Individual variation in parental tradeoffs between the number and size of offspring at the pre- and post-natal stages

Life-history theory predicts that parents refer to the resources they hold to determine their breeding strategy. In multi-brooded species, it is hypothesized that single-brooded parents produce larger clutches and raise offspring with a brood survival strategy, whereas multi-brooded parents only do this under good breeding conditions. Under poor conditions, they produce smaller clutches and raise offspring with a brood reduction strategy. We tested this hypothesis in the Brown-cheeked Laughing Thrush Trochalopteron henrici, which can breed twice a year on the Tibetan Plateau, by investigating the life-history traits and provisioning behaviours of single- and double-brooded parents. Single-brooded parents laid larger clutches of smaller eggs and produced more and larger fledglings than double-brooded parents in their first brood. Double-brooded parents produced smaller clutches of larger eggs but fledged larger nestlings in their first brood than in their second brood. As single-brooded parents only need to raise one brood a year, then producing and raising as many offspring as possible (i.e. the brood survival strategy in a large brood) can maximize their reproductive success. For double-brooded parents, producing and raising fewer offspring in the first brood (i.e. the brood survival strategy in a small brood) can ensure their nesting success during a short breeding cycle. Additionally, producing more offspring but raising larger nestlings in the second brood (i.e. the brood reduction strategy in a large brood) can select for offspring of higher quality within the brood. Our findings indicate that different tradeoffs between single- and double-brooded parents in egg-laying and nestling-raising may be an adaptation to the seasonal variation in environmental conditions.     

Experimental dissociation of individual quality, food and timing of breeding effects on double-brooding in a migratory songbird

Annual reproductive success in many species is influenced by the number of breeding attempts within a season. Although previous studies have shown isolated effects of female quality, food, and timing of breeding on the probability of female birds producing second broods, to our knowledge, none have tested the relative importance of multiple factors and their interactions using simultaneous manipulations within populations of free-living birds. In this study, we show that individual quality and timing of breeding interact to affect the probability of double-brooding in female mountain bluebirds (Sialia currucoides). High-quality females (those that naturally initiated clutches early in the season) were more likely to double-brood, regardless of whether their hatching date was advanced or delayed, whereas later breeding, lower quality females were much less likely to double-brood when their first attempt was delayed. This indicates that annual fecundity of poorer quality (or younger) female bluebirds may be more sensitive to seasonal variation in environmental conditions. In addition, birds that were provided with supplemental food throughout first breeding attempts were more likely to double-brood in one of the study years, suggesting that female bluebirds may be energetically limited in their capacity to initiate a second brood. Females that had their first brood delayed also had a shorter inter-brood interval and were moulting fewer feathers during second broods compared to controls, while females in better condition showed more advanced moult in second breeding attempts. Taken together, our results demonstrate the combined effects of age- or individual quality-mediated energetic trade-offs between current and future reproduction, and between investments in offspring and self-maintenance, on annual fecundity of female birds.     

Brood provisioning rates and fledgling behavior of Cordilleran Flycatchers in southwestern Colorado

The behavior of young songbirds after fledging is one of the least understood phases of the breeding cycle, although parental provisioning rates and movement of fledglings are key to understanding life history evolution. We studied Cordilleran Flycatchers (Empidonax occidentalis) at two sites in southwestern Colorado, USA, from 2012 to 2017. We banded and sexed breeding adults to determine the relative contributions of males and females to nestling and fledgling care, and attached radio‐transmitters to nestlings to facilitate observations of brood behavior after fledging. Females made 60% and 78% of total observed feedings of nestlings and fledglings, respectively. Parental provisioning rates increased with nestling age, and per‐nestling provisioning rates increased with brood size. Parental provisioning rates declined just before fledging, then increased just after fledging. Fledging times of individuals in broods were asynchronous and concentrated during the late afternoon and early evening. Males stopped caring for fledglings before females even though this species is single‐brooded, with some late‐season broods being abandoned by males. Broods spent the first three weeks after fledging within 400 m of nests, after which they began to disperse. Most aspects of the breeding biology of Cordilleran Flycatchers in our study, including the duration of nestling and fledging periods, female‐dominated provisioning, and movement patterns of fledglings, were similar to those of other Empidonax species. However, the times when young fledged were not concentrated in the morning as reported in most other songbirds, and this result warrants additional study of the timing of fledging in ecologically and taxonomically similar species. The increased per‐nestling provisioning rate with increasing brood size was unexpected, and additional study is needed to determine if this increase results from a trade‐off between adult annual survival and productivity favoring increased provisioning of young in larger broods, or from the existence of high‐quality individuals where larger clutches and higher provisioning rates are linked.     

