Benefits and costs of earwig (Forficula auricularia) family life

The evolution of parental care and family group formation critically depends on offspring survival benefits and parental fecundity costs of care under given ecological conditions. Investigations of the functional significance of care in insect species that exhibit facultative parental care have been relatively rare but may be of particular interest for better understanding of benefit and cost schedules at an early evolutionary stage. In this study, aspects of benefits and costs of care were addressed in the sub-social European earwig (Forficula auricularia; Dermaptera: Forficulidae) by manipulating the presence of tending mothers and brood size in a fully crossed experimental design. Larvae growing in broods tended by their mother or of reduced size showed a higher survival probability than larvae growing in untended or large broods, as predicted if maternal care is beneficial and shaped by a trade-off between number and quality of offspring. Analysis of patterns of food consumption and developmental time further suggested that the benefit of maternal attendance is mediated by the maternal provisioning of food, while the quality–quantity trade-off seemed to be driven by sibling rivalry. Further, tending mothers delayed the production of a second clutch, indicating a potential cost of care in terms of lifetime fecundity. This study experimentally shows benefits and potential costs of maternal care and family group formation in the European earwig. More detailed behavioural experiments will be required to fully understand how behavioural interactions among family members mediate these reproductive outcomes.     

Parentally biased favouritism in relation to offspring sex in zebra finches

Family conflicts over parental care result in offspring attempting to exert control using solicitation behaviours, whilst the parents are potentially able to retaliate through provisioning rules. However, the evolutionary interests of one parent may not necessarily support the evolutionary interests of the other parent, and such conflicts of interest may be expressed in how the two parents allocate the same form of parental care to individual offspring. Theory suggests that such parentally biased favouritism is a universally predicted outcome of evolutionary conflicts of interest, and empirical evidence suggests that parentally biased favouritism occurs in relation to offspring size and solicitation behaviours. However, unequivocal empirical evidence of parentally biased favouritism in relation to offspring sex is absent, due to being strongly confounded by sex differences in size and solicitation behaviours. Here, we present strong evidence for parentally biased favouritism in relation to offspring sex in zebra finches (Taeniopygia guttata), independent of the effects of chick size and begging intensity. Mothers preferentially provisioned sons over daughters, whilst fathers showed no bias, meaning that sons received more food than daughters. Parentally biased favouritism in relation to offspring sex facilitates parental control over evolutionary conflicts of interest and is probably more widespread than previously realised.     

Rising costs of care make spiny chromis discerning parents

When the costs of parental care do not scale with the number of offspring being cared for, inclusion of non-descendant young into broods can be advantageous, leading to systems of alloparental care. However, if the cost of care scales with the number of offspring, selection may act against misdirected parental care. The spiny chromis, Acanthochromis polyacanthus, is a marine fish with extended biparental care, and broods that increase in size over the care period strongly suggest that alloparental care occurs in this species. However, A. polyacanthus parents directly provision their offspring by producing ectodermal mucus for their fry to feed on. The costs of such provisioning may scale with brood size, potentially increasing the costs of parental care. Using wild A. polyacanthus pairs, we tested whether foreign offspring are accepted into established broods, and measured how brood defence effort and mucal feeding scale with brood size. We found that A. polyacanthus discriminate between their own and foreign young, vigorously expelling experimentally introduced foreign offspring. Although defensive effort did not scale with brood size, mucal feeding was strongly dependent on brood size, and this increasing cost of care likely acts as the primary selective force on parental discrimination and rejection of foreign fry in A. polyacanthus.     

Brood reduction in response to manipulated brood sizes in the common swift (Apus apus)

Following a brood size manipulation experiment on the common swift (Apus apus) in which different levels of parental effort were created, brood reduction occurred in all five nests manipulated to four chicks and in two of the five manipulated to three young. This provided an opportunity for looking in detail at the parental investment decisions concerning allocation strategies between parent and young during the process of brood reduction. The data recorded here were analysed on a visit-by-visit basis regarding changes in parental and chick body masses, the mass of prey delivered and the estimated mass of parental self-feeding. The results show that food delivery rates did not increase in proportion to the number of chicks in the broods. This reduction in delivery rates per chick resulted in lower chick masses and ultimately in the death of one or more chicks in the larger broods. The resulting low parental body masses for adult birds feeding larger broods suggested that these parents could not have raised all their young without risking their own survival. There was a tendency for parents from nests that experienced brood reduction to feed themselves instead of allocating most of the food gathered to the young. After brood reduction, parents regained lost body mass and their young fledged at masses only slightly lower than normal. The differential allocation decisions regarding parental self-feeding which resulted in brood reduction were largely responsible for keeping the parents in good enough condition to care for the surviving nestlings. Therefore, this study clearly demonstrates that brood reduction can operate through the differential allocation of food between parent and young in a way that can have adaptive consequences for the survival of parents and their young. Correspondence to: T.L.F. Martins     

