Individual Breeding Performance Research Articles

Climate warming induced a stretch of the breeding season and an increase of second clutches in a passerine breeding at its altitudinal limits.

The increase in the average air temperature due to global warming has produced an early onset of the reproduction in many migratory birds of the Paleartic region. According to the “mismatch hypothesis” this response can lead to a decrease in the breeding output when the conditions that trigger the departure from the wintering areas do not match the availability of food resources in the breeding ground. We used 653 brooding events registered during the period 1991–2013 to investigate the link between climatic variables and individual breeding performance of a partially migratory passerine, the Rock Sparrow Petronia petronia, breeding at the altitude limit of its distribution. The laying date (LD) of the earliest first clutch was associated with local spring (minimum) temperatures but did not show a significant trend during the period considered. The LD of the latest first clutch had a positive and statistically significant trend, unrelated to local covariates and resulting in a longer breeding season (∼1.5 days/year). A longer breeding season allowed birds to produce more second clutches, which proportion increased from 0.14 to 0.25. The average breeding success was also positively correlated with the average temperature in July and with the duration of the breeding season. Contrary to expectations, the most important climate-dependent effect was a stretch of the breeding season due to a significant increase of the LD of the latest first-clutches rather than an earlier breeding onset. We show how climate changes act on bird populations through multiple paths and stress the need to assess the link between climatic variables and several aspects of the breeding cycle.

Evidence for high inter-generational individual quality in yellow-eyed penguins.

Longitudinal studies focusing on lifetime reproductive success (LRS) have been used to measure individual breeding performance and identify commonalities among successful breeders. By extending the focus to subsequent generations we identify a proportion of high-quality individuals that contribute disproportionately to the population over multiple generations. We used 23 years of yellow-eyed penguin (Megadyptes antipodes) breeding data from one breeding area to identify the proportion of individual birds that raised successful breeders, which in turn raised offspring. We explored which life-history components influenced LRS, as this knowledge would enable conservation resources to be focused on high-performing individuals in this endangered population. From 2,147 birds marked as chicks, 370 (17.2%) survived to adulthood and recruited to their natal location, of which 219 (10.2%) fledged offspring: 124 (56.6%) of the 219 birds produced offspring that recruited as breeders. Only 102 birds (4.8% of 2,147) fledged first-generation offspring that in turn fledged offspring (second-generation offspring, or grand-offspring). We found that ∼25% of the birds that survived to breed had above-average LRS as well as above-average numbers of grand-offspring, and were more likely to have produced first-generation chicks that recruited and also produced above-average numbers of second-generation chicks. Our findings suggest that there is a core of “super-breeders” that contribute disproportionately to the population over successive generations. Lifespan and age-at-first-breeding were correlated with LRS. We suggest that traits of birds relating to longevity, health (e.g., immunocompetence) and fitness could be examined to identify potential links with high LRS and inter-generational fecundity. “Super-breeders” appear to consistently achieve high LRS and long lifespans in a stochastic environment, demonstrating greater resilience in the face of extreme events.

Low Impact of Avian Pox on Captive-Bred Houbara Bustard Breeding Performance.

Avian pox, a disease caused by avipoxviruses, is a major cause of decline of some endangered bird species. While its impact has been assessed in several species in the wild, effects of the disease in conservation breeding have never been studied. Houbara bustard species (Chlamydotis undulata and Chlamydotis macqueenii), whose populations declined in the last decades, have been captive bred for conservation purposes for more than 20 years. While mortality and morbidity induced by avipoxviruses can be controlled by appropriate management, the disease might still affect bird breeding performance and jeopardize the production objectives of conservation programs. Impacts of the disease was studied during two outbreaks in captive-bred juvenile Houbara bustards in Morocco in 2009–2010 and 2010–2011, by modeling the effect of the disease on individual breeding performance (male display and female egg production) of 2,797 birds during their first breeding season. Results showed that the impact of avian pox on the ability of birds to reproduce and on the count of displays or eggs is low and mainly non-significant. The absence of strong impact compared to what could be observed in other species in the wild may be explained by the controlled conditions provided by captivity, especially the close veterinary monitoring of each bird. Those results emphasize the importance of individual management to prevent major disease emergence and their effects in captive breeding of endangered species.

