Individual changes underlie age-specific pattern of laying date and egg-size in female common terns Sterna hirundo

It has been suggested that breeding performance differs between young and old birds due to the appearance and disappearance of phenotypes through differential survival (selection hypothesis) or differential recruitment (delayed breeding hypothesis) of high-quality individuals, but each bird may show constant breeding performance over its life. We tested constant egg-volume and laying date by modelling their variability on the basis of the 109 known-age females of common tern Sterna hirundo with data available from 1 to 9 years. Longitudinal analyses showed a significant advancement of laying date, as well as a steady increase in egg-volume, in young age classes from 2 to 5–7 years old, indicating individual intrinsic changes in performance with age. In our model, female effect accounted for 74% and 8% of variance in egg-volume and laying date, respectively, suggesting that if correlation between breeding performance and survival or recruitment exists, population patterns of age-specific performance may emerge. However, we found no evidence that birds that did not return to breed during young age classes laid later or smaller eggs than returned breeders. Likewise, we found no evidence that recruiting birds laid earlier or larger eggs than same aged birds recruited in preceding years. Thus, this study shows that age-specific patterns in timing of breeding and egg-size in common terns result from individuals intrinsic changes, and we reject the selection and the delayed breeding hypotheses as a major factor shaping age-specific patterns at population level.Communicated by F. Bairlein     

Resources and reproductive performance in Mute swans Cygnus olor

Around Oxford, breeding pairs of mute swans maintained territories throughout most of the year and these territories varied considerably in potential food availability in the form of aquatic vegetation, bread supplied by the public and pasture for grazing. On territories with abundant aquatic vegetation females laid earlier larger clutches than on those with little aquatic vegetation. Clutches were also larger on territories with a high potential bread supply. In addition, adult females were heavier on territories with a high diversity of aquatic vegetation. However, contrary to expectation, there were no direct relationships between territory quality measures and other aspects of reproductive performance including number of young fledged, egg volume, cygnet weight or size, or adult male weight. Instead, the number of young fledged was related to clutch size; egg volume was related to laying date; and cygnet weight was related to age and egg volume. Adult male weight was positively related to age.     

Seasonal changes in male and female bill knob size in the mute swan Cygnus olor

Sexual selection theory predicts that sexually selected ornaments are costly to maintain and, as condition-dependent signals, are likely to vary in attractiveness with season and age. Mute swans Cygnus olor possess a black, fleshy knob at the base of the bill, which is present in both sexes. Using measures calculated from digital photographs taken over two years we monitored changes in the size of the bill knob in individual swans throughout the breeding season. Our longitudinal data show that bill knob size is highly dynamic. Relative bill knob size was larger in males than females and was consistently greater for breeding males than for non-breeders. For males, relative bill knob size peaked during cygnet hatching, when male protection of the brood is most important, and was smallest during moult. In females, breeders had larger bill knobs than non-breeders at all times apart from immediately after egg-laying and incubation, when the reverse was true, presumably reflecting the costs of reproduction. Body mass was a highly significant predictor of relative bill knob size in both sexes, as was age, with an initial increase and then later a decline in relative ornament size across the lifetime of male birds. The bill knob ornament in mute swans thus appears to be a condition-dependent, highly malleable trait. It accurately reflects the differing pressures experienced by individual birds as they progress through the breeding season, suggesting selection by both intra- and inter-sexual forces.     

Coexistence and population genetic structure of the whooper swan <Emphasis Type="Italic">Cygnus cygnus</Emphasis> and mute swan <Emphasis Type="Italic">Cygnus olor</Emphasis> in Lithuania and Latvia

Two closely related swan species, the mute swan Cygnus olor and the whooper swan Cygnus cygnus, were formerly allopatric throughout their breeding ranges, but during the last decades a sympatric distribution has become characteristic of these species in the Baltic Sea region. The whooper swan has gradually replaced the mute swan in many suitable habitats in Lithuania and Latvia. Marked differences in the genetic population structure of both species may partially explain the dominance of the whooper swan, as genetic population divergence can be a major factor affecting inter-specific competition. A homogenous genetic population structure was defined for mute swans breeding in Lithuania, Latvia, Poland and Belarus. Breeding mute swans in this region are mostly of naturalised origin. A diverse population genetic structure characterizes whooper swans breeding in Lithuania and Latvia.     

