Group composition and reproductive success of cooperatively breeding white‐winged choughs (Corcorax melanorhamphos) in urban and non‐urban habitat

Abstract: White‐winged choughs (Corcorax melanorhamphos, Corcoracidae) are a common, breeding resident in and around the city of Canberra in the Australian Capital Territory. We compared five measures of reproductive success between the urban and non‐urban populations of choughs to investigate the effect of urbanization on this cooperatively breeding species. Urban choughs initiated breeding earlier than their non‐urban counterparts and were more likely to suffer nest failures. However, there was no difference in the number of successful nests in a season or the number of fledglings produced per successful nesting attempt. A greater proportion of fledglings survived their first 12 months in the non‐urban habitat. We suggest that increased rates of nest predation and fledgling mortality in the urban environment may have a negative effect on reproductive success and remove any advantage that might be gained through a longer breeding season. Possible effects of urbanization on the social and genetic structure of white‐winged choughs are also discussed.     

Decision analysis for conservation breeding: Maximizing production for reintroduction of whooping cranes

Captive breeding is key to management of severely endangered species, but maximizing captive production can be challenging because of poor knowledge of species breeding biology and the complexity of evaluating different management options. In the face of uncertainty and complexity, decision-analytic approaches can be used to identify optimal management options for maximizing captive production. Building decision-analytic models requires iterations of model conception, data analysis, model building and evaluation, identification of remaining uncertainty, further research and monitoring to reduce uncertainty, and integration of new data into the model. We initiated such a process to maximize captive production of the whooping crane (Grus americana), the world's most endangered crane, which is managed through captive breeding and reintroduction. We collected 15 years of captive breeding data from 3 institutions and used Bayesian analysis and model selection to identify predictors of whooping crane hatching success. The strongest predictor, and that with clear management relevance, was incubation environment. The incubation period of whooping crane eggs is split across two environments: crane nests and artificial incubators. Although artificial incubators are useful for allowing breeding pairs to produce multiple clutches, our results indicate that crane incubation is most effective at promoting hatching success. Hatching probability increased the longer an egg spent in a crane nest, from 40% hatching probability for eggs receiving 1 day of crane incubation to 95% for those receiving 30 days (time incubated in each environment varied independently of total incubation period). Because birds will lay fewer eggs when they are incubating longer, a tradeoff exists between the number of clutches produced and egg hatching probability. We developed a decision-analytic model that estimated 16 to be the optimal number of days of crane incubation needed to maximize the number of offspring produced. These results show that using decision-analytic tools to account for uncertainty in captive breeding can improve the rate at which such programs contribute to wildlife reintroductions. © 2011 The Wildlife Society.     

Comparative nest survival of three sympatric loon species breeding in the Arctic

Identifying factors influencing nest survival among sympatric species is important for understanding and managing sources of variation in population dynamics of individual species. Three species of loons nest sympatrically in northern Alaska and differ in body size, life history characteristics, and population trends. We tested the effects of competition, nest site selection, and water level variations on nest survival of Pacific Gavia pacifica, yellow‐billed G. adamsii, and red‐throated loons G. stellata on the Arctic Coastal Plain in Alaska. Although overall nest survival rates did not differ between species, the factors influencing nest survival varied. Nest site selection influenced nest survival for Pacific and yellow‐billed loons, with both species having high nest survival when nesting on islands and peninsulas, likely due to a reduction in access by terrestrial predators. However, on mainland shorelines, Pacific loons had lower nest survival than yellow‐billed loons, and used a higher proportion of vegetation mats for nest sites suggesting that their smaller body size makes them less adept at nest defense. Nest site selection did not influence nest survival of red‐throated loons corresponding to our result of no nest site preferences by this species. Initiation date had a strong influence on nest survival for Pacific and yellow‐billed loons with nests laid earlier having higher survival. Pacific and yellow‐billed loon nests were susceptible to flooding due to precipitation, which contrasted with red‐throated loons that nest on smaller lakes with lower water level variations. Competition did not affect nest survival for any of the species likely due to most territorial encounters occurring prior to incubation. The only influence we found on red‐throated loon nest survival was differences among years. Our results indicate that loons chose nest sites based on predation risk and that factors influencing breeding success of closely related species may differ under similar breeding conditions.     