Habitat fragmentation and reproductive success: a structural equation modelling approach

1. There is great interest on the effects of habitat fragmentation, whereby habitat is lost and the spatial configuration of remaining habitat patches is altered, on individual breeding performance. However, we still lack consensus of how this important process affects reproductive success, and whether its effects are mainly due to reduced fecundity or nestling survival. 2. The main reason for this may be the way that habitat fragmentation has been previously modelled. Studies have treated habitat loss and altered spatial configuration as two independent processes instead of as one hierarchical and interdependent process, and therefore have not been able to consider the relative direct and indirect effects of habitat loss and altered spatial configuration. 3. We investigated how habitat (i.e. old forest) fragmentation, caused by intense forest harvesting at the territory and landscape scales, is associated with the number of fledged offspring of an area-sensitive passerine, the Eurasian treecreeper (Certhia familiaris). We used structural equation modelling (SEM) to examine the complex hierarchical associations between habitat loss and altered spatial configuration on the number of fledged offspring, by controlling for individual condition and weather conditions during incubation. 4. Against generally held expectations, treecreeper reproductive success did not show a significant association with habitat fragmentation measured at the territory scale. Instead, our analyses suggested that an increasing amount of habitat at the landscape scale caused a significant increase in nest predation rates, leading to reduced reproductive success. This effect operated directly on nest predation rates, instead of acting indirectly through altered spatial configuration. 5. Because habitat amount and configuration are inherently strongly collinear, particularly when multiple scales are considered, our study demonstrates the usefulness of a SEM approach for hierarchical partitioning of habitat amount vs. habitat configuration in landscape ecology that may have bearing on biological conclusions.

Disentangling site and mate fidelity in a monogamous population under strong nest site competition

Site and mate fidelity are important aspects of animal mating systems and have implications for animal population structures. In birds, they are often related to the age, breeding experience, prior residence, pair duration or breeding performance of individuals. These variables are often correlated, and site and mate fidelity may also be associated with each other. Most previous studies focused on either site fidelity or mate fidelity, and the relationship between them remains to be explored. In this study, we used path analysis to examine the correlates of site and mate fidelity of Lanyu scops owls, Otus elegans botelensis, and to investigate the relationship between site and mate fidelity. Site fidelity of female owls was related to previous breeding success, following a ‘win-stay, lose-switch’ model. Site fidelity of male owls increased with both previous breeding success and years of prior residence. Mate fidelity was related to female site fidelity and pair duration. Although mate fidelity was higher after successful breeding attempts, it was a result of the breeding success-dependent site fidelity of the owls. Many studies used the positive correlation between previous breeding success and mate fidelity to support the better option hypothesis or the incompatibility hypothesis of divorce. Our study identified the most likely causal pathway for the complex site and mate fidelity pattern in the Lanyu scops owl, and demonstrated that it is important to identify the underlying driving forces to avoid erroneous conclusions.

Heterogeneity in detection probability along the breeding season in Black-legged Kittiwakes: implications for sampling design

In wild animal population studies, capture heterogeneity is likely to be prevalent and can reduce the accuracy of vital rate estimates. Here, we test how individual detection probabilities vary through the breeding season in a population of a cliff-nesting colonial seabird, the Black-legged Kittiwake (Rissa tridactyla). Specifically, we expected detection probability to be affected by changes in brooding behavior and nest attendance associated with the breeding phenology and the local breeding performance of individuals. As predicted, we found that strong heterogeneities in detection probability can occur in relation to the breeding performance of individuals, the breeding performance of their neighbors, and the timing of surveys. Detection probability is highest and most homogeneous at the beginning of the breeding season. Later in the season, it is lower and can vary dramatically among groups of breeding individuals. A simulation approach was used to assess the implications of these results for the performance (bias and precision) of different study designs. Clearly, investing sampling effort early in the season is an efficient way to improve the accuracy of parameter estimates in this species. Our findings stress the importance of establishing study designs that take into account the population and behavioral ecology of the focal species.