Effects of mute swans on wetlands: a synthesis

The increases in mute swan (Cygnus olor Gmelin) population size have caused concern among stakeholders, who sometimes consider it as a pest species. Here, we aim to review existing studies on the ecological effects that mute swans have on wetlands. Claim that mute swans threaten other waterbirds were partly supported: mute swans sometimes behave territorially towards conspecifics and other waterbird species, but this does not systematically occur. A second common claim, that mute swans damage aquatic plant beds, was upheld in that the species did indeed affect aquatic plant communities in several studies. However, grazing by mute swans does not systematically have negative effects on aquatic plants. Habitat patch size, distance between habitat patches, resource availability and water velocity affect habitat selection process by mute swans, with varying effects depending on season and mute swan breeding status. Scientific knowledge does not support the idea that mute swan population increase can be considered as a biological invasion in Europe. Conversely, there is a genuine risk of biological invasion in North America. In light of the literature review, we discuss the relevance of mute swan population management in Europe and in North America, and propose future research avenues.     

Breeding mute swan habitat selection when accounting for detectability: a plastic behaviour consistent with rapidly expanding populations

A number of native and exotic animal species show dramatic population increases in terms of both numbers and geographic range. Understanding the habitat selection processes behind such increases is crucial to implement adequate management measures. Mute swan (Cygnus olor) populations have experienced a tremendous demographic and geographic expansion in Western Europe during the twentieth century, colonizing a wide variety of aquatic habitats. We aimed at assessing how swans select nesting sites during the pre-laying and laying periods on medium to large fishponds (from 10 to 50 ha) in Eastern France, while accounting for detectability biases and testing for the effects of fishpond spatial configuration, vegetation resources, human disturbance and habitat management. Our results demonstrate that the mute swan is a non-selective species regarding its nesting habitat among such fishponds, using these independently from the parameters considered although fishpond characteristics varied. Although mute swan is one of the least cryptic Anatidae, owing to its white colour and large size, detection of breeding pairs remained imperfect for each over several sampling occasions. However, because we repeated the sampling sessions, detection of swan pairs by the end of the monitoring period was as high as 0.94. These results are consistent with previous assertions that the mute swan is a species of high ecological plasticity, which may partly explain its recent colonization rates. Given that even swan breeding events were imperfectly detected on each occasion, we highlight the fact that most studies of breeding ducks (which are more cryptic) would be considerably improved by better considering detection biases.     

Geographical variation in reproductive ageing patterns and life‐history strategy of a short‐lived passerine bird

We investigated differences in ageing patterns in three measures of breeding performance in populations of barn swallows Hirundo rustica L. from Spain and Denmark differing in breeding latitude and hence migration distance and duration of the breeding season. We found differences in ageing patterns between populations. Generally, young (i.e. yearling) and old females (i.e. ≥ 5 years of age) laid their first eggs later and produced smaller clutches than middle‐aged females (i.e. 2–4 years of age) in both populations. The southernmost population (i.e. Spanish) showing the shorter migratory distance experienced a greater within‐individual increase in timing of breeding and clutch size in early life and a greater within‐individual decrease in laying date but not in clutch size during senescence compared with the northernmost population (i.e. Danish). We also found that the number of fledglings produced annually was related to the age of the two members of the breeding pairs with pairs composed of young and old females performing less well than breeding pairs composed of middle‐aged females. We did not find reproductive senescence for the age of the male while controlling for the age of the female on the number of fledglings produced annually by the breeding pair. Differential survival between individuals did not explain age effects on laying date or annual clutch size in neither population. However, the increase in the number of fledglings produced annually with age was partly explained by the disappearance of poor‐quality members of the pairs, mainly poor‐quality males. Age‐related breeding success (i.e. number of fledglings) was similar for barn swallows from Spain and Denmark. Therefore, the study of ageing patterns and life‐history strategies in free‐ranging animals from more than a single population can throw new light on life‐history theory, population dynamics and evolutionary studies of senescence.     

