Do Pied Flycatchers Use Personal or Social Information for Replacement Clutch Decisions? A Field Experiment

Life is a continuous stream of decisions: organisms are repeatedly confronted with multiple options from which to choose. To make the best decisions and maximise their fitness, individuals need to collect information: from direct interactions with the environment (personal information) or from observing other individuals interacting with the environment (social information). Yet, very little is known about the details of decision‐making by animals in their natural environments, including the sources of information they use in the process. We set out to determine the relative importance of social and personal information in breeding habitat decisions in the migratory pied flycatcher, Ficedula hypoleuca. In a field experiment in Finland, we manipulated flycatcher breeding success and then allowed individuals to choose among breeding sites labelled with artificial cues (arbitrary symbols) associated with their own (manipulated) and neighbour's reproductive success, respectively, for laying a replacement clutch. Birds did not appear to distinguish among the symbols representing their own, their neighbour's or a neutral symbol, indicating that they did not use either information source when deciding about a replacement clutch. Also, the choice did not influence subsequent investment into breeding (clutch size). This may suggest that pied flycatchers utilise other cues (e.g. breeding success or predation risk on larger scales) when having to make a decision about replacement clutches. We discuss our results in light of constraints imposed by a short breeding season of migratory birds.     

Prospecting in the collared flycatcher: gathering public information for future breeding habitat selection?

Visiting breeding patches can allow prospecting individuals to gather information on local patch quality to make optimal decisions about selecting a breeding habitat in the following year. This prospecting behaviour has been described in many bird species. However, the nature of the information gathered by prospectors often remains unknown. We collected data on prospecting behaviour in the collared flycatcher, Ficedula albicollis, a small hole-nesting passerine bird, to investigate whether prospectors could gather information on their conspecifics' reproductive success, that is, ‘public information’. If they could, they would be expected to inspect conspecific nests, prospect at the time when public information is reliable, and be attracted to, and spend time at, successful sites. Prospecting at conspecific nests was frequent. Prospectors were mainly males, and prospecting activity closely matched the peak of nesting activity. The probability of observing a prospector at a nest increased with parental activity, measured by feeding rate and vigilance behaviour, but did not depend on direct measures of success of the inspected nest (nestling number and condition), or on female characteristics. Because feeding rate and vigilance behaviour were predictors of breeding success at fledging, prospectors were attracted to the most successful nests when cueing on conspicuous parental activity. Therefore, prospectors could gather accurate information on local conspecific reproductive success that may be used for breeding patch choice in the following year. We discuss alternative explanations for our results, and the need to test experimentally whether prospectors gather public information. We also discuss the role of breeding constraints on, and the origin of sex differences in, prospecting.     

Manipulating Individual Decisions and Environmental Conditions Reveal Individual Quality in Decision-Making and Non-Lethal Costs of Predation Risk

Habitat selection is a crucial decision for any organism. Selecting a high quality site will positively impact survival and reproductive output. Predation risk is an important component of habitat quality that is known to impact reproductive success and individual condition. However, separating the breeding consequences of decision-making of wild animals from individual quality is difficult. Individuals face reproductive decisions that often vary with quality such that low quality individuals invest less. This reduced reproductive performance could appear a cost of increased risk but may simply reflect lower quality. Thus, teasing apart the effects of individual quality and the effect of predation risk is vital to understand the physiological and reproductive costs of predation risk alone on breeding animals. In this study we alter the actual territory location decisions of pied flycatchers by moving active nests relative to breeding sparrowhawks, the main predators of adult flycatchers. We experimentally measure the non-lethal effects of predation on adults and offspring while controlling for effects of parental quality, individual territory choice and initiation of breeding. We found that chicks from high predation risk nests (<50 m of hawk) were significantly smaller than chicks from low risk nests (>200 m from hawk). However, in contrast to correlative results, females in manipulated high risk nests did not suffer decreased body condition or increased stress response (HSP60 and HSP70). Our results suggest that territory location decisions relative to breeding avian predators cause spatial gradients in individual quality. Small adjustments in territory location decisions have crucial consequences and our results confirm non-lethal costs of predation risk that were expressed in terms of smaller offspring produced. However, females did not show costs in physiological condition which suggests that part of the costs incurred by adults exposed to predation risk are quality determined.     