Azure‐winged Magpies Cyanopica cyanus trade off reproductive success and parental care by establishing a size hierarchy among nestlings

It is common in birds that the sizes of nestlings vary greatly when multiple young are produced in one nest. However, the methods used by parents to establish size hierarchy among nestlings and their effect on parental provisioning pattern may differ between species. In the Azure‐winged Magpie Cyanopica cyanus, we explored how and why parents controlled the sizes of nestlings. Asynchronous hatching was the main cause of size hierarchy within the brood, although the laying of larger eggs later in the laying sequence reduced this effect. Parents with asynchronous broods produced more eggs and fledged more nestlings than those with synchronous broods but their brood provisioning rates, food delivery per feeding bout and feeding efficiency did not differ. We performed a cross‐fostering experiment to synchronize some asynchronous broods. Provisioning rates of asynchronous broods were lower than those of synchronized broods, but the daily growth rates and fledging body mass of their nestlings were not different. Our findings indicate that parents of asynchronous broods can achieve higher reproductive success than those of synchronous broods based on the same parental care, and the same reproductive success as those of synchronized broods based on less parental care. It appears that parent birds can better trade off reproductive success and parental care by establishing a size hierarchy among nestlings.     

Factors affecting the probability of double brooding by Southern House Wrens

Seasonal fecundity of birds is influenced by clutch sizes and the number of successful breeding attempts during a breeding season. As such, understanding the factors that determine the decision to initiate multiple broods within a season and the consequences of this reproductive tactic is important. We examined the frequency of double brooding by Southern House Wrens (Troglodytes aedon musculus) in eastern Argentina. We analyzed inter‐ and intraseasonal variation in double brooding and evaluated the effect of weather conditions and laying date on the frequency and occurrence of this behavior. Finally, we assessed the effect of double brooding on the seasonal and lifetime productivity of female Southern House Wrens. During our 8‐year study, we found that ～43% (range = 17–83% each year) of breeding pairs attempted a second brood after successfully raising a first brood. The probability of females having a second brood was affected by the laying date of the first nesting attempt, but was independent of the number of young fledged. About 65% of females that started laying eggs before the first quarter of each breeding season produced a second brood, and this percentage decreased to ～40% after this period. In addition, variation in double‐brooding frequency among years was related to weather conditions, with the proportion of pairs double brooding increasing with increased precipitation early in the breeding season. More precipitation likely contributed to an increase in insect abundance. Although double brooding increased the seasonal and lifetime productivity of female Southern House Wrens, additional study of the survival and fate of fledglings from first and second broods is needed to assess the importance of multi‐brooding in the reproductive success of these wrens.     