Supplementary feeding during the chick-rearing period is ineffective in increasing the breeding success in the bearded vulture (Gypaetus barbatus)

One of the most widespread tools in the conservation of scavenger species is the provision of supplementary food. However, scientific studies on its effectiveness have been rarely conducted. Here, we present the first results of an experimental specific supplementary feeding programme applied from hatching to 45-60 days, aimed at increasing the breeding success of an obligate brood reducer, the threatened bearded vulture Gypaetus barbatus. We tested its effectiveness in the Spanish Pyrenees, the locale of the most important population of this species in the European Union, in which a regressive trend in breeding parameters has been observed in recent years. We compared the breeding success in nests with supplementary food to non-supplemented control nests. Supplementary food did not significantly increase global breeding success (supplemented nests, 0.793±0.193 chicks per pair with eggs hatched vs non-supplemented nests, 0.771±0.185) or the individual breeding performance of the territories (supplemented period, 0.712±0.307 vs non- supplemented period, 0.642±0.311). The similar values obtained suggest that the specific supplementary feeding programme applied during the chick-rearing period is apparently ineffective at increasing breeding success. The results suggest that, at least in the study area, factors that provoke breeding failure after hatching continue to operate independently of the supplementary feeding programme. Although more research on this subject is required, these preliminary conclusions should be taken into account by managers in order to optimise the investment of economic resources and to better prioritise the future establishment of alternative conservation actions.

Reproductive performance of little penguinsEudyptula minorin relation to year, age, pair‐bond duration, breeding date and individual quality

We measured breeding performance of little penguinsEudyptula minorat Phillip Island, Victoria, Australia, during a 21‐year period. All birds considered in this paper (n=307) were of known age (2–22 y) and sex, and most were of known, or closely‐estimated, pair‐bond status (1–8 mates per bird; pair‐bond durations 1–13 y). Breeding dates and breeding performance varied markedly from year to year; measures of annual performance were not associated with early breeding. Measures of individual breeding performance (clutch‐size, hatching success, chick masses and productivity) were related to early laying, parental age, and duration of pair‐bond. Dependence of breeding performance on parental age was curvilinear, levelling off at about 8 y of age. Productivity declined significantly among birds older than 8 y; this decline was not due to events in the last year of breeding (“terminal illness”). Breeding performance increased with duration of pair‐bond at least through y 5. Early breeding was significantly related to age and duration of pair‐bond. Most of these relationships were stronger among males than among females, and many of them were not significant when females were considered alone. After controlling for other factors, breeding performance varied significantly among birds, but autocorrelations were low and limited to intervals of one year. Parental quality (defined for birds studied in six or more years as the individual bird term in a GLM for productivity controlling for other factors) was not correlated with lifespan or other demographic parameters, but high‐quality birds were less prone to change mates and burrows than low‐quality birds. We know of no previous study in which simultaneous effects of laying date, age and pair‐bond duration on breeding performance were measured, while controlling for year, individual quality and terminal illness.

Do great tits rely on inadvertent social information from blue tits? A habitat selection experiment

Sympatric species sharing requirements are competitors, but recent evidence suggests that heterospecifics may also be used as a source of information. The heterospecific habitat copying hypothesis proposes that individuals of one species might use information inadvertently produced by the breeding performance of individuals of other species to assess habitat quality whenever the two species share needs. In this study, we provide the first experimental test of this hypothesis by examining whether the manipulated reproductive success of blue tits (Cyanistes caeruleus) is used as heterospecific inadvertent social information (ISI) in breeding-habitat selection by sympatric great tits (Parus major). The reproductive success of blue tits was manipulated 1year at the scale of patches by transferring nestlings from decreased to increased patches. No evidence was found of great tits using the reproductive success of blue tits as a source of heterospecific ISI. However, dispersal decisions by adult great tits correlated with information on con- and heterospecific densities, which constitute other sources of ISI. As density and breeding performance are tightly intertwined forms of information, the difficulty in distinguishing between them might lead great tits to use heterospecific ISI more in the form of density than breeding performance when making dispersal decisions.