Food supply differentially affects sibling negotiation and competition in the barn owl (Tyto alba)

In contrast to most birds, nestling barn owls (Tyto alba) vocalise not only when parents are at the nest but also in their absence. Calls produced in their absence have been shown to facilitate sibling negotiation over the impending food resource. Since nestlings vocalise more vigorously in the presence of parents, they may be calling not to negotiate resources but to compete amongst each other over parental food distribution. A critical issue is to determine whether offspring need differentially affects sibling negotiation and sibling competition, that is vocalisation in the absence and presence of parents. To answer this question, I manipulated the food supply of 26 broods by adding or removing prey items. In the absence of parents, food-added broods vocalised at a significantly lower level than food-removed ones. In contrast, once a parent arrived at the nest, the vocalisation level was not significantly related to the manipulation of brood food supply. This suggests that in the absence of parents, it is more important for food-removed nestlings to vocalise intensely, and that in their presence, the relationship between begging and offspring need tends to vanish. In other words, brood food supply may affect sibling negotiation to a larger extent than sibling competition.     

Lactation performance of guinea pigs (Cavia porcellus) does not respond to experimental manipulation of pup demands

To ensure proper development of young, mothers should react to offspring signals of need. Studies of such parent-offspring interaction often manipulated litter size to measure effects of changed offspring food demand. We used the extreme precociality of guinea pig (Cavia porcellus) pups to increase offspring demand without changing any other litter characteristic. As pups contribute to their own energy demands from birth by independent feeding, their food demand can be increased by withholding solid food early in lactation. We studied whether mothers reacted to increased food demand of offspring by enhanced parental care, especially by changes in the pattern of milk production and nursing. Pups deprived of solid food early in lactation grew more slowly and were in poorer body condition than pups in control litters, even after the former had access to solid food in late lactation. Mothers of deprived young reacted to offspring long-term need by maintaining nursing behaviour for longer than control mothers. However, this change in behaviour did not occur early in lactation when pup short-term need was greatest nor did it result in increased milk transfer at any time. Energy allocation of mothers measured by changes in their food intake, maternal body reserves, and milk production stayed the same whether offspring had early access to solid food or not. Thus mothers did not increase energy allocation to pups even though they apparently had information about the pups' poor state.     

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.     

Begging signals more than just short-term need: cryptic effects of brood size in the pied flycatcher (Ficedula hypoleuca)

The begging of nestling birds is known to reliably signal short-term nutritional need, which is used by parents to adjust rates of food delivery and patterns of food distribution within broods. To test whether begging signals reflect more than just short-term feeding history, we experimentally created 18 "small" (4-nestling) and 18 "large" (8-nestling) broods in the pied flycatcher (Ficedula hypoleuca). Compared to small broods, large broods were provisioned by parents at a greater rate, but at a lower visit rate per nestling and with no obvious differences in load mass per visit. However, lower rates of food mass delivery per nestling in large broods did not result in any measurable reduction in nestling growth (i.e. "long-term need") or in any increase in the begging effort per individual nestling whilst in the nest. Mid-way through the nestling period we also used hand-feeding laboratory trials to assess in more detail individual begging behaviour and digestive performance of the three mid-ranking nestlings from each brood. More food items were required at the start of each trial to satiate nestlings from large broods, but despite this initial control for "short-term need", nestlings from large broods went on to beg at consistently higher rates and at different acoustic frequencies. Large brood nestlings also produced smaller faecal sacs, which were quantitatively different in content but did not differ in frequency. We suggest that different nutritional histories can produce cryptic changes in nestling digestive function, and that these can lead to important differences in begging signals despite controlling for short term need.     

Parental feeding rate in relation to begging behavior in asynchronously hatched broods of the great tit Parus major

Summary Variations in the begging behaviour of the nestlings of altricial birds can provide the parents with information about the nestlings' nutritional needs and thus influence the parental feeding rate. In a series of four experiments, the stimulus situation encountered by great tits on their feeding visits to the brood was manipulated to explore its effect on feeding rate.A higher feeding rate was observed under the following conditions: (1) after a period of food deprivation, as compared with both normal conditions and satiation through artificial feeding; (2) in periods when recorded begging calls were played during feeding visits, as compared with control periods; (3) after temporary removal from the nest of heavier, as compared with lighter, siblings. The lighter nestlings in the brood benefitted more —in terms of gain in weight — from the increase in parental feeding rate following the playing of begging calls than did the heavier nestlings. Differences in weight spread within broods did not affect the amount of food the parents brought.We conclude that parental feeding rate is affected not simply by the begging of the hungriest nestling, but rather by the behaviour of all the nestlings in the brood, which makes possible an adjustment of the feeding rate to the average hunger level of the brood. The effects of hatching asynchrony and sibling competition on parental feeding rate are discussed.     