Anthropogenically-Mediated Density Dependence in a Declining Farmland Bird

Land management intrinsically influences the distribution of animals and can consequently alter the potential for density-dependent processes to act within populations. For declining species, high densities of breeding territories are typically considered to represent productive populations. However, as density-dependent effects of food limitation or predator pressure may occur (especially when species are dependent upon separate nesting and foraging habitats), high territory density may limit per-capita productivity. Here, we use a declining but widespread European farmland bird, the yellowhammer Emberiza citrinella L., as a model system to test whether higher territory densities result in lower fledging success, parental provisioning rates or nestling growth rates compared to lower densities. Organic landscapes held higher territory densities, but nests on organic farms fledged fewer nestlings, translating to a 5 times higher rate of population shrinkage on organic farms compared to conventional. In addition, when parental provisioning behaviour was not restricted by predation risk (i.e., at times of low corvid activity), nestling provisioning rates were higher at lower territory densities, resulting in a much greater increase in nestling mass in low density areas, suggesting that food limitation occurred at high densities. These findings in turn suggest an ecological trap, whereby preferred nesting habitat does not provide sufficient food for rearing nestlings at high population density, creating a population sink. Habitat management for farmland birds should focus not simply on creating a high nesting density, but also on ensuring heterogeneous habitats to provide food resources in close proximity to nesting birds, even if this occurs through potentially restricting overall nest density but increasing population-level breeding success.     

The Process and Causes of Fledging in a Cavity-Nesting Passerine Bird, the House Wren (Troglodytes aedon)

Little is known about the process or causes of fledging or nest‐leaving in passerine birds because researchers can rarely predict when fledging will occur in a given nest. We used continuous videotaping of nests to both document the process of fledging in the house wren, Troglodytes aedon, a small, cavity‐nesting songbird, and test hypotheses as to what might cause fledging to begin. Fledging began any time from 14 to 19 d after hatching commenced. Slower‐developing broods fledged later than faster‐developing broods. Fledging typically began within 5 h of sunrise and over 80% of all nestlings fledged before noon. All nestlings fledged on the same day at 65% of nests and over two consecutive days in most other nests. We found no evidence that fledging was triggered by changes in parental behaviour. Parental rate of food delivery to nestlings did not decline during a 3‐h period leading up to the first fledging, nor was the rate of feeding just prior to the first fledging lower than the rate at the same time the day before. Moreover, parents did not slow the rate of food delivery to nests after part of the brood had fledged. Hatching is asynchronous in our study population which creates a marked age/size hierarchy within broods. At most nests, the first nestling to fledge was the most well‐developed nestling in the brood or nearly so (as measured by feather length). This suggests that fledging typically begins when the most well‐developed nestlings in the brood reach some threshold size. However, at about one‐fifth of nests, the first nestling to fledge was only moderate in size. At these nests, severe competition for food may have caused smaller, less competitive nestlings to fledge first to increase their access to food. We found no strong support for the suggestion that the oldest nestlings delay fledging until their least‐developed nestmate reaches some minimum size, although further experimental work on this question is warranted.     

Horned larks on the Tibetan Plateau adjust the breeding strategy according to the seasonal changes in the risk of nest predation and food availability

Songbirds in seasonal environments often adjust their breeding strategy according to spatial or temporal changes in breeding conditions. Here we investigate how horned larks Eremophila alpestris, a multi‐brooded songbird on the Tibetan Plateau, responded to the changing risk of nest predation and food availability across breeding attempts. We showed that both nest concealment and food supply increased with plant growth, and horned larks adjusted their breeding strategies accordingly. First they selected nest‐sites where predator density was low, which enhanced nest survival. Second, clutch size increased with improving breeding conditions. They did not adopt an ‘egg‐size’ strategy as egg size did not change with laying sequence or breeding attempt. Instead, they adopted the ‘brood survival (feeding later‐hatched nestlings more)’ and ‘brood reduction (feeding early‐hatched nestlings more)’ strategies during early and later attempts. Moreover, nestlings’ growth varied with breeding attempt: more energy was invested into the growth of body mass during the first attempt but more energy was expended on the growth of linear structures during later attempts. This difference in energy allocation reflected changing food availability. We suggest that temporal changes of environmental factors are also the important force driving the evolution of avian breeding strategies.     