Using net sacks to examine the relationship between egg size and young size in Common Pochards

ABSTRACT.   Investigators studying the relationship between egg size and young size often face difficulties in assigning particular young to particular eggs. We present a new method based on the use of separate net sacks for each egg that allowed us to study this phenomenon in Common Pochards ( Aythya ferina ) without excessive disturbance of breeding birds. We found a positive isometric relationship between duckling body mass and estimated fresh egg mass ( P < 0.001). However, we found no relationship between estimated fresh egg mass and structural measurements of ducklings (tarsus and bill length, both P > 0.29). In addition, we found a weak positive relationship between estimated egg mass and the size of ducklings ( P = 0.049). Greater hatchling mass typically means a larger yolk supply, and more yolk may provide energy crucial for survival during the first few days after hatching. In addition, ducklings with longer tarsi may be better swimmers and better at finding food and escaping from predators. Although our method has certain limitations and investigators should consider risks when applying it in over-water nests or in nests with large clutches, we believe it is a reliable way to assign young to particular eggs in waterfowl and possibly other bird species.     

Comparative reproductive biology of sympatric species: nest and chick survival of American avocets and black‐necked stilts

Identifying differences in reproductive success rates of closely related and sympatrically breeding species can be useful for understanding limitations to population growth. We simultaneously examined the reproductive ecology of American avocets Recurvirostra americana and black‐necked stilts Himantopus mexicanus using 1274 monitored nests and 240 radio‐marked chicks in San Francisco Bay, California. Although there were 1.8 times more avocet nests than stilt nests, stilts nonetheless fledged 3.3 times more chicks. Greater production by stilts than avocets was the result of greater chick survival from hatching to fledging (avocet: 6%; stilt: 40%), and not because of differences in clutch size (avocet: 3.84; stilt: 3.77), nest survival (avocet: 44%; stilt: 35%), or egg hatching success (avocet: 90%; stilt: 92%). We reviewed the literature and confirmed that nest survival and hatching success are generally similar when avocets and stilts breed sympatrically. In addition to species, chick survival was strongly influenced by age, site, and year. In particular, daily survival rates increased rapidly with chick age, with 70% of mortalities occurring ≤ 1 week after hatch. California gulls Larus californicus caused 55% of avocet, but only 15% of stilt, chick deaths. Differential use of micro‐habitats likely reduced stilt chick's vulnerability to gull predation, particularly during the first week after hatch, because stilts nested in vegetation 2.7 times more often than avocets and vegetation height was 65% taller at stilt nests compared with avocet nests. Our results demonstrate that two co‐occurring and closely related species with similar life history strategies can differ markedly in reproductive success, and simultaneous studies of such species can identify differences that limit productivity.     

Incubation temperature impacts nestling growth and survival in an open‐cup nesting passerine

For oviparous species such as birds, conditions experienced while in the egg can have long‐lasting effects on the individual. The impact of subtle changes in incubation temperature on nestling development, however, remains poorly understood, especially for open‐cup nesting species with altricial young. To investigate how incubation temperature affects nestling development and survival in such species, we artificially incubated American robin (Turdus migratorius) eggs at 36.1°C (“Low” treatment) and 37.8°C (“High” treatment). Chicks were fostered to same‐age nests upon hatching, and we measured mass, tarsus, and wing length of experimental nestlings and one randomly selected, naturally incubated (“Natural”), foster nest‐mate on days 7 and 10 posthatch. We found significant effects of incubation temperature on incubation duration, growth, and survival, in which experimentally incubated nestlings had shorter incubation periods (10.22, 11.50, and 11.95 days for High, Low, and Natural eggs, respectively), and nestlings from the Low treatment were smaller and had reduced survival compared to High and Natural nestlings. These results highlight the importance of incubation conditions during embryonic development for incubation duration, somatic development, and survival. Moreover, these findings indicate that differences in incubation temperature within the natural range of variation can have important carryover effects on growth and survival in species with altricial young.     