Asymmetric sibling rivalry and nestling growth in red-winged blackbirds (Agelaius phoeniceus)

Many birds hatch their offspring asynchronously, and the adaptive significance of this trait, if any, is controversial. David Lack suggested long ago that by facilitating brood reduction when resources are scarce, hatching asynchrony provides relief from the effects of overcrowding. Some field workers interpret this to mean that the growth and survival of survivors should rise following partial brood loss. Here we show in a 6-year study of red-winged blackbirds (Agelaius phoeniceus) that the presence or absence of marginal offspring in experimentally manipulated broods had virtually no effect upon the growth of core offspring, whereas alterations of the size of core brood had strong and significant effects. Nestling growth was, not surprisingly, slower in broods with partial brood loss. Intriguingly, marginal offspring showed significantly greater variation in mass. Core offspring are less sensitive to, but not exempt from, the inimical effects of resource shortfall than are marginal offspring. The phenotypic handicap appears to marginal offspring a caste of high-variance progeny whose fitness prospects rest upon levels of parental input (stochastic resources) and the size of the core brood (stochastic development). Received: 21 June 1999 / Revised: 5 June 2000 / Accepted: 25 June 2000     

Common goldeneyes adjust maternal effort in relation to prior brood success and not current brood size

Parental investment theory predicts that parental effort should be related to the reproductive value of the current brood. This depends on both the number of young and the survival prospects of each of them. Thus parents may provide more care to larger broods either because of (1) the direct effect of brood size per se on reproductive value (the “brood size” hypothesis) or because (2) past mortality, reflected in current brood size, predicts future mortality of the brood and hence its reproductive value (the “brood success” hypothesis). Earlier studies have not attempted to distinguish between these alternatives. We tested the hypotheses in the precocial, nidifugous common goldeneye Bucephala clangula, a species with uniparental female care. Maternal effort was measured as the time spent by the female in rearing the brood. We found that brood size itself is not associated with maternal effort, but that females modify their maternal effort according to the mortality already experienced by the brood, supporting the prediction of the brood success hypothesis. We also found that brood mortality varied considerably between broods and that previous mortality predicts future mortality within broods, basic assumptions of the brood success hypothesis. Received: 30 January 1996 / Accepted after revision: 27 October 1996     

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     

Influence of behavior and mating success on brood-specific contribution to fish recruitment in ponds

One source of uncertainty in predicting the response of populations to exploitation is individual differences within a population in both vulnerability to capture and contribution to population renewal. For species with parental care, individuals engaged in nesting behavior are often targeted for exploitation, but predicting outcomes of this nonrandom vulnerability will depend in part on an understanding of how parental traits are related to potential for brood contribution to the population. Variation in brood-specific contribution to recruitment of largemouth bass (Micropterus salmoides), a fish species with extended parental care, was quantified to determine if differences in mating success, parental care behaviors, and timing of reproduction influenced offspring recruitment. Dependence of these relationships on brood predation was tested in communities that differed in the presence of bluegill, Lepomis macrochirus, an important nest predator. Daily snorkel surveys were conducted in experimental ponds during spring to monitor male spawning and parental care behaviors in populations of largemouth bass. Tissue samples collected from larvae in nests were used to develop brood-specific DNA fingerprints for determining nest origins of fall recruits. Largemouth bass spawning period in bluegill ponds was longer and more variable in duration, with lower, more variable mating success, than in ponds without bluegill. In all populations, only one or two broods provided the majority of recruits, and these were broods produced during the earliest days of spawning by the oldest, largest males. In bluegill ponds, brood contribution from earliest nests also increased with brood size. Earliest nesters were the oldest males, and recruits from these nests were often above average in body size. Offspring needed to be guarded to at least swim-up larval stage to contribute any recruits. Termination of parental protection before offspring were free swimming mainly occurred with broods guarded by smaller males in ponds with brood predators. These age- and size-specific differences in timing of spawning and duration of parental care are consistent with influences of residual reproductive value and energetic constraints on reproductive behavior. Furthermore, these patterns of individual contribution to recruitment imply that fisheries that selectively target either nesting individuals or larger, older males could potentially decrease recruitment at the population scale.     