The effect of migration strategy and food availability on White Stork Ciconia ciconia breeding success

In the mid 1970s, the breeding populations of the migrant White Stork Ciconia ciconia were close to extinction in the northeastern region of France (Alsace). A re-introduction project was implemented, resulting in the year-round settlement of some individuals in the region, which rely on additional food supplied by humans during the winter. Today, both resident and migrant birds breed in the same areas and take food from rubbish dumps and humans (farmers). The effects of these anthropogenic influences, altering Stork behaviour, on Stork reproductive success are not known. The aim of this study was to test the influence of bird status (resident vs. migrant) and food availability (control nests vs. nests that benefit from high food supply) on reproductive success. In control nests, the mean laying date was earlier in resident than in migrant White Storks. There was also a clear seasonal decline in clutch size. For all nests, the numbers of eggs and hatchlings were higher in resident birds than in migrants, which can be attributed to the earlier breeding of resident Storks. The large broods of resident birds showed a high mortality rate, leading to the same fledgling success (fledglings/hatchlings) and number of fledglings as in migrants. Fledgling success and the number of fledglings were higher for nests close to a reliable food supply. In summary, although resident birds can breed earlier and produce more eggs than migrants, we found no advantage in terms of number of fledglings. The higher mortality rate of chicks found in pairs with a large brood could be caused by the deterioration of their habitat. Thus, the year-round settlement of Storks may not present a biological advantage if the quality of their habitat is not guaranteed by the conservation of their grasslands.     

Within-season divorce in Blue Tits (Cyanistes caeruleus)

Divorce between breeding seasons, i.e. mate change while the old breeding partner is still alive, has been well-studied in several bird species. It can be viewed either as the result of reproductive decisions of individuals to maximize their own fitness or as the outcome of intra-sexual competition for mating opportunities. In contrast, divorce within a breeding season—also referred to as rapid mate switching—has received much less attention. Within-season divorce may occur after sudden changes in environmental conditions, such as those causing the disappearance or devaluation of nesting sites or territories, or when better breeding partners become available. Within-season divorce can be initiated by a member of the pair, or it can be the result of a take-over by an unpaired individual that competes for access to the resource. During a field experiment investigating the effects of limiting nesting sites on reproductive behaviour in Blue Tits, we recorded several cases of within-season divorce. The rate of divorce was not related to the experimental nest-site limitation, and pairs that changed their partner suffered reduced reproductive success compared to faithful pairs. Although there were no differences in the timing of breeding, clutch size or hatching success, pairs with a new partner also suffered a reduced fledging success, which was partly explained by complete brood failures. This study highlights that the pair bond prior to egg laying can be unstable when conditions force individuals to compete for a new partner or nest site and indicates the importance of the correct timing of divorce within the breeding cycle.     

Annual and seasonal reproductive trends in the Lesser Spotted Woodpecker Dendrocopos minor

Annual and seasonal variation in reproductive timing and performance were studied in a population of the Lesser Spotted Woodpecker Dendrocopos minor over 10 years in southern Sweden. The median laying date of the first egg varied by up to 17 days between years, being generally larger than the variation of laying dates within years. Neither clutch size, brood size in successful nests, fledging success in successful nests nor mean nestling weight differed significantly between years. There was no trend for mean clutch size to vary between early and late years. In spite of a more than threefold variation in population size, no reproductive variable demonstrated an apparent density-dependence. Within the season, clutch size declined steeply with increasing clutch initiation date, whereas fledging success and nesting success did not, leading to a trend in brood size almost identical to the trend in clutch size. The survival prospects of fledged young declined with increasing clutch initiation date, and it is argued that the clutch size laid is a strategic adjustment to laying date. Out of 124 breeding attempts, 34% did not produce fledged young. In 9% of the breeding attempts, pairs laid no eggs. At least 20% of the breeding attempts failed after egg-laying. The most common cause of breeding failure was loss of the breeding partner followed by nest abandonment (40% of the failures). Only 16–28% of the failures were due to predation on the nest. Most complete failures, and also partial losses from nests, occurred at the early breeding stages. It is argued that the early nestling phase may be a critical stage, which the woodpeckers adjust to coincide with the seasonal food peak, explaining the strikingly late breeding season compared with other non-migrant species.     