Nest monitoring does not affect nesting success of Whinchats Saxicola rubetra

It is important to assess the effect that research activities may have on animals in the wild, especially when key parameters, such as breeding success, could potentially be influenced by observer activity. For birds, some studies have suggested that nest monitoring can increase the chances of nest failure due to predation, whereas others suggest that human nest visits may actually deter mammalian predators. Nest monitoring visits can also influence breeding success more indirectly by altering parental provisioning behaviour. Here, the influence of monitoring activities on nest success was examined in a ground‐nesting grassland bird, the Whinchat Saxicola rubetra. During the egg phase, a sample of nests were not visited between the initial finding event and the estimated hatching date; instead, the nest status was assessed at a distance. Daily survival rates (DSR) for these nests were compared with that of nests visited every 2 days. During the nestling phase, the effects of observer nest visits on parental provisioning behaviour were determined. Nest visits were found not to affect egg DSR significantly, and parental provisioning was disrupted for a maximum of 20 min (0.52% of the nestling period) following an observer visit. Given the variation in response to nest visits across species, we suggest that consideration should be given to observer impact in all studies where predation risk is high. Here, we illustrate a method for researchers to assess the impact of their nest visits to ensure they are not biasing estimates of breeding success.     

Consequences of a changing environment on the breeding phenology and reproductive success components in a long-distance migratory bird

Migratory birds have a narrow time window to breed, especially in the Arctic, where early nesting typically yields the highest reproductive success. We assessed temporal changes (1991–2015) in reproductive success components in relation to timing of breeding in greater snow geese (Chen caerulescens atlantica). This species breeds in the Canadian Arctic, a region that has experienced a strong warming trend. We tested the effect of laying or hatching date, year and their interaction on six reproductive components: Total clutch laid, nesting success, egg survival, hatching success, prefledging, and postfledging survival. Over 25 years, mean laying date changed little, even though it advanced 1.8 days in early breeders and was delayed 3 days in late breeders. Likewise, the number of eggs in nests initiated early in the season decreased by 0.6 egg, whereas in late nests it increased by 0.3 egg. Success of nests initiated early and late in the season was lower than nests initiated near the population mean, and consistently increased over time. The proportion of eggs surviving to partial predation and postfledging survival decreased with laying date but the pattern did not change over time. In contrast, prefledging survival was not affected by laying date initially but declined in nests initiated late in the season toward the end of the study. Overall, nests initiated close to the population mean showed little temporal change for most components of reproductive success and seem to be less affected by environmental change than nests initiated early and late in the season.     

Rates and consequences of relaying in little auks Alle alle breeding in the High Arctic an experimental study with egg removal

Most seabirds have a small clutch size. Thus, replacement of a clutch after loss can make important contributions to an individual’s lifetime reproductive success. However, in the condition of short polar summer, relaying propensity may be time‐constrained. In this study, we investigated rates and consequences of relaying in a small High Arctic seabird, the little auk Alle alle. We performed an experiment in which we removed the single egg from 20 nests of early‐laying breeders. We measured relaying rates, and compared chick body mass and breeding success between the experimental and control nests. Despite the narrow window of the Arctic summer and the closely synchronized breeding, 75% of females produced a replacement egg just 2.7% smaller in volume than the first egg. This indicates that in little auks, the demographic effects of disruptions to breeding attempts (by predators, adverse weather or human activity) may be mitigated to some extent by replacement clutches. However, peak body mass and fledging body mass were lower in the experimental than the control chicks. This effect was rather a consequence of late hatching – chicks from replacement clutches followed seasonal decline in peak body mass and fledging mass. Finally, breeding success and chick survival up to 20 d in the experimental nests were respectively 34 and 37% lower than in the control nests. Thus, the quality and post‐fledging survival of chicks from the replacement clutches were probably lower compared to the chicks hatched from the first‐laid eggs.     