Food supplementation affects egg albumen content and body size asymmetry among yellow-legged gull siblings

Parental decisions can determine offspring experience of environmental conditions. Such ‘maternal’ effects act both before and after hatching via, e.g., egg quality or the social milieu predisposed by parents. Resource availability may constrain the expression of adaptive maternal effects, and the specific pattern of allocation of these effects among offspring depending on their sex or birth order can result in different fitness payoffs to parents. Declining egg mass with laying order observed in several bird species may constitute an adaptive strategy of parental favouritism towards early hatching offspring with larger reproductive value but may also result from nutritional constraints on laying effort. A previous study has shown that the small size of the third, last laid (c-)egg in yellow-legged gull (Larus michahellis) clutches depends on food availability and that food-supplemented mothers increase the size of their female but not male c-eggs. Here, we show that increased mass of c-eggs laid by females supplemented with food after clutch initiation depends on increased albumen mass, which, in turn, enhances the size of daughters at hatching. Because asynchronous hatching results in a competitive disadvantage of c-chicks, present results suggest that mothers relieved from nutritional constraints enhance the size of daughters to compensate for their larger susceptibility to hatching last. The study also confirms the role of egg albumen content in determining hatchling size, previously experimentally detected only in one species in the wild. The effect of increased egg mass on offspring size persisted at least until day 8 after hatching, when, however, it did not vary with sex, suggesting intense negative selection on small female c-chicks in control broods. Hence, maternal effects mediated by egg albumen content had persistent effects on offspring size.     

The dynamics of brood desertion among communally breeding females in the treehopper, Publilia concava

In species that exhibit extended parental care, females sometimes lay eggs communally in order to redistribute the costs of offspring care. Communal egg-laying often involves redundancy in female effort, such that the number of females contributing to reproduction is more than is needed to provide adequate parental care. As a result, a subset of females will often abandon the communal brood, with the time of departure ranging from immediate abandonment after egg-laying (brood parasitism) to delayed abandonment with prolonged care (cooperative breeding). In this paper I approach the parental care dynamics of female-female broods as a desertion game similar to that of mate desertion in species with bi-parental care. I describe a field study of the insect Publilia concava (Hemiptera: Membracidae), a species that exhibits communal oviposition and a full range of egg guarding. This species exhibits full redundancy in female care, with no difference in survival between singly and doubly guarded broods. I find that double guarding is extremely rare in the population, with most communal broods having only one female guard. While this guard was usually the female that initiated the brood, these same females were more likely to abandon when secondary females arrived and when secondary females exhibited longer guarding durations. Paradoxically, the secondary females usually abandoned the broods they visited, resulting in up to 50% of broods with double abandonment. These unguarded broods suffered a 50% reduction in hatching success, reflecting an important risk for primary females that abandon egg masses to secondary females. Overall, P. concava exhibits desertion dynamics similar to mate desertion in vertebrates and it is likely that the theoretical work in this area will be useful for future work that addresses the allocation of parental care among communal breeders.     

Parentage and paternal care: consequences of intersexual selection in Savannah sparrows?

Empirical relationships between parentage and male parental care are commonly interpreted in the context of life-history models that consider increased offspring survivorship as the only benefit of paternal effort. However, indirect benefits associated with male care can also influence a male's response to cuckoldry: if females allocate paternity according to their prior experience with male parental care, it may pay for males to provision extra-pair young in early broods. Here, I assess the relationship between first-brood parentage and paternal care in a population of Savannah sparrows (Passerculussandwichensis) where a male's fertilization success in the second brood appears to be influenced by his prior parental performance. Based on the multi-locus DNA fingerprinting of 17 first broods, male feeding effort was influenced by parentage (percent of brood resulting from within-pair fertilizations) but not by brood size, male mating status (monogamous versus polygynous), timing of breeding (hatching date), structural size (wing length) or condition (mass). Males provided more care to broods that contained few within-pair young. This result supports the idea that males provision young to increase their future mating success, but alternative hypotheses involving male quality and timing of breeding cannot be excluded. Received: 13 August 1996 / Accepted after revision: 22 February 1997     

Food-dependent benefits of hatching asynchrony in American kestrels Falco sparverius