Maturation increases early reproductive investment in Adélie Penguins Pygoscelis adeliae

Correlations between female investment in egg production and age, breeding experience and laying date have been reported in several seabird species. In general, clutch and egg sizes increase with female age and breeding experience but decrease with laying date. Positive correlations of clutch and egg size with age and breeding experience can be caused by an increase in reproductive investment with maturation or they may be an artefact of lower survival rates for individuals with poor-quality phenotypes. Negative correlations of clutch and egg size with laying date might signal an adaptive reduction in egg production or be due in part to variation among individuals. We examined the interactions of female age, breeding experience, laying date and clutch and egg size in Adélie Penguins Pygoscelis adeliae . Breeding experience strongly affected clutch size with 87.3% of all one-egg clutches laid by first-time breeders. In addition, increasing age had a positive influence on egg size and was associated with earlier laying dates. However, there was little evidence to suggest that either clutch or egg sizes are influenced by laying date. Laying dates and clutch and egg sizes did not affect a female's probability of returning to breed in the following year, indicating that increased investment is a product of maturation and not of the loss of poor-quality breeders from the population. Our results suggest that as female Adélie Penguins gain foraging and breeding experience they are able to initiate breeding earlier, to lay complete clutches of two eggs and to lay larger eggs.     

Age-related fledgling production in great cormorants Phalacrocorax carbo: influence of individual competence and disappearance of phenotypes

Age-related reproductive performance of great cormorants Phalacrocorax carbo sinensis was studied in a tree nesting colony in Denmark in relation to age-related improvements of competence and progressive disappearance of phenotypes. Within-individual changes in fledgling production were measured, and cross-sectional analyses were applied. The within-individual analyses showed that competence improved with age and/or that individuals showed restraint to optimize their reproductive effort. The within-individual improvements were three to six times higher among individuals that survived and returned to breed beyond the fourth breeding attempt than among individuals disappearing from the breeding population before the fourth breeding attempt. Taking this into account the within-individual improvements explained 70–90% of the age-effect observed in the population over the youngest ages. Effects of breeding experience were significant for females, but only within the group of individuals that were present in the breeding population beyond the age of five years. In males, improvements arose because of unknown factors related to age. Individual great cormorants that bred beyond the age of five years had higher reproductive success, on average, than birds disappearing from the breeding population earlier in life. This supports the differential survival hypothesis. However, the effect on the population mean was partly counterbalanced by late recruitment of other inferior breeders. It is concluded that the enhancement in fledgling production with increasing age was primarily an effect of age-related improvements of competence and secondly an effect of progressive disappearance of phenotypes.     

Effects of seasonality,isolation and patch quality for habitat selection processes by mute swans Cygnus olor in a fishpond landscape

Foragers in patchy environments do not only select sites for single patch characteristics, but also have to consider the local environment of such patches. We studied habitat selection by mute swans Cygnus olor in a wide and heterogeneous fishpond region (the Dombes, eastern France). In this study, we considered fishpond isolation, resource quality within fishponds and breeding status of mute swans during both summer and winter. Mute swans did not select aquatic habitat randomly within the landscape. During summer, the population spread preferentially on medium to large fishponds, in subregions with numerous or closely related waterbodies, without generating a clumped distribution of birds. In addition to a positive effect of local fishpond number (2 km radius), breeding birds also responded positively to fishpond size. Non‐breeders selected fishponds mainly according to their size. Intraspecific territoriality did not appear to limit the presence of non‐breeders (i.e. moulting flocks), since both breeders and non‐breeders could coexist on the larger fishponds. During winter, mute swans used medium to large reflooded fishponds after summer drainage. The surrounding aquatic environment of fishponds played a minor role in determining flocking, compared to actual patch quality. Flocking occurred on large fishponds that had reflooded after having dried the summer before, whatever the agricultural cultivation practiced in the summer following drainage. The results suggest that geographical aspects should be taken into account when considering the potential impact of this expanding species within such ecosystems, and also in more general management policies dealing with aquatic habitats for waterbird populations.     