Breeding performance, mate fidelity, and nest site fidelity in a long‐lived seabird: behaving against the current?

There is evidence that breeding failure is associated with divorce and dispersal in many bird species. However, deviations from the general pattern “success‐stay/failure‐leave” seem to be common, suggesting that factors other than breeding performance may importantly influence mate and habitat selection. Moreover, variability in response to performance suggests coexistence of different evolutionary strategies of mate and site selection within a population. In this study, we assessed how individuals conform to the success‐stay/failure‐leave pattern in kittiwakes (Rissa tridactyla), and aimed to identify categories of individuals presenting different behavioural patterns. We considered individual attributes (experience, prior residence at the nest site, performance in multiple breeding attempts), pair attributes (arrival asynchrony, timing of failure, pair duration), and productivity in habitat patches. Timing of failure was an important factor. Pair reunion probability was close to 0.5 in failed pairs, but it was consistently higher in early failed than in late failed pairs. Prior residence better explained variability in probability of reunion in failed pairs than pair duration. However, the positive influence of prior residence on the probability of reunion was perceptible only in early failed pairs. Divorce probability in successful pairs increased with arrival asynchrony, and was higher in first‐time than in experienced breeders. Local productivity positively influenced site fidelity probability in early failed birds, but not in late failed ones. Using memory models, we found that dispersal decisions integrate information on individual breeding performance in a temporal scale longer than one year. This study contributed to the identification of relevant states to be considered when addressing mate and nest site choice. Natural selection may operate on slight fitness differences that cannot be detected without high levels of stratification according to the appropriate individual and habitat attributes.

Phenotypic assortative mating and within-pair sexual dimorphism and its influence on breeding success and offspring quality in Magellanic penguins

We examined within-pair sexual dimorphism and phenotypic assortative mating in Magellanic penguins (Spheniscus magellanicus) breeding in six colonies located on the Patagonian coast (Argentina). All measured phenotypic traits except the number of pectoral spots differed between the sexes; bill depth and flipper length were the most and least dimorphic traits, respectively. We found assortative mating by bill depth and body mass. The similarity in body condition within pairs was close to significant. When we performed separate correlations for birds that bred successfully, i.e., raised one or two offsprings, and birds that did not attempt to breed or bred unsuccessfully, only the successful breeders showed assortative mating by body mass. In addition, we attempted to relate the body size of each member of the pair and the degree of sexual dimorphism within pairs to the breeding performance of individuals, which was measured as brood size, and body condition and immunocompetence of offspring. We found that pairs that were less dimorphic in flipper length raised more offspring. This effect was due to female flipper length per se and not to the relative difference in flipper length between members of the pair. Females with larger flippers had a higher probability of raising two chicks. No effects of body measurements or degree of sexual dimorphism on body condition or T-cell-mediated immune response of offspring were found. We discuss these results in the context of potential factors responsible for the maintenance of sexual size dimorphism in this species.