Food supply and hatching asynchrony were manipulated for 90 broods of American kestrels (Falco sparverius) during 1989–1991. We measured the growth and mortality of nestlings within four treatment groups (asynchronous, synchronous, food-supplemented, unsupplemented) to test the brood reduction hypothesis of Lack (1947, 1954). Fledging success did not differ between synchronous and asynchronous broods when food was poor but consistent with the brood reduction hypothesis, nestlings died at a younger age in asynchronous broods. When food was supplemented, mortality did not occur in the synchronous broods but youngest nestlings still died in asynchronous nests despite apparently adequate food for the brood. Oldest nestlings in asynchronous broods fledged with a greater mass than their younger siblings, also consistent with Lack's hypothesis. Average nestling quality in synchronous broods was very dependent on food levels. Synchronous young that were supplemented were, on average, the heaviest of any treatment group but young from unsupplemented synchronous broods were the lightest. Overall, patterns of mortality and growth for kestrels support the brood reduction hypothesis when food is limited, but not when it is abundant. This food-dependent benefit of asynchrony in the nestling period is a prerequisite for facultatively adjusted hatching spans during laying.     

Extra-pair paternity and tail ornamentation in the barn swallow Hirundo rustica

Females of socially monogamous species may copulate with attractive non-mates to obtain access to the genes of such males, and a preference for attractive copulation partners may result in sexual selection. Extra-pair copulations are common in the socially monogamous barn swallow Hirundorustica, and a 2-year study of paternity using multi-locus DNA fingerprinting demonstrated that 33% of 63 broods and 28% of 261 offspring were sired by extra-pair males. The frequency of extra-pair offspring within broods was highly skewed with the majority of all broods having either no extra-pair offspring or only extra-pair offspring. Individual pairs were consistent in their frequency of extra-pair paternity among broods, and the repeatability of extra-pair paternity of multiple broods of the same female was statistically significant. The proportion of extra-pair offspring was negatively related to the tail length of the male attending the nest. Behavioural observations showed that extra-pair fertilizations were more likely in broods raised by females that had been observed to engage in extra-pair copulations. The frequency of extra-pair offspring was unrelated to the intensity of two male paternity guards, mate guarding and the rate of intra-pair copulations. In an analysis of extra-pair paternity and male parental care in different broods of the same male, male barn swallows fed their offspring relatively less frequently if the brood contained more extra-pair offspring. Therefore, female barn swallows pursue extra-pair copulations with attractive males, which may result in sexual selection, even though extra-pair paternity is costly for females due to the reduction of paternal care by their social mates. Received: 24 January 1997 / Accepted after revision: 2 August 1997     

Hatching asynchrony reduces the duration, not the magnitude, of peak load in breeding green-rumped parrotlets (Forpus passerinus)

The peak load reduction hypothesis suggests that hatching asynchrony in altricial birds is adaptive because it reduces parental workload during the most energetically costly time in brood rearing. By staggering the ages of their offspring, parents may ensure that all nestlings do not reach maximum energy demand simultaneously. To test the hypothesis, we used the doubly labeled water technique to measure the energy expenditure of green-rumped parrotlets (Forpus passerinus) that reared experimentally manipulated synchronous and asynchronous broods. Peak metabolic rates of the two experimental groups did not differ, but parents of asynchronous broods metabolized significantly less energy than did parents of synchronous broods throughout the first half of the brood-rearing period. Our results suggest that hatching asynchrony in parrotlets substantially shortens the temporal duration of high brood energy demand, but does not reduce the magnitude of peak energy demand. Received: 16 July 1998 / Accepted after revision: 13 December 1998     

Resource allocation varies with parental sex and brood size in the asynchronously hatching green-rumped parrotlet (<Emphasis Type="Italic">Forpus passerinus</Emphasis>)

When eggs hatch asynchronously, offspring arising from last-hatched eggs often exhibit a competitive disadvantage compared with their older, larger nestmates. Strong sibling competition might result in a pattern of resource allocation favoring larger nestlings, but active food allocation towards smaller offspring may compensate for the negative effects of asynchronous hatching. We examined patterns of resource allocation by green-rumped parrotlet parents to small and large broods under control and food-supplemented conditions. There was no difference between parents and among brood sizes in visit rate or number of feeds delivered, although females spent marginally more time in the nest than males. Both male and female parents preferentially fed offspring that had a higher begging effort than the remainder of the brood. Mean begging levels did not differ between small and large broods, but smaller offspring begged more than their older nestmates in large broods. Male parents fed small offspring less often in both brood sizes. Female parents fed offspring evenly in small broods, while in large broods they fed smaller offspring more frequently, with the exception of the very last hatched individual. These data suggest male parrotlets exhibit a feeding preference for larger offspring—possibly arising from the outcome of sibling competition—but that females practice active food allocation, particularly in larger brood sizes. These differential patterns of resource allocation between the sexes are consistent with other studies of parrots and may reflect some level of female compensation for the limitations imposed on smaller offspring by hatching asynchrony.