and phenology. For these species, coexistence may be facilitated by contrasting incidence in relation to patch size and isolation.     

Contribution of males to brood care can compensate for their food consumption from a shared resource

The sharing of the same food source among parents and offspring can be a driver of the evolution of family life and parental care. However, if all family members desire the same meal, competitive situations can arise, especially if resource depletion is likely. When food is shared for reproduction and the raising of offspring, parents have to decide whether they should invest in self‐maintenance or in their offspring and it is not entirely clear how these two strategies are balanced. In the burying beetle Nicrophorus vespilloides, parents care for their offspring either bi‐ or uniparentally at a vertebrate carcass as the sole food source. The question of whether biparental care in this species offers the offspring a better environment for development compared with uniparental care has been the subject of some debate. We tested the hypothesis that male contribution to biparental brood care has a beneficial effect on offspring fitness but that this effect can be masked because the male also feeds from the shared resource. We show that a mouse carcass prepared by two Nicrophorus beetles is lighter compared with a carcass prepared by a single female beetle at the start of larval hatching and provisioning. This difference in carcass mass can influence offspring fitness when food availability is limited, supporting our hypothesis. Our results provide new insights into the possible evolutionary pathway of biparental care in this species of burying beetles.     

Sibling negotiation

Current discussions of offspring begging typically assume eitherthat it is a signal directed at parents or that it representsa form of scramble competition to gain access to them. However,offspring might also display to inform nest mates that theywill contest the next food item to be delivered; in other words,begging (possibly in the absence of parents) might serve purelyas a form of negotiation among siblings. Here, we develop agame-theoretical model of this possibility. We assume that offspringvary in their need for food, which influences how intenselythey compete for access to parents. Before parental arrival,however, young may exchange signals informing each other oftheir state, and these signals may influence their subsequentcompetitive behavior. We focus on the possibility that a costlydisplay during the "negotiation" phase can serve to inform rivalsof a chick's need for resources and thereby deter them fromcompeting intensely when the parent arrives. We show that thisform of negotiation is more likely to prove stable when thefood delivered by parents is indivisible, and when it is hardfor one chick to monopolize access to resources. Investmentin negotiation (as opposed to competition) is predicted to increasewith nestling relatedness; in addition, all other things beingequal, hungrier chicks are expected to invest relatively moreeffort in displaying to their rivals, whereas weaker or smallerchicks are expected to invest less.     

Maternal investment in a spider with suicidal maternal care, Stegodyphus lineatus (Araneae, Eresidae)

Providing parental care is costly for the parent, but generally beneficial for the young whose survival, growth and reproductive value can be increased. Selection should strongly favour an optimal distribution of parental resources, depending on the relationship between the costs and benefits for parents and their offspring. Parental care is characterized by trade offs in investment, for example between egg size and number of young or providing resources at the egg stage versus the post-hatching stage. Females of the spider Stegodyphus lineatus (Eresidae) produce a single small brood with small eggs and provide the young with regurgitated fluid and later, with their body contents via matriphagy. We asked whether females adjust the investment of resources differentially into eggs, regurgitation feeding and matriphagy, and how maternal investment affects the size of the young at dispersal. We followed the growth of young of broods in the lab and in the field and manipulated brood size in order to determine the pattern of resource allocation. We found that brood size was positively correlated with body mass: larger females had larger broods. Females provided 95% of their body mass to the young, allocating more resources to regurgitation than to matriphagy. Females provided regurgitated food to the young according to the brood size, providing less food when the brood was reduced. Maternal resources had a large influence on offspring mass at dispersal, which is likely to affect their future fitness. The study shows the importance of the female's body mass and her resource allocation decisions for her reproductive outcome.     

Life history and the evolution of family living in birds

The reason why some bird species live in family groups is an important question of evolutionary biology that remains unanswered. Families arise when young delay the onset of independent reproduction and remain with their parents beyond independence. Explanations for why individuals forgo independent reproduction have hitherto focused on dispersal constraints, such as the absence of high-quality breeding openings. However, while constraints successfully explain within-population dispersal decisions, they fail as an ultimate explanation for variation in family formation across species. Most family-living species are long-lived and recent life-history studies demonstrated that a delayed onset of reproduction can be adaptive in long-lived species. Hence, delayed dispersal and reproduction might be an adaptive life-history decision rather than 'the best of a bad job'. Here, we attempt to provide a predictive framework for the evolution of families by integrating life-history theory into family formation theory. We suggest that longevity favours a delayed onset of reproduction and gives parents the opportunity of a prolonged investment in offspring, an option which is not available for short-lived species. Yet, parents should only prolong their investment in offspring if this increases offspring survival and outweighs the fitness cost that parents incur, which is only possible under ecological conditions, such as a predictable access to resources. We therefore propose that both life-history and ecological factors play a role in determining the evolution of family living across species, yet we suggest different mechanisms than those proposed by previous models.