Age‐specific survival and recruitment of piping plovers Charadrius melodus in the Great Lakes region

Juvenile survival and age at first breeding (i.e. recruitment) are critical parameters affecting population dynamics in birds, but high levels of natal dispersal preclude measurement of these variables in most species. We used multi‐state capture–recapture models to measure age‐specific survival and recruitment probabilities of piping plovers Charadrius melodus in the Great Lakes region during 1993–2012. This federally endangered population is thoroughly monitored throughout its entire breeding range, minimizing concerns that measures of survival and recruitment are confounded by temporary or permanent emigration. First‐year survival (± SE) averaged 0.284 ± 0.019 from mean banding age (9 d) and 0.374 ± 0.023 from fledging age (23 d). Factors that increased first‐year survival during the pre‐fledging period (9–23 d) included earlier hatching dates, older age at banding, greater number of fledglings at a given site, and better body condition at time of banding. However, when chicks that died prior to fledging were excluded from analysis, only earlier hatching dates improved first‐year survival estimates. Females had a higher probability (0.557 ± 0.066) of initiating breeding at age one than did males (0.353 ± 0.052), but virtually all plovers began breeding by age three. Adult survival was reduced by increased hurricane activity on the southeast U.S. Atlantic coast where Great Lakes piping plovers winter and by higher populations of merlins Falco columbarius. Mean annual adult survival declined from 1993 to 2012, and did not differ between males and females. Enhanced body condition led to higher survival to fledge and early breeding led to improved first‐year survival; therefore, management actions focused on ensuring access to quality feeding habitat for growing young and protecting early nests may increase recruitment in this federally endangered population.     

Egg temperature and initial brood patch area determine hatching asynchrony in Magellanic penguin Spheniscus magellanicus

In birds, the adaptive significance of hatching asynchrony has been under debate for many years and the parental effects on hatching asynchrony have been largely assumed but not often tested. Some authors suggest that hatching asynchrony depends on the incubation onset and many factors have been shown to influence hatching asynchrony in different species. Our objective was to analyze the exact timing of the onset of incubation and if this affects hatching asynchrony; and, in addition, which other factors (brood patch development, incubation position, adult body condition, intra‐clutch egg dimorphism, laying date and year) affect hatching asynchrony in Magellanic penguins Spheniscus magellanicus. We first estimated the eggshell temperature at which embryo development starts, with a non‐destructive and novel method. We then recorded individual egg temperatures in 61 nests during incubation, and related them, and other breeding parameters, to hatching asynchrony. We also observed incubation positions in 307 nests. We found a significant positive relationship between hatching asynchrony and the temperature that the first‐laid egg experienced during egg laying and between hatching asynchrony and the initial brood patch area. We also found a negative relationship between hatching asynchrony and the difference in temperature between second and first‐laid eggs within a clutch, measured after the egg‐laying period was finished. We ruled out position of the eggs during incubation, adult body condition, egg volume, laying date, and study year as factors influencing hatching asynchrony. The egg temperature during laying and the difference in temperature between eggs of a clutch are determinants of hatching asynchrony in Magellanic penguins.     

Life‐history tradeoffs revealed by seasonal declines in reproductive traits of Arctic‐breeding shorebirds