Male parental care promotes early fledging in an open-nester, the Willow Warbler Phylloscopus trochilus

The importance of male parental care to female reproductive success was investigated in the monogamous Willow Warbler Phylloscopus trochilus by removing the male parent at two different stages of the breeding cycle. Females that were widowed at the start of egg-laying continued breeding and managed to raise their brood on their own with no apparent reductions in numbers fledged or fledgling body-mass. The widowed females compensated for the loss of male assistance by increasing their own food provisioning rate as compared with control females. However, widows spent less time brooding the small young, and the growth rate of nestlings was reduced. In nests where the male parent was removed 7 days after the eggs hatched, the subsequent growth rate of nestlings was still affected, which suggests that male care is influential throughout the nestling period. On average, broods reared by widows fledged 2 days later than did broods of control females. An extension of the nestling period may appreciably affect reproductive success, since 68% of nests failed due to predation, mostly during the nestling period. We suggest that the main role of male parental care in the Willow Warbler is to assure a high growth rate of nestlings, which leads to early fledging and hence a reduced risk of nest predation.     

A more informative method for analyzing reproductive success

ABSTRACT.   Few investigators have examined how a female's prior history (i.e., number of clutches laid previously and whether those nesting attempts were successful or not) might influence the success of a particular clutch. Our objective was to determine the seasonal change in the mean number of fledglings successfully leaving nests in a population of Prairie Warblers ( Dendroica discolor ). To do so, we calculated the success of each clutch in the laying sequence (i.e., first, second, third, …, n ^th) and sorted them further by whether they were laid before (first-brood clutches) or after (second-brood clutches) the first brood fledged. In our sample population of 70 females we knew the fate of all clutches laid during a breeding season. Although the number of fledglings per successful clutch varied slightly, the probability that a clutch would be successful (i.e., produce at least one fledgling) and the number of fledglings per nest attempt increased during the breeding season. Thus, later clutches were more productive than earlier clutches. The low probability of producing a successful first clutch early in the breeding season would seem to favor selection for females that wait until later in the season to begin breeding. However, the seasonal reproductive success of females may be more dependent on breeding early (increasing the number of clutches laid in a season) than on the higher probability of success with later clutches. Our results indicate that further study of seasonal changes in reproductive success and their causes is needed.     

Cascading costs of reproduction in female house wrens induced to lay larger clutches

In many species, females produce fewer offspring than they are capable of rearing, possibly because increases in current reproductive effort come at the expense of a female's own survival and future reproduction. To test this, we induced female house wrens (Troglodytes aedon) to lay more eggs than they normally would and assessed the potential costs of increasing cumulative investment in the three main components of the avian breeding cycle – egg laying, incubation and nestling provisioning. Females with increased clutch sizes reared more offspring in the first brood than controls, but fledged a lower proportion of nestlings. Moreover, nestlings of experimental females were lighter than those of control females as brood size and prefledging mass were negatively correlated. In second broods of the season, when females were not manipulated, experimental females laid the same number of eggs as controls, but experienced an intraseasonal cost through reduced hatchling survival and a lower number of young fledged. Offspring of control and experimental females were equally likely to recruit to the breeding population, although control females produced more recruits per egg laid. The reproductive success of recruits from broods of experimental and control females did not differ. The manipulation also induced interseasonal costs to future reproduction, as experimental females had lower fecundity than controls when breeding at least 2 years after having their reproductive effort experimentally increased. Finally, females producing the modal clutch size of seven eggs in their first broods had the highest lifetime number of fledglings.     

Parental care trade‐offs in the inter‐brood phase in Barn Swallows Hirundo rustica

In seasonal environments with limited time and energy resources, double‐brooded birds face trade‐offs in the timing of their two reproductive attempts and in the effort allocated to the first and the second broods. In the Barn Swallow Hirundo rustica a long care period for the first brood enhances the survival of first‐brood chicks, but also delays the start of the second brood, which in turn reduces the survival prospects of second‐brood chicks. Probably as a response to this trade‐off, double‐brooded Barn Swallows reduce the period of post‐fledging care for first‐brood fledglings. By radiotracking whole families, we investigated the determinants of this behaviour and its consequences for the survival of the first‐brood fledglings. The end of the females’ investment in post‐fledging care of the first brood was related to the beginning of egg synthesis for the second clutch. With the start of egg synthesis, females significantly reduced provisioning rates to the first‐brood fledglings to less than one‐fifth of the previous rates, while the proportion of time they spent foraging remained high. Assuming that the females’ foraging success was constant, we conclude that their energy income was allocated to egg production rather than fledgling provision. Males did not compensate for the females’ reduced feeding rates. Thus the start of egg production for the second clutch had a marked effect on the quantity of food received by first‐brood fledglings. In parallel with the changes in parental behaviour and provisioning rates, we observed a marked drop in the daily survival rate of first‐brood chicks. These results support the hypothesis that females face a strong trade‐off in the allocation of energy to subsequent broods. Energy allocation to a second clutch involves a cost in terms of reduced provisioning, and as a result the survival of first‐brood chicks is compromised. This is probably outweighed by the improved success of an early second brood.     