CONSPECIFIC REPRODUCTIVE SUCCESS AND BREEDING HABITAT SELECTION: IMPLICATIONS FOR THE STUDY OF COLONIALITY

Habitat selection is a crucial process in the life cycle of animals because it can affect most components of fitness. It has been proposed that some animals cue on the reproductive success of conspecifics to select breeding habitats. We tested this hypothesis with demographic and behavioral data from a 17-yr study of the Black-legged Kittiwake (Rissa tridactyla), a cliff-nesting seabird. As the hypothesis assumes, the Black-legged Kittiwake nesting environment was patchy, and the relative quality of the different patches (i.e., breeding cliffs) varied in time. The average reproductive success of the breeders of a given cliff was predictable from one year to the next, but this predictability faded after several years. The dynamic nature of cliff quality in the long term is partly explained by the autocorrelation of the prevalence of an ectoparasite that influences reproductive success. As predicted by the performance-based conspecific attraction hypothesis, the reproductive success of current breeders on a given cliff was predictive of the reproductive success of new recruits on the cliff in the following year. Breeders tended to recruit to the previous year’s most productive cliffs and to emigrate from the least productive ones. Consequently, the dynamics of breeder numbers on the cliffs were explained by local reproductive success on a year-to-year basis. Because, on average, young Black-legged Kittiwakes first breed when 4 yr old, such a relationship probably results from individual choices based on the assessment of previous-year local quality. When breeders changed breeding cliffs between years, they selected cliffs of per capita higher reproductive success. Furthermore, after accounting for the potential effects of age and sex as well as between-year variations, the effect of individual breeding performance on breeding dispersal was strongly influenced by the average reproductive success of other breeders on the same cliff. Individual breeding performance did not appear to influence the probability of dispersing for birds breeding on cliffs with high local reproductive success, whereas individual breeding performance did have a strong effect on dispersal for birds that bred on cliffs with lower local reproductive success. This suggests that the reproductive success of locally breeding conspecifics may be sufficient to override an individual’s own breeding experience when deciding whether to emigrate. These results, which are supported by behavioral observations of the role of prospecting in recruitment, suggest that both first breeders and adults rely on the reproductive success of conspecifics as “public information” to assess their own chances of breeding successfully in a given patch and to make settling decisions. A corollary prediction is that individuals should attempt to breed near successful conspecifics (a form of social attraction) in order to benefit from the same favorable local environmental conditions. Such a performance-based conspecific attraction mechanism can thus lead to an aggregative distribution of nests and may have played a role in the evolution of coloniality.

Conspecific Reproductive Success and Breeding Habitat Selection: Implications for the Study of Coloniality

Habitat selection is a crucial process in the life cycle of animals because it can affect most components of fitness. It has been proposed that some animals cue on the reproductive success of conspecifics to select breeding habitats. We tested this hypothesis with demographic and behavioral data from a 17-yr study of the Black-legged Kittiwake (Rissa tridactyla), a cliff-nesting seabird. As the hypothesis assumes, the Black-legged Kittiwake nesting environment was patchy, and the relative quality of the different patches (i.e., breeding cliffs) varied in time. The average reproductive success of the breeders of a given cliff was predictable from one year to the next, but this predictability faded after several years. The dynamic nature of cliff quality in the long term is partly explained by the autocorrelation of the prevalence of an ectoparasite that influences reproductive success. As predicted by the performance-based conspecific attraction hypothesis, the reproductive success of current breeders on a given cliff was predictive of the reproductive success of new recruits on the cliff in the following year. Breeders tended to recruit to the previous year’s most productive cliffs and to emigrate from the least productive ones. Consequently, the dynamics of breeder numbers on the cliffs were explained by local reproductive success on a year-to-year basis. Because, on average, young Black-legged Kittiwakes first breed when 4 yr old, such a relationship probably results from individual choices based on the assessment of previous-year local quality. When breeders changed breeding cliffs between years, they selected cliffs of per capita higher reproductive success. Furthermore, after accounting for the potential effects of age and sex as well as between-year variations, the effect of individual breeding performance on breeding dispersal was strongly influenced by the average reproductive success of other breeders on the same cliff. Individual breeding performance did not appear to influence the probability of dispersing for birds breeding on cliffs with high local reproductive success, whereas individual breeding performance did have a strong effect on dispersal for birds that bred on cliffs with lower local reproductive success. This suggests that the reproductive success of locally breeding conspecifics may be sufficient to override an individual’s own breeding experience when deciding whether to emigrate. These results, which are supported by behavioral observations of the role of prospecting in recruitment, suggest that both first breeders and adults rely on the reproductive success of conspecifics as “public information” to assess their own chances of breeding successfully in a given patch and to make settling decisions. A corollary prediction is that individuals should attempt to breed near successful conspecifics (a form of social attraction) in order to benefit from the same favorable local environmental conditions. Such a performance-based conspecific attraction mechanism can thus lead to an aggregative distribution of nests and may have played a role in the evolution of coloniality.