Seasonal declines in breeding performance are widespread in wild animals, resulting from temporal changes in environmental conditions or from individual variation. Seasonal declines might drive selection for early breeding, with implications for other stages of the annual cycle. Alternatively, selection on the phenology of nonbreeding stages could constrain timing of the breeding season and lead to seasonal changes in reproductive performance. We studied 25 taxa of migratory shorebirds (including five subspecies) at 16 arctic sites in Russia, Alaska, and Canada. We investigated seasonal changes in four reproductive traits, and developed a novel Bayesian risk‐partitioning model of daily nest survival to examine seasonal trends in two causes of nest failure. We found strong seasonal declines in reproductive traits for a subset of species. The probability of laying a full four‐egg clutch declined by 8–78% in 12 of 25 taxa tested, daily nest survival rates declined by 1–12% in eight of 22 taxa, incubation duration declined by 2.0–2.5% in two of seven taxa, and mean egg volume declined by 5% in one of 15 taxa. Temporal changes were not fully explained by individual variation. Across all species, the proportion of failed nests that were depredated declined over the season from 0.98 to 0.60, while the proportion abandoned increased from 0.01 to 0.35 and drove the seasonal declines in nest survival. An increase in abandonment of late nests is consistent with a life‐history tradeoff whereby either adult mortality increased or adults deserted the breeding attempt to maximize adult survival. In turn, seasonal declines in clutch size and incubation duration might be adaptive to hasten hatching of later nests. In other species of shorebirds, we found no seasonal patterns in breeding performance, suggesting that some species are not subject to selective pressure for early breeding.     

Iodine deficiency affects hatchability of endangered captive kaki (Black Stilt,Himantopus novaezelandiae)

A significant increase in hatching and late‐incubation deaths during two breeding seasons in a captive population of endangered black stilts (kaki, Himantopus novaezelandiae) prompted an investigation into the potential role of iodine deficiency. During the 1997 and 1998 breeding seasons, the survival rate of captive‐laid eggs during artificial incubation and hatching was lower than that of eggs laid by wild kaki, which were collected and incubated artificially in the same environment. Necropsy results revealed abnormalities in thyroid structure. These abnormalities had been occurring in captive birds since 1994, but they peaked in 1997 and 1998 at 30% and 81%, respectively, of all diagnosed necropsies. Necropsies of pied stilts, hybrids, and one kaki from the wild revealed no thyroid abnormalities. Gross morphological symptoms observed at hatching in chicks that died were consistent with those described for iodine deficiency in poultry. Serum levels of thyroxine (T4) were considerably higher in wild pied stilts and hybrid chicks than in captive adult, subadult, and kaki chicks. In five individuals from the captive population, serum levels increased after about 160 days of iodine supplementation and approached values in the wild. The constraints associated with working with such a critically endangered species (e.g., small sample size and an urgent need to effect changes) precluded the use of an experimental approach to determine the role iodine deficiency played in peri‐hatching deaths. However, we believe the evidence presented here strongly supports a case for the impact of iodine on survival within the captive population. Our experience also highlights the need for vigilance in captive programs with respect to potential dietary deficiencies when endangered species are raised in captivity for conservation purposes. Zoo Biol 23:1–13, 2004. © 2004 Wiley‐Liss, Inc.     

Delayed egg‐laying and shortened incubation duration of Arctic‐breeding shorebirds coincide with climate cooling

Biological impacts of climate change are exemplified by shifts in phenology. As the timing of breeding advances, the within‐season relationships between timing of breeding and reproductive traits may change and cause long‐term changes in the population mean value of reproductive traits. We investigated long‐term changes in the timing of breeding and within‐season patterns of clutch size, egg volume, incubation duration, and daily nest survival of three shorebird species between two decades. Based on previously known within‐season patterns and assuming a warming trend, we hypothesized that the timing of clutch initiation would advance between decades and would be coupled with increases in mean clutch size, egg volume, and daily nest survival rate. We monitored 1,378 nests of western sandpipers, semipalmated sandpipers, and red‐necked phalaropes at a subarctic site during 1993–1996 and 2010–2014. Sandpipers have biparental incubation, whereas phalaropes have uniparental incubation. We found an unexpected long‐term cooling trend during the early part of the breeding season. Three species delayed clutch initiation by 5 days in the 2010s relative to the 1990s. Clutch size and daily nest survival showed strong within‐season declines in sandpipers, but not in phalaropes. Egg volume showed strong within‐season declines in one species of sandpiper, but increased in phalaropes. Despite the within‐season patterns in traits and shifts in phenology, clutch size, egg volume, and daily nest survival were similar between decades. In contrast, incubation duration did not show within‐season variation, but decreased by 2 days in sandpipers and increased by 2 days in phalaropes. Shorebirds demonstrated variable breeding phenology and incubation duration in relation to climate cooling, but little change in nonphenological components of traits. Our results indicate that the breeding phenology of shorebirds is closely associated with the temperature conditions on breeding ground, the effects of which can vary among reproductive traits and among sympatric species.     