Factors determining the frequency and productivity of double brooding of Barn Owls Tyto alba

Capsule: Early nesting Barn Owls Tyto alba and those that switched nest sites fledged most chicks overall because they could fit two, more productive, nesting attempts into a breeding season.

Aims: To determine the frequency and productivity of double broods in Barn Owls, and for double brooders, to determine what affects the probability of nest switching and how that affects productivity.

Methods: We monitored the first egg date of each nesting attempt, whether it was in a ‘vole year’, whether a breeding attempt was first or a second annual attempt, the number of chicks fledged from each attempt, and whether a pair switched nest sites, if breeding twice, from 602 Barn Owl breeding attempts in an area of lowland England from 1996 to 2007. General linear models were used to determine predictors of the probability that a pair had a second brood and the number of chicks fledged in each nesting attempt, and then for those owls that double brooded, which variables best predicted the probability of switching, and the number of chicks fledged from the second nest. Finally, we tested whether switching resulted in a shorter laying interval and higher annual productivity.

Results: Early nesting birds were more likely to double brood, although this was relaxed in vole years when later nesting birds also double brooded. Productivity (through increased numbers of chicks fledged or reduced chick loss) was higher the earlier a nest occurred, and there were more chicks fledged in good vole years and in second nesting attempts. Productivity, brood depletion, first clutch date and vole years did not determine whether a double brooding pair switched nesting sites. Productivity in the second nest did not change with a switch but productivity increased for early first nests and second nests with a shorter interval between the first and second nest. Switching however decreased nesting interval and nesting interval was also shorter if there were fewer fledglings from the first nest. Overall productivity was higher for pairs that switched.

Conclusions: Double brooding in Barn Owls increased seasonal productivity substantially and its occurrence depended on vole abundance or early nesting. Nest switching between broods may be a strategy for earlier laying of the second brood. Provision of alternative nest sites, close together in a Barn Owl’s home range, may allow earlier re-nesting and so increase productivity.     

Females compensate for moult‐associated male nest desertion in Hooded Warblers

Uniparental offspring desertion occurs in a wide variety of avian taxa and usually reflects sexual conflict over parental care. In many species, desertion yields immediate reproductive benefits for deserters if they can re‐mate and breed again during the same nesting season; in such cases desertion may be selectively advantageous even if it significantly reduces the fitness of the current brood. However, in many other species, parents desert late‐season offspring when opportunities to re‐nest are absent. In these cases, any reproductive benefits of desertion are delayed, and desertion is unlikely to be advantageous unless the deserted parent can compensate for the loss of its partner and minimize costs to the current brood. We tested this parental compensation hypothesis in Hooded Warblers Setophaga citrina, a species in which males regularly desert late‐season nestlings and fledglings during moult. Females from deserted nests effectively doubled their provisioning efforts, and nestlings from deserted nests received just as much food, gained mass at the same rate, and were no more likely to die from either complete nest predation or brood reduction as young from biparental nests. The female provisioning response, however, was significantly related to nestling age; females undercompensated for male desertion when the nestlings were young, but overcompensated as nestlings approached fledging age, probably because of time constraints that brooding imposed on females with young nestlings. Overall, our results indicate that female Hooded Warblers completely compensate for male moult‐associated nest desertion, and that deserting males pay no reproductive cost for desertion, at least up to the point of fledging. Along with other studies, our findings support the general conclusion that late‐season offspring desertion is likely to evolve only when parental compensation by the deserted partner can minimize costs to the current brood.     