Reproductive responses of birds to experimental food supplementation: a meta-analysis

Introduction

Food availability is an important environmental cue for animals for deciding how much to invest in reproduction, and it ultimately affects population size. The importance of food limitation has been extensively studied in terrestrial vertebrate populations, especially in birds, by experimentally manipulating food supply. However, the factors explaining variation in reproductive decisions in response to food supplementation remain unclear. By performing meta-analyses, we aim to quantify the extent to which supplementary feeding affects several reproductive parameters in birds, and identify the key factors (life-history traits, behavioural factors, environmental factors, and experimental design) that can induce variation in laying date, clutch size and breeding success (i.e., number of fledglings produced) in response to food supplementation.

Results

Food supplementation produced variable but mostly positive effects across reproductive parameters in a total of 201 experiments from 82 independent studies. The outcomes of the food effect were modulated by environmental factors, e.g., laying dates advanced more towards low latitudes, and food supplementation appeared not to produce any obvious effect on bird reproduction when the background level of food abundance in the environment was high. Moreover, the increase in clutch size following food addition was more pronounced in birds that cache food, as compared to birds that do not. Supplementation timing was identified as a major cause of variation in breeding success responses. We also document the absence of a detectable food effect on clutch size and breeding success when the target species had poor access to the feed due to competitive interactions with other animals.

Conclusions

Our findings indicate that, from the pool of bird species and environments reviewed, extra food is allocated to immediate reproduction in most cases. Our results also support the view that bird species have evolved different life-history strategies to cope with environmental variability in food supply. However, we encourage more research at low latitudes to gain knowledge on how resource allocation in birds changes along a latitudinal gradient. Our results also emphasize the importance of developing experimental designs that minimise competition for the supplemented food and the risk of reproductive bottle-necks due to inappropriate supplementation timings.

     

Nosy neighbours: large broods attract more visitors. A field experiment in the pied flycatcher, <Emphasis Type="Italic">Ficedula hypoleuca</Emphasis>

Life is uncertain. To reduce uncertainty and make adaptive decisions, individuals need to collect information. Individuals often visit the breeding sites of their conspecifics (i.e., “prospect”), likely to assess conspecifics’ reproductive success and to use such information to identify high-quality spots for future breeding. We investigated whether visitation rate by prospectors and success of visited sites are causally linked. We manipulated the reproductive success (enlarged, reduced, and control broods) in a nest-box population of migratory pied flycatchers, Ficedula hypoleuca, in Finland. We measured the visitation rates of prospectors at 87 nest-boxes continuously from manipulation (day 3 after hatching) to fledging. 302 adult pied flycatchers prospected 9194 times on these manipulated nests (at least 78% of detected prospectors were successful breeders). While the number of visitors and visits was not influenced by the relative change in brood size we induced, the resulting absolute brood size predicted the prospecting behaviour: the larger the brood size after manipulation, the more visitors and visits a nest had. The parental provisioning rate at a nest and brood size pre-manipulation did not predict the number of visitors or visits post-manipulation. More visitors, however, inspected early than late nests and broods in good condition. Our study suggests that individuals collect social information when visiting conspecific nests during breeding and provides evidence that large broods attract more visitors than small broods. We discuss the results in light of individual decision-making by animals in their natural environments.     