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.     

Determining Great Bittern Botaurus stellaris laying date from egg and chick biometrics

Capsule Great Bittern breeding phenology can be estimated from egg and chick biometrics.

Aims To estimate egg or chick ages in order to back-calculate egg-laying dates.

Methods Bittern nests were searched for in six French and three Belarussian sites between 1999 and 2004. Eggs and chicks were measured at each visit. By using a subsample of nests with known egg-laying (or hatching) dates, regression equations are determined using egg density and tarsus length in order to estimate, respectively, egg and chick ages. Additionally in Belarus, the ‘water test’ was used to estimate the incubation stage of the clutch.

Results A total of 141 Bittern nests were found. Egg density decreased linearly from 1.063 at laying to 0.915 the day before hatching. A regression equation therefore allows estimation of egg age from its density. A scale was also constructed to estimate egg age from its position in water, and the accuracy of the two methods is compared. Chick growth rates were similar between the two countries. Before the age of 25 days, chicks are best aged by tarsus length compared to other measurements (weight, bill length). No data were available after that age because chicks were no longer found on nests.

Conclusions Egg-laying date can be estimated to within ±3 days using egg density, and to within ±5 days, using the ‘water test’. Tarsus length can be used until the age of 25 days to age chicks to within ±2 days. These simple measurements provide efficient and accurate methods to record the breeding calendar of this endangered species.     

An experimental approach to the brood reduction hypothesis in Magellanic penguins

In many bird species, eggs in a brood hatch within days of each other, leading to a size asymmetry detrimental to younger siblings. Hatching asynchrony is often thought of as an adaptive strategy, and the most widely studied hypothesis in relation to this is the ‘brood reduction hypothesis’. This hypothesis states that when food resources are unpredictable, hatching asynchrony will allow the adjustment of the brood size maximizing fledging success and benefitting parents. The Magellanic penguin Spheniscus magellanicus is an appropriate species to test this hypothesis because it has a 2‐egg clutch that hatches over a 2‐d interval with a broad range of variation (–1 to 4 d), it shows facultative brood reduction, and food abundance between breeding seasons is variable. We performed a manipulative study at Isla Quiroga, Argentina, during three breeding seasons (2010–2012) by forcing broods to hatch synchronously (0 d) or asynchronously (2 or 4 d). Years were categorized based on estimated food abundance. Our study provided mixed results because in the low estimated food abundance year asynchronous broods did not have higher nestling survival than synchronous broods, and the second‐hatchling in asynchronous broods did not die more often than those in synchronous broods. On the other hand, younger siblings of 4‐d asynchronous broods starved earlier than those of synchronous broods, and 2‐d asynchronous broods fledged heavier young than synchronous broods. Asynchronous hatching would seem to benefit reproduction in this species, not with respect to survival, but in terms of the advantages it can accord to nestlings and, in terms of lower costs, for parents raising nestlings.     

Hatching order affects offspring growth,survival and adult dominance in the joint‐laying Pukeko Porphyrio melanotus melanotus

In birds with asynchronous hatching, hatching order is an important factor in determining offspring phenotype. Many previous studies have demonstrated that later‐hatched offspring show reduced growth and survival during development. However, few studies have followed individuals from hatching to adulthood to test whether the effects of hatching order persist into later life. Here, we explore patterns of hatching order and fitness‐related traits in the Pukeko Porphyrio melanotus melanotus, a cooperatively breeding bird that lives in stable social groups that form linear dominance hierarchies. Pukeko groups sometimes contain two breeding females that lay eggs in the same nest (joint‐laying). Thus, competition between nest‐mates can influence the relative fitness of each laying female. We show that in both single‐clutch and joint‐clutch nests, earlier‐hatched Pukeko chicks grow faster and survive better than later‐hatched brood‐mates. Moreover, earlier‐hatched chicks achieve higher dominance ranks as adults, making this study one of the first to find a relationship between hatching order and adult dominance in wild birds. Finally, we show that in groups with two breeding females, the chicks of the primary female hatch earlier than the chicks of the secondary female. As a result, the offspring of the primary female may be at a competitive advantage, which could have important implications for social dynamics in this species.     