Paternal care and male mate-attraction effort in the European starling is adjusted to clutch size

In facultative polygynous birds with biparental care, a trade-off may occur between male parental care and attraction of additional mates. If there is a cost associated with reduced male parental care, the relative benefit of mate attraction may be predicted to decrease as the size of a male's clutch or brood increases. We tested this prediction in monogamous pairs of facultatively polygynous European starlings (Sturnus vulgaris). The larger the clutch, the more time the male spent incubating and the less time he spent attracting an additional female (i.e. singing near and carrying green nesting material into adjacent empty nest-boxes). Reduced paternal incubation resulted in lower overall incubation (the female did not compensate) and lower hatching success. Immediately after experimental reduction of clutches, males spent significantly less time incubating and more time singing and carrying greenery, and vice versa for experimentally enlarged clutches. Males with experimentally reduced clutches attracted a second female more often than males with experimentally enlarged clutches. This is the first study, to our knowledge, to provide experimental evidence for an adjustment of paternal care and male mate-attraction effort to clutch size. However, a trade-off between paternal nestling provisioning and mate attraction was not revealed, probably due to the absence of unpaired females by that time in the breeding season. Experiments showed that the relative contribution of the male and female to nestling provisioning was unrelated to brood size.     

Predators as stressors? Physiological and reproductive consequences of predation risk in tropical stonechats (Saxicola torquata axillaris).

Tropical birds usually lay smaller clutches and are less likely to initiate a second brood than their temperate-zone relatives. This reduction in annual fecundity is generally explained as an adaptation either to higher rates of nest predation or to a more limited food supply concurrent with higher adult survival in the tropics. However, the physiological parameters associated with lower annual fecundity in tropical birds have not been well investigated. We compared the annual fecundity, behaviour and a number of physiological parameters of stonechat parents feeding fledged juveniles in territories with and without fiscal shrikes, a predator on adult and fledged birds. Stonechat pairs in territories with shrikes were less likely to initiate a second brood and delayed successive broods compared to pairs in territories without shrikes. After fledging of their young, males showed a greater propensity than females to initiate distraction calls after a human intrusion into their territory and, therefore, invested more in the defence of their young. In territories with shrikes stonechat males had higher initial plasma corticosterone levels and lower body conditions than males in territories without shrikes, suggesting that they were chronically stressed. In contrast, the females from both types of territory had low initial plasma corticosterone levels. We conclude that shrike presence might account for the delay in initiation of a second brood and the reduction in the tendency to initiate a second brood. Whether these effects are mediated by the elevated levels of corticosterone remains to be demonstrated.     

Male‐skewed adult sex ratio,survival, mating opportunity and annual productivity in the Snowy Plover Charadrius alexandrinus

Sex differences in adult mortality may be responsible for male‐skewed adult sex ratios and male‐skewed parental care in some birds. Because a surplus of breeding males has been reported in serially polyandrous populations of Snowy Plover Charadrius alexandrinus, we examined sex ratio, early‐season nesting opportunities, adult survival and annual reproductive success of a Snowy Plover population at Monterey Bay, California. We tested the hypotheses that male adult survival was greater than female survival and that a sex difference in adult survival led to a skewed adult sex ratio, different mating opportunities and different annual productivity between the sexes. Virtually all females left chicks from their first broods to the care of the male and re‐nested with a new mate. As a result, females had time to parent three successful nesting attempts during the lengthy breeding season, whereas males had time for only two successful attempts. Among years, the median population of nesting Plovers was 96 males and 84 females (median difference = 9), resulting in one extra male per eight pairs. The number of potential breeders without mates during the early nesting period each year was higher in males than in females. Adult male survival (0.734 ± 0.028 se) was higher than female survival (0.693 ± 0.030 se) in top‐ranked models. Annually, females parented more successful clutches and fledged more chicks than their first mates of the season. Our results suggest that in C. alexandrinus a sex difference in adult survival results in a male‐skewed sex ratio, which creates more nesting opportunities and greater annual productivity for females than for males.     