Geographical variation in reproductive investment across avian assemblages in Europe: effects of environmental drivers differ between altricial and precocial species

Reproductive traits provide information about the ways by which available resources are allocated during breeding. We tested for environmental drivers of large scale geographical patterns in assemblage mean clutch size, number of broods and overall reproductive investment per breeding season in European birds. We combined data about geographical distribution with published information about reproductive traits, and calculated mean trait values for avian assemblages occurring in 50 × 50 km grid cells. In total, we employed data from 499 species and 2059 assemblages. As the time available for breeding and the amount of food limit the reproductive effort, we related the geographical variation in reproductive traits to the length of breeding season, normalized difference vegetation index (NDVI) as a surrogate of resource availability, and its seasonality. Geographical patterns in traits may differ between reproductive modes, thus we performed the analyses separately for altricial Passerines (n = 203) and precocial non‐passerine species (n = 164) and controlled for the effect of taxonomy. Large clutches dominated in areas with high NDVI and, in precocial birds, also in areas with high annual seasonality and a long breeding season. High number of broods and high overall reproductive investment dominated in areas with a long breeding season, and high number of broods was found also in areas with low annual seasonality, but only in precocial species. High overall reproductive investment dominated in highly productive areas and also in areas with low annual seasonality in both groups. The increase in reproductive investment is caused mostly by an increase in the number of broods related to the length of season and partly by increase in clutch size related to NDVI. We found a negative correlation between clutch size and the number of broods in Passerines, which might suggest a trade‐off between these traits. Processes behind trait patterns differ between altricial and precocial species.     

Effects of a food supplementation experiment on reproductive investment and a post-mating sexually selected trait in magpies Pica pica

Food availability is an important factor affecting breeding success in birds. Food supplementation experiments in birds have in general focused on the effects on reproductive success in terms of female investment (laying date, clutch size, egg size), however, it is also known that the estimation of mate quality based on sexually selected signals influences female reproductive investment. In the particular case of magpies, females use nest size, a post-mating sexually selected signal, to assess male's likelihood to invest in reproduction, and accordingly adjust reproductive investment (clutch size). Then, the possible effects of food supplementation on female reproductive investment could be mediated by other variables related to parental quality, such as nest size in magpies. In the present study, we explore if higher food availability in a magpie territory affected both male sexually selected traits (i.e. nest size) and female reproductive investment (laying date, egg size, clutch size). We performed a food supplementation experiment in which we experimentally increased food availability in several magpie territories, keeping others as controls. In food-supplemented territories, males built significantly larger nests and females significantly increased egg size by 4.1% compared to control females. Results suggest that the continuous provisioning of protein rich food allowed magpie females to increase egg size. However, laying date and clutch size did not differ between control and food-supplemented magpie pairs. Food availability also affected the relationship between female reproductive investment and nest size. In control territories, females decreased their egg size in response to a larger nest, whereas a tendency for the opposite relationship was revealed in food-supplemented territories. We discuss the possibility that magpie females adopt different strategies for reproductive investment according to food availability.     

Relationship between clutch size,egg volume and hatching success in a Yellow-legged Gull Larus michahellis colony in south-eastern Tunisia

This study aimed to test the hypothesis that clutch size covaries with egg volume and hatching success in the Yellow-legged Gull Larus michahellis. We determined clutch size and egg volume in a sample of 131 nests, and we used the data to check whether egg volume varied among nests according to clutch size, while taking into account the effects of egg laying order. We also estimated hatching success rate and investigated the relationship between hatching success and clutch size. Egg volume varied among clutches according to clutch size, with eggs being larger in three-egg clutches than in two-egg clutches. Moreover, three-egg clutches showed higher daily survival rates, and hence hatching success, than two-egg clutches. Overall, our results suggest that in the Yellow-legged Gull clutch size covaries with egg volume and hatching success, which could possibly reflect an age effect through different mechanisms. Indeed, older females could be hypothesised to exhibit greater breeding performance than younger females because of their higher experience in tapping energy resources for egg formation and defending nests from dangers. Moreover, due to their age, older females are likely to have lower residual reproductive potential and should invest more heavily in current breeding attempts.     

Breeding biology of the Little Egret Egretta garzetta on the southern coast of the Bay of Biscay

ABSTRACT

Capsule: Annual reproductive parameters of Little Egrets Egretta garzetta in an Atlantic coastal colony showed strong variation in a 20-year study, mainly due to extreme events.