Reproductive performance of little penguins Eudyptula minor in relation to year, age, pair-bond duration, breeding date and individual quality

We measured breeding performance of little penguins Eudyptula minor at 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.     

Age effects on clutch size and laying dates of individual female Lesser Snow Geese Anser caerulescens

It is well known in many species of birds that mean clutch size increases and laying date advances with increasing age of the female. This has been interpreted widely as being due to some maturation process in which the performance of individual birds improves as they age. There are two alternative explanations, however. Birds which lay small clutches and lay late may have a higher mortality and be under-represented in the later year samples, or birds with a propensity to lay large and early clutches may enter the breeding population at a later age. In an attempt to discriminate between these alternatives, the reproductive performances of individual Snow Geese were examined over a series of years. Clutch size increased and laying date advanced for these individuals as it did for the population as a whole, thus confirming the maturation hypothesis.     

Costs and Benefits of Subadult Plumage in Mute Swans: Testing Hypotheses for the Evolution of Delayed Plumage Maturation

In some avian species, young birds capable of reproducing diminish their prospects of doing so by molting into a subadult plumage that accurately signals their subadult status. Several hypotheses have been proposed to explain the evolution of delayed plumage maturation, but testing them usually has involved interspecific comparisons that are hard to interpret. Mute swans (Cygnus olor) exhibit two phenotypes that differ in whether the birds have a gray subadult plumage (SAP phenotype) or molt immediately into an all white adult plumage (AP phenotype). The AP phenotype results from a recessive allele on the X chromosome; both phenotypes occur in the same population and even in the same brood. We compared costs and benefits of both phenotypes in mute swans on the Chesapeake Bay in 1972-1980 and on Long Island Sound in 1982-1989. Swans with the SAP phenotype had higher survival rates from hatching to fledging than AP swans. In the fall, when AP cygnets began to molt into their white plumage, their parents often attacked and drove them off while allowing SAP cygnets from the same brood to remain on their territories for several more months. SAP males had higher survival rates during their first 2 yr of life than AP males, but AP swans bred at a younger age than SAP swans. The only proposed hypothesis for the evolution of delayed plumage maturation that can explain its occurrence in mute swans is the status-signaling hypothesis. This hypothesis argues that males with subadult plumage honestly advertise their age and subordinate status while AP swans are cheaters and engaging in dishonest communication. SAP males acquire a longer period of parental care, suffer less aggression from older birds, and increase their survival but forgo the opportunity to breed at an early age. This is a unique example of how a single gene resulted in either honest or dishonest communication, changed a bird's relationship with its parents and potential mates, and altered the bird's chances to survive and to reproduce.     

FACTORS AFFECTING THE AGE OF FIRST BREEDING OF THE KITTIWAKE RISSA TRIDACTYLA

At a Kittiwake colony in Northumberland, 80% of those birds which returned to their natal colony to breed were males and these supplied 52% of all male recruits. More females breed away from their natal colony than males. There was no differences in the proportions of young fledged from sites in the centre or at the edge of the colony, or by parents of different experience, which returned to breed. Kittiwakes breed for the first time at ages from 3 to 8 years, but most at 4 or S years old. Males arrive back at the colony at an earlier age than females and breed for the first time one year earlier. Males obtaining sites at the centre of the colony first breed at an earlier age than those at the edges. Neither the age nor the area of first breeding appear to be transmitted from parent to offspring. Males breeding first aged 4 years or younger produced more young than those which first bred aged 5 years or older, despite their partners laying smaller clutches. This difference was most marked among those males recruited to sites in the centre of the colony. The advantage of this earlier breeding is counteracted by a lower survival rate among those males which start to breed at the younger ages. In all breeding Kittiwakes, annual reproductive output increases with experience while annual survival rates decrease. Once they had started to breed, many birds failed to breed in one subsequent season. Nearly 60% of these cases of intermittent breeding occurred in the year following first breeding. Intermittent breeding was most frequent among young birds and among females. It is suggested that each breeding involves a cost to the individual in terms of reduced survival, and that deferred and intermittent breeding are means of guarding survival. A model is proposed whereby the age at which a bird starts to breed, the nesting site which it obtains, and its subsequent breeding strategy result in each individual producing an optimal number of reproducing offspring in its lifetime, relative to its quality.     