Food availability and the male's role in parental care in double-brooded Treecreepers Certhia familiaris

The aim of this work was to examine differences in paternal and maternal care in a double-brooded, monogamous species, the Treecreeper Certhia familiaris, in relation to food availability. As a measure of parental care, we recorded the hourly feeding activity of parents when the nestlings from their first and second breeding attempts were 7 and 12 days old. Feeding frequency of the first brood increased with the age of the nestlings and also with the brood size when 12 days old. While the feeding activities of the females were similar with respect to the first and second broods, the males were less active and failed to provide any food to their nestlings in 15 cases out of 28 second broods. In spite of this, the fledglings from the second broods were heavier than those in the first. Such a pattern of male behaviour was possible without being a disadvantage to the chicks because the food supply increased during the breeding season and the female could provide food for the young alone. Thus paternal care was particularly important in times of poor food supply, i.e. during the first brood, where the extent of these males' activity in feeding the 7-day-old nestlings was positively correlated with the average mass of the nestlings. Our results support the idea that the male of monogamous, altricial bird species often makes important contributions to raising the young, especially during periods when it is difficult for the female to do so alone. Males show flexibility in their pattern of parental care, and male Treecreepers change their contribution to the first and second broods within the same season.     

Parental provisioning behaviour in a flock-living passerine, the Vinous-throated Parrotbill Paradoxornis webbianus

The amount of food delivered by parents to their chicks is affected by various life history traits as well as environmental and social factors, and this investment ultimately determines the current and future fitness of parents and their offspring. We studied parental provisioning behaviour in the Vinous-throated Parrotbill Paradoxornis webbianus, a species with an unusual social system that is characterised by flock-living, weak territoriality and variable nesting dispersion. Parental provisioning rate had a positive influence on chick mass gain, suggesting that provisioning rate is an effective measure of parental investment in this species. Males and females fed nestlings at approximately the same rate, and no other carers were observed at nests. Parents coordinated provisioning rates so that they mostly fed chicks synchronously. However, the extent to which parents coordinated provisioning was associated with their social environment, synchrony being positively related to local breeding density and negatively to nearest neighbour distance. The rate at which parents provisioned nestlings showed the same relationships with social measures, being greatest at higher density and when neighbours were closer. Visit rate was also related to chick age, but not to brood size, brood sex ratio, extra pair paternity, laying date, temperature, parents’ body characters, time of day or year. We conclude that a breeding pairs’ social environment plays an important role in determining parental investment, probably through its effects on the opportunities that parents have for foraging with conspecifics.     

Age-dependent reproductive costs and the role of breeding skills in the Collared flycatcher

This study addressed whether there are any age‐related differences in reproductive costs. Of especial interest was whether young individuals increased their reproductive effort, and thereby their reproductive cost, as much as older birds when brood size was enlarged. To address these questions, a brood‐size manipulation experiment with reciprocal cross‐fostering of nestlings of young and middle‐aged female Collared flycatchers, Ficedula albicollis, was performed on the Swedish island of Gotland. Nestlings’ body mass, tarsus length and survival were recorded to estimate the parental ability and parental effort of the experimental female birds. Female survival and clutch size were recorded in the following years to estimate reproductive costs. We found that middle‐aged female flycatchers coped better with enlarged broods than younger females or invested more in reproduction. In the following year, young female birds that had raised enlarged broods laid smaller clutches than the females from all the other experimental groups. This result shows that the young female birds pay higher reproductive costs than the middle‐aged females. Both young and middle‐aged female flycatchers seemed to increase their reproductive effort when brood size was increased. However, such an increase resulted in higher reproductive costs for the young females. The difference in reproductive costs between birds of different ages is most likely a result of insufficient breeding skills of the young individuals.