Aims: To describe the breeding biology of Little Egrets in the Bay of Biscay and compare it with that of the Mediterranean basin. Also, to explore relationships between breeding parameters and colony size with some climatic indicators.

Methods: Phenology, number of nests, clutch size, number of hatchlings, brood size, and hatching and breeding success were recorded over 20 years.

Results: Median laying date of 214 nests was 1 May (range: 1 April–25 June) and 54% of the clutches were laid in the second half of April and the first half of May. Over the 20-year study period mean clutch size of 270 nests was 4.0, mean number of hatchlings was 3.1, and mean brood size was 2.3. Hatching success ranged from 46.1% to 100% and breeding success from zero to 100%. Number of nests was negatively associated with clutch and brood size. The highest clutch size and lowest brood size were recorded at the beginning of the season. Significant relationships were found between the number of hatchlings and the rainfall during the pre-breeding season, and between brood size and the summer rainfall.

Conclusions: Reproductive parameters showed significant variation over the study period, which highlights the importance of using long-term data sets. Breeding occurred one month later than in natural colonies of the Mediterranean basin. Negative relationships between the number of nests and clutch and brood size suggest some degree of density-dependent effects. Ranges of clutch size, number of hatchlings, and brood size were within those reported in Mediterranean populations. The effect of rainfall on reproductive parameters was weak. Extreme weather and predation events caused low rates of hatching and breeding success that affected the growth of the colony.     

青海湖鸬鹚繁殖成功率初报

2000年3—8月对青海湖西山鸬鹚岛鸬鹚(Phalacrocorax carbo)不同繁殖阶段的成功率做了初步观察。鸬鹚卵的窝卵数为3．47枚,孵化率为37．9％,雏鸟成活率为61．1％,繁殖成功率为23．2％。孵化期的前3d,鸬鹚巢的损失率高达36．1％;育雏期前10d雏鸟损失率为25％。这些数据可看作鸬鹚种群-青海湖湿地生态系统的一个重要组分——动态监测的起始数据之一,可在未来进一步的比较分析中得到应用。     

Intraspecific variation in size and density of Avocet colonies: effects of nest-distances on hatching and breeding success

In many colonial bird species there is considerable intraspecific variation in colony size and inter‐nest distance (colony density). Possible causes of this variation and its effects on hatching success (survival of eggs) and breeding success (probability of a pair raising chicks) were studied in 48 Avocet Recurvirostra avosetta colonies in Schleswig‐Holstein (Germany) between 1991 and 1996. Colony density was influenced by time of year and habitat (categories: island or mainland, close to or far from feeding grounds). Colonies on islands had the highest densities. When all available space at a colony site was used, colonies became very dense (mean nearest‐neighbour nest distance less than 1 m). Colony size (number of clutches) was influenced by time of year, but not by habitat. Hatching success was low in high density colonies and in very low density ‘colonies’ (single nests) and high over a broad range of intermediate nest densities. The low success rate of single nests was caused by a very high predation rate, whereas the low success rate in very dense colonies was caused by a high rate of nest abandonment. Nest abandonment in very dense colonies was associated with a high level of aggressiveness among Avocets during the egg‐laying period. Due to territorial behaviour, Avocets seemed to be expelled from the densest breeding sites. In very dense colonies, high frequencies of clutches of unusual size occurred due to conspecific nest parasitism. The number of Avocets taking part in attacks on potential egg predators was small and (in colonies of more than one clutch) depended neither on colony size nor on colony density. Despite a low hatching success in very dense colonies, individuals breeding in the densest colonies had significantly better chances of raising chicks than Avocets breeding in less dense colonies. Coloniality seemed to be obligatory for Avocets in order to ensure hatching success. The size and density of colonies seemed to be associated with the availability of suitable nesting habitats (islands).     