The Effects of Birth Weight and Maternal Care on Survival of Juvenile Steller Sea Lions (Eumetopias jubatus)

Steller sea lions were listed as endangered following a collapse of the western distinct population beginning in the late 1970s. Low juvenile survival has been implicated as a factor in the decline. I conducted a multistate mark-recapture analysis to estimate juvenile survival in an area of the western population where sea lions are showing signs of recovery. Survival for males and females was 80% between 3 weeks and 1 year of age. Approximately 20% of juveniles continued to be nursed by their mothers between ages 1 and 2 and 10% between ages 2 and 3. Survival for juveniles that suckled beyond 1 year was 88.2% and 89.9% to ages 2 and 3, respectively. In contrast, survival for individuals weaned by age 1 was 40.6% for males and 64.2% for females between ages 1 and 2. Birth mass positively influenced survival for juveniles weaned at age 1 but had little effect on individuals continuing to suckle. Cumulative survival to age 4 was double that estimated during the population decline in this region. Evidence suggests that western Steller sea lions utilize a somewhat different maternal strategy than those in the eastern distinct population. Western adult females generally invest more in their pups during the first year but wean offspring by age 1 more often. This results in better survival to age 1, but greater mortality between ages 1 and 3 compared to the eastern population. Different maternal strategies may reflect density dependent pressures of populations at opposite levels of abundance.     

No effect of partner age and lifespan on female age‐specific reproductive performance in blue tits

Studies of age‐specific reproductive performance are fundamental to our understanding of population dynamics and the evolution of life‐history strategies. In species with bi‐parental care, reproductive ageing trajectories of either parent may be influenced by their partner's age, but this has rarely been investigated. We investigated within‐individual age‐specific performance (laying date and number of eggs laid) in wild female blue tits Cyanistes caeruleus and evaluated how the age and longevity of their male partner indirectly influenced the females’ reproductive performance. Females showed clear age‐dependence in both laying date and number of eggs laid. We found that female reproductive performance improved in early life, before showing a decline. Longer‐lived females had an earlier laying date throughout their lives than shorter‐lived females, but there was no difference in number of eggs laid between longer‐ and shorter‐lived females. Within breeding pairs, the female's (age‐specific) reproductive performance was not dependent on the age and longevity of the male partner. We conclude that the age and quality of the male partner may be of little importance for traits that are under direct female control.     

Life history of breeding partners alters age‐related changes of reproductive traits in a natural population of blue tits

Reproductive senescence, an intra‐individual decline in reproductive function with age, is widespread, but proximate factors determining its rate remain largely unknown. Most studies of reproductive senescence focus on females, leaving senescence in male function and its implications for female function largely understudied. We constructed linear mixed models to explore the interactive effects of paternal and maternal age and a life‐history trait (i.e. age at first reproduction) on four fitness components (i.e. laying date, clutch size, number of fledglings and number of recruits) measured in a wild, breeding population of blue tits Cyanistes caeruleus ogliastrae where individual breeding success has been followed for over 30 years (our dataset spanned 29 years). Previous studies have shown that, across female lifespan, laying date decreases and subsequently increases; earlier laying dates result in higher fitness because hatchlings have greater access to a seasonal food source. Our analyses reveal that females that initiate reproduction early in life show a greater delay in laying date with old age. In addition to delayed laying dates, older females lay smaller clutches. However, the magnitude of female age effects was influenced by the age at first reproduction of their breeding partners. Senescence of laying date and clutch size was reduced when females mated with males that reproduced early in life compared to males that delayed reproduction. We confirmed that both laying date and clutch size were significantly correlated with reproductive fitness suggesting that these dynamics early in the breeding cycle can have long‐term consequences. These complex phenotypic interactions shed light on the proximate mechanisms underlying reproductive senescence in nature and highlight the potential importance of cross‐sex age by life‐history interactions.     