Nest building under the risk of predation: safe nests are not always the best option

Nest predation is a widespread demographic and evolutionary force in avian reproduction, but few studies have considered the circumstances under which birds might invest in the construction of safe nests. We examined this question using a stochastic simulation model based on a basic passerine breeding season. Nest safety functions were used to translate time invested in nest building into an increase in daily nest survival; that increase could be rapid, requiring only a few days to achieve a safe nest, or slow, taking many days to do so. The maximum achievable safety differed across nest safety functions. Given a limited length to the breeding season, a greater time investment in nest safety detracts from the time available for re‐nesting following successful or unsuccessful nesting attempts. In many circumstances, the best option is a quick‐build ‘minimal’ nest that provides adequate support for young, but little additional safety from attacks. This is especially true for scenarios that allow for multiple nesting attempts across a season. However, relatively safe nests that can be built fairly quickly are uniformly favored options. Safe, long‐build nests are favored only when they provide a great deal of safety over other nest‐building options, but greater safety alone is not sufficient for such investment. Simulations allowing only a single nesting attempt generally favor a greater investment in nest safety. Parental survival is another important factor in nest investment. Increased danger to the parent during nest building strongly favors a low investment in nests. However, substantial investment in a safe nest is favored when that safety extends to the incubating parent. Our results provide some insight into the prevalence of seemingly unsafe, open‐cup nests across the bird world, but the range of nest types that could potentially be built by a given species is an open question.     

Effects of intraclutch variation in egg size and hatching asynchrony on nestling development and survival in semi‐precocial Herring Gulls

Intraclutch egg size variation may non‐adaptively result from nutritional/energetic constraints acting on laying females or may reflect adaptive differential investment in offspring in relation to laying/hatching order. This variation may contribute to size hierarchies among siblings already established due to hatching asynchrony, and resultant competitive asymmetries often lead to starvation of the weakest nestling within a brood. The costs in terms of chick mortality can be high. However, the extent to which this mortality is egg size‐mediated remains unclear, especially in relation to hatching asynchrony which may operate concomitantly. I assessed effects of egg size and hatching asynchrony on nestling development and survival of Herring Gulls (Larus argentatus), where the smaller size and later hatching of c‐eggs may represent a brood‐reduction strategy. To analyze variation in egg size, I recorded the laying order and laying date of 870 eggs in 290 three‐egg clutches over a 3‐yr period (2010–2012). I measured hatchlings and monitored growth and survival of 130 chicks from enclosed nests in 2011 and 2012. The negative effect of laying date (β = ?0.18 ± SE 0.06, P = 0.002) on c‐egg size possibly reflected the fact that late breeders were either low quality or inexperienced females. The mass, size, and condition of hatchling Herring Gulls were positively related to egg size (all P < 0.0001). C‐chicks suffered from increased mortality risk during the first 12 d, identified as the brood‐reduction period in my study population. Although intraclutch variation in egg size was not directly related to patterns of chick mortality, I found that smaller relative egg size interactively increased differences in relative body condition of nestlings, primarily brought about by the degree of hatching asynchrony during this brood‐reduction period. Thus, the value of relatively small c‐eggs in Herring Gulls may lie in reinforcing brood reduction through effects on nestling body condition. A reproductive strategy Herring Gulls might have adopted to maintain a three‐egg clutch, but that also enables them to adjust the number of chicks they rear relative to the prevailing environmental conditions and to their own condition during the nestling stage.