Don'T put all your eggs in one nest: spread them and cut time at risk

Abstract In many egg-laying animals, some females spread their clutch among several nests. The fitness effects of this reproductive tactic are obscure. Using mathematical modeling and field observations, we analyze an unexplored benefit of egg spreading in brood parasitic and other breeding systems: reduced time at risk for offspring. If a clutch takes many days to lay until incubation and embryo development starts after the last egg, by spreading her eggs a parasitic female can reduce offspring time in the vulnerable nest at risk of predation or other destruction. The model suggests that she can achieve much of this benefit by spreading her eggs among a few nests, even if her total clutch is large. Field data from goldeneye ducks Bucephala clangula show that egg spreading enables a fecund female to lay a clutch that is much larger than average without increasing offspring time at risk in a nest. This advantage increases with female condition (fecundity) and can markedly raise female reproductive success. These results help explain the puzzle of nesting parasites in some precocial birds, which lay eggs in the nests of other females before laying eggs in their own nest. Risk reduction by egg spreading may also play a role in the evolution of other breeding systems and taxa-for instance, polyandry with male parental care in some birds and fishes.     

Fear factor: prey habitat selection and its consequences in a predation risk landscape

Predation risk influences prey use of space. However, little is known about how predation risk influences breeding habitat selection and the fitness consequences of these decisions. The nest sites of central-place foraging predators may spatially anchor predation risk in the landscape. We explored how the spatial dispersion of avian predator nests influenced prey territory location and fitness related measures. We placed 249 nest boxes for migrant pied flycatchers Ficedula hypoleuca , at distances between 10 and 630 m, around seven different sparrowhawk nests Accipiter nisus . After closely monitoring flycatcher nests we found that flycatcher arrival dates, nest box occupation rates and clutch size showed a unimodal relationship with distance from sparrowhawk nests. This relationship suggested an optimal territory location at intermediate distances between 330 and 430 m from sparrowhawk nests. Furthermore, pied flycatcher nestling quantity and quality increased linearly with distance from sparrowhawk nests. These fitness related measures were between 4 and 26% larger in flycatcher nestlings raised far from, relative to those raised nearby, sparrowhawk nests. Our results suggest that breeding sparrowhawk affected both flycatcher habitat selection and reproductive success. We propose that nesting predators create predictable spatial variation in predation risk for both adult prey and possibly their nests, to which prey individuals are able to adaptively respond. Recognising predictable spatial variation in perceived predation risk may be fundamental for a proper understanding of predator-prey interactions and indeed prey species interactions.     

Plasma mammalian leptin analogue predicts reproductive phenology,but not reproductive output in a capital‐income breeding seaduck

To invest in energetically demanding life history stages, individuals require a substantial amount of resources. Physiological traits, particularly those related to energetics, can be useful for examining variation in life history decisions and trade‐offs because they result from individual responses to environmental variation. Leptin is a protein hormone found in mammals that is proportional to the amount of endogenous fat stores within an individual. Recently, researchers have confirmed that a mammalian leptin analogue (MLA), based on the mammalian sequence of leptin, is present with associated receptors and proteins in avian species, with an inhibitory effect on foraging and body mass gain at high circulating levels. While MLA has been both quantified and manipulated in avian species, little is currently known regarding whether plasma MLA in wild‐living species and individuals is associated with key reproductive decisions. We quantified plasma MLA in wild, Arctic‐nesting female common eiders (Somateria mollissima) at arrival on the breeding grounds and followed them to determine subsequent breeding propensity, and reproductive phenology, investment, and success. Common eiders are capital‐income breeding birds that require the accumulation of substantial fat stores to initiate laying and successfully complete incubation. We found that females with lower plasma MLA initiated breeding earlier and in a shorter period of time. However, we found no links between plasma MLA levels and breeding propensity, clutch size, or reproductive success. Although little is still known about plasma MLA, based on these results and its role in influencing foraging behaviors and condition gain, plasma MLA appears to be closely linked to reproductive timing and is therefore likely to underlie trade‐offs surrounding life history decisions.     