Régulation par le recrutemet, la fidélité et la non-reproduction chez un oiseau colonial, la mouette tridactyle (Rissa tridactyla)

A long term study of the Kittiwake colonies of Cap Sizun (Brittany, France) shows that, even though Kittiwakes usually recruit at the age of 4, the population growth rate from year to year is highly correlated with the breeding success of the previous year. This is due to a strong recruitment, a large proportion of breeders, and a high adult fidelity after years of high fecondity. Furthermore, for a given individual breeding performance (breeding success or failure), adult fidelity to the colony and the proportion of breeders in the following year increase with the breeding success of the colony in which they bred in the previous year. This leads us to propose a mechanism of population regulation through Attraction, Recruitment, Non-breeding, and Adult Fidelity, in which breeding success and social context — two characteristics which reflect environmental quality — play prominent roles. These two parameters are the main components of the attraction exerted by a breeding site, i.e. its capacity to recruit new breeders (young or adults) and to retain its own adults as breeders. Breeding success and social context are assessed by potential recruits when prospecting different breeding places. If fecondity and social context are good, prospectors may settle there and recruit in the following year. Prospectors are attracted to nests with chicks, on which they squat when they are momentarily left unattended by the parents after the middle of the rearing period. These squatters are mainly prebreeders or failed breeders ; both are actively looking for a breeding site. Males are more prone to squat on chicks. Prebreeding squatters on chicks show a high proportion of recruitment in the following year : 34 % of prospecting males recruit on one of the nests they have squatted during the previous year, and 24 % on the most consistently squatted nest. Thus, squatting is an appropriation behaviour of a breeding site. The density of parasites influences the colony breeding success : breeding groups with high densities of parasites on the chicks have the lowest fecondity. The appearence of parasites is correlated with a drop in fecondity of the kittiwakes and parasites are likely to be the cause of this reduced fecondity. This acts on the numbers through prospectors’ and adults’ behaviour which in turn can explain the positive correlation between the fecondity in a given year and the rate of population increase in the following year.

Diet Choice in the Herring Gull: Constraints Imposed by Reproductive and Ecological Factors

Studies of diet choice in carnivorous animals typically concentrate on maximizing rates of energy intake, and rarely emphasize either constraints on choice or alternate currencies. We examined diet choice in a generalist carnivore, the Herring Gull, Larus argentatus, in relation to both nesting habitat and reproductive performance over a 2—yr period. During prelaying and incubation periods 75—80% of Herring Gulls specialized on either intertidal organisms, human refuse, or other seabirds, while only 20—25% had generalized diets. Specific types of specialists tended to nest in particular habitats. Foraging tactics associated with each of three diets were related to time budgets and ecological constraints, i.e., levels of predation or intraspecific competition in specific habitats. Variation in diet choice was also strongly related to individual breeding performance. Intertidal specialists laid eggs earlier, produced larger and heavier clutches, and had higher rates of hatching than generalists and other specialists. This pattern held true over both years of the study even though weather conditions, and food accessibility, differed between years. Despite differences in performance related to laying and hatching, there were no significant differences in fledging rates, either between years or among diets. These results suggest that (a) choice of nesting habitat may constrain diet choice, (b) diet choice may have major effects on breeding performance, and (c) energy may not be the proper currency for examining diet choice in carnivores subject to a variety of constraints.