Improvement of reproductive performance with age and breeding experience depends on recruitment age in a long‐lived seabird

Reproductive success increases with age in many species, however, the underlying mechanisms remain unclear and several hypotheses have been proposed to explain age‐related improvements in reproductive output. In this contribution we investigated the effects of age, recruitment age, breeding experience and sex on reproductive performance during the early breeding career in the common tern Sterna hirundo using long‐term individual‐based data. We used measurements of performance, which spanned the entire breeding process: clutch size, hatching success, fledging success and fledglings per pair. Longitudinal analyses within individuals showed a clear increase with age in all performance measures. Furthermore, a significant change in reproductive performance was found between first time‐ and experienced breeders. Recruitment age had a strong influence on hatching and fledging success: two‐year‐old recruits had significantly lower reproductive success than birds which recruited at older ages, but the increase in breeding performance with experience was stronger in young recruits. Comparing age and experience effects, age effects were more pronounced during the first breeding attempts, whereas experience effects were also visible in subsequent breeding attempts. The degree of intra‐individual improvements in reproductive performance is due to a complex interplay of age at first breeding and experience. The results strongly support the constraint hypothesis.     

Experience versus effort: what explains dynamic heterogeneity with respect to age?

Age‐related patterns of survival and reproduction have been explained by accumulated experience (‘experience hypothesis’), increased effort (‘effort hypothesis’), and intrinsic differences in phenotypes (‘selection hypothesis’). We examined the experience and effort hypotheses using a 40‐year data set in a population of Leach's storm‐petrels Oceanodroma leucorhoa, long‐lived seabirds for which the effect of phenotypic variation has been previously demonstrated. Age was quantified by time since recruitment (‘breeding age’). The best model of adult survival included a positive effect of breeding age (1, 2, 3+ years), sex (male > female), and year. Among‐individuals variation (fixed heterogeneity) accounted for 31.6% of the variance in annual reproductive success. We further examined within‐individual patterns in reproductive success (dynamic heterogeneity) in the subset of individuals with at least five breeding attempts. Three distinct phases characterized reproductive success – early increase, long asymptotic peak, late decline. No effect of early reproductive output on longevity was found, however, early success was positively correlated with lifetime reproductive success. Reproductive success was lower earlier than later in life. Among the few natally philopatric individuals in the population, age of first breeding had no effect on longevity, lifetime reproductive success, or early reproductive success. No support for the effort hypothesis was found in this population. Instead, age‐specific patterns of survival and reproduction in these birds are best explained by the experience hypothesis over and above the effect of intrinsic differences among individuals.     

Parental age and lifespan influence offspring recruitment: a long-term study in a seabird

Recent studies of wild populations provide compelling evidence that survival and reproduction decrease with age because of senescence, a decline in functional capacities at old ages. However, in the wild, little is known about effects of parental senescence on offspring quality. We used data from a 21-year study to examine the role of parental age on offspring probability of recruitment in a long-lived bird, the blue-footed booby (Sula nebouxii). Offspring probability of recruiting into the breeding population varied over the life of parents and effects age were similar in mothers and fathers. Offspring recruitment was high when parents were roughly 6-12 years old and low before and after then. Effects of parental age on offspring recruitment varied with lifespan (parental age at last reproduction) and previous breeding experience. Offspring recruitment from young and old parents with long reproductive lifespans was greater than that of offspring from parents with short lifespans at young and old ages. For parents with little previous breeding experience recruitment of offspring decreased with their hatch date, but experienced parents were no similarly affected. We found evidence of terminal effects on offspring recruitment in young parents but not in older parents, suggesting that senescence is more likely a gradual process of deterioration than a process of terminal illness. Failure to recruit probably reflects mortality during the first years after independence but also during the fledgling transition to full independence. Our results show effects of parental age and quality on offspring viability in a long-lived wild vertebrate and support the idea that wild populations are composed of individuals of different quality, and that this individual heterogeneity can influence the dynamics of age-structured populations.