Body condition,migration, and timing of reproduction in snow geese: a test of the condition-dependent model of optimal clutch size

The seasonal decline of avian clutch size may result from the conflict between the advantage of early breeding (greater offspring value) and the advantage of a delay in lay date (improved body condition and hence clutch size). We tested predictions of a condition-dependent individual optimization model based on this trade-off (Rowe et al. 1994) in a long-distance migrant, the greater snow goose (Chen caerulescens atlantica), using data on condition, migration, and reproductive decisions of individuals. We closely tracked radio-marked females at their main spring staging area and on their breeding grounds. Our results were consistent with predictions of the model. Early-arriving females had a longer prelaying period and initiated their nests earlier than late arrivals. After controlling statistically for arrival date, we determined that females with high premigration condition had an earlier lay date than those in low condition. After controlling for the seasonal decline (i.e., lay date), we observed that clutch size was not related to premigration condition. Moreover, we took advantage of an unplanned manipulation of the prebreeding condition that occurred during our long-term study. We found that a reduction in condition caused a delay in lay date. However, after controlling for the seasonal decline, it did not affect clutch size. Our study indicates that geese simultaneously adjust their lay date and clutch size according to their premigration condition and migratory behavior as predicted by the condition-dependent optimization model.     

Drivers of age-specific survival in a long-lived seabird: contributions of observed and hidden sources of heterogeneity

1.?We assessed the relative influence of variability in recruitment age, dynamic reproductive investment (time-specific reproductive states) and frailty (unobserved differences in survival abilities across individuals) on survival in the black-legged kittiwake. Furthermore, we examined whether observed variability in survival trajectories was best explained by immediate reproductive investment, cumulative investment, or both. 2.?Individuals that delayed recruitment (≥ age 7) suffered a higher mortality risk than early recruits (age 3), especially later in life, suggesting that recruitment age may be an indicator of individual quality. Although recruitment age helped explain variation in survival, time-varying reproductive investment had a more substantial influence. 3.?The dichotomy of attempting to breed or not explained variability in survival across life better than other parameterizations of reproductive states such as clutch size, brood size or breeding success. In the kittiwake, the sinequanon condition to initiate reproduction is to hold a nest site, which is considered a very competitive activity. This might explain why attempting to breed is the key level of investment that affects survival, independent of the outcome (failure or success). 4.?Interestingly, the more individuals cumulate reproductive attempts over life, the lower their mortality risk, indicating that breeding experience may be a good indicator of parental quality as well. In contrast, attempting to breed at time t increased the risk of mortality between t and t + 1. We thus detected an immediate trade-off between attempting to breed and survival in this population; however, the earlier individuals recruited, and the more breeding experience they accumulated, the smaller the cost. 5.?Lastly, unobserved heterogeneity across individuals improved model fit more (1·3 times) than fixed and dynamic sources of observed heterogeneity in reproductive investment, demonstrating that it is critical to account for both sources of individual heterogeneity when studying survival trajectories. Only after simultaneously accounting for both sources of heterogeneity were we able to detect the 'cost' of immediate reproductive investment on survival and the 'benefit' of cumulative breeding attempts (experience), a proxy to individual quality.     

Female coloration indicates female reproductive capacity in blue tits

It is poorly understood whether female morphological and behavioural traits can be used as 'signals'. In particular, experimental tests of the hypothesis that female ornaments reflect quality are scarce. Here, we experimentally examine whether female plumage coloration might signal maternal quality in the blue tit, Cyanistes caeruleus by forcing half of the females breeding in our population to produce a replacement clutch. Using statistical models that controlled for the effects of male coloration, and the effects of age and condition of both parents, we found that carotenoid-based female coloration was positively linked to key proxies of bird lifetime reproductive success: clutch size, fledgling success and recruitment. Importantly, the relationships between maternal yellow carotenoid coloration and both clutch size and recruitment were stronger in the experimental group than in the control group, indicating that breeding females with higher values of yellow coloration were better able to handle the cost of producing a second clutch. Finally, UV-blue female coloration was positively linked to female survival and marginally linked to laying date. Taken together, these results show for the first time in a natural population that female coloration can indicate individual and maternal quality under natural and adverse reproductive conditions. They highlight the potential for the evolution of female ornamental traits through sexual selection.