Modelling the population consequences of age- and sex-related differences in winter mortality in the oystercatcher, Haematopus ostralegus

A modelling approach is used to explore the effect of age and sex differences in oystercatcher ( Haematopus ostralegus ) winter mortality on population size, population structure and the population response to habitat loss or change. Increasing the mortality of first and second year birds reduced population size, but had very little effect on the proportion of the population that were adults. Increasing female mortality reduced population size and resulted in a male-biased population. A sex bias amongst birds of breeding age meant that there were fewer potential breeding pairs for a given population size, reducing the size of the breeding population and the breeding output. Increasing the mortality of one sex relative to the other reduced population size, even when mean adult mortality rates remained unchanged. Increasing the strength of density-dependent mortality in young birds caused a greater reduction in population size as habitat was lost. Increasing the strength of female density-dependent mortality had the same effect, even though male density-dependent mortality had been correspondingly reduced. Increasing density-independent or density-dependent winter mortality in one sex relative to another also exaggerated the disproportional effect of winter habitat loss on separate breeding subpopulations using the same overwintering area. These results suggest that any study of population dynamics should be aware of both age and sex differences in mortality. Conservationists should be particularly aware of any age or sex differences in diet or habitat use that may result in a differential response to environmental change.     

Effects of food abundance, density and climate change on reproduction in the sparrowhawk Accipiter nisus

The reproductive success of predators depends on abiotic environmental conditions, food abundance and population density, and food abundance, density and their interactions may respond to changes in climatic conditions. Timing of reproduction by five of the eight numerically most common prey of the sparrowhawk Accipiter nisus advanced significantly since 1971, during a period of temperature increase. There was no evidence that mean laying date or any other reproductive parameter of sparrowhawks changed consistently during the study period 1977–1997. Laying date advanced and percentage of unsuccessful female sparrowhawks decreased with beech mast in the current year, an index of food abundance for avian prey. Mean laying date of sparrowhawks was advanced in warmer springs, and although mean clutch size was not larger in warm than in cold springs, mean brood size of successful pairs and breeding success increased in such springs, showing that sparrowhawks enjoyed a fitness gain when reproducing early. The timing of sparrowhawk reproduction with respect to the peak in abundance of fledgling prey increased, from a good match between mean timing of fledging by prey and maximum demand for food by the predator in 1977, to reproduction occurring later than the peak in fledging prey availability in 1997. The size of the breeding population of sparrowhawks was not predicted by mean spring temperature, the size of the breeding population the previous year or beech mast crop. The size of the post-breeding population was predicted by size of the breeding and post-breeding population the previous year and by the proportion of unsuccessful females the current year. These findings imply that sparrowhawks did not respond to change in climate, although climate changed the timing of reproduction by the main prey species.     

Ecological and phenological covariates of offspring sex ratio in barn swallows

Non-random sex allocation in relation to parental, ecological and phenological factors has been investigated in several correlational studies of birds, mostly based on few breeding seasons and relatively small sample sizes, which have led to different results. We investigated sex ratio of nestling barn swallows (Hirundo rustica) in relation to adult sex ratio, laying date, clutch size, colony size and meteorological conditions in a sample of 553 broods (>2200 nestlings) during 10 years. At the population level, nestling sex ratio varied among years and deviated from parity in two years. Sex ratio among adults did not predict offspring sex ratio in the current or the following year. At the within-family level, the proportion of sons increased with laying date in large clutches, did not vary among clutches of intermediate size, and tended to decline with laying date in small clutches. Large colonies harbored more sons. The proportion of males increased with temperature during laying whereas the effects of temperature during the pre- or post-laying periods and that of rainfall were non-significant. These patterns of variation of offspring sex ratio did not differ between years. Thus, we identified several potential causal sources of variation in barn swallow offspring sex ratio, including temporal, phenological and ecological factors. The observation of an association of offspring sex with temperature during laying is novel for birds and may be mediated by effects on maternal steroid hormones profile. The ecological and evolutionary implications of present findings are discussed in the light of adaptive sex allocation theory.     

Teleconnections and local weather orchestrate the reproduction of tit species in the Carpathian Basin

Variation in climatic conditions is an important driving force of ecological processes. Populations are under selection to respond to climatic changes with respect to phenology of the annual cycle (e.g. breeding, migration) and life‐history. As teleconnections can reflect climate on a global scale, the responses of terrestrial animals are often investigated in relation to the El Niño‐Southern Oscillation and North Atlantic Oscillation. However, investigation of other teleconnections and local climate is often neglected. In this study, we examined over a 33‐year period the relationships between four teleconnections (El Niño‐Southern Oscillation, North Atlantic Oscillation, Arctic Oscillation, East Atlantic Pattern), local weather parameters (temperature and precipitation) and reproduction in great tits Parus major and blue tits Cyanistes caeruleus in the Carpathian Basin, Hungary. Furthermore, we explored how annual variations in the timing of food availability were correlated with breeding performance. In both species, annual laying date was negatively associated with the Arctic Oscillation. The date of peak abundance of caterpillars was negatively associated with local temperatures in December–January, while laying date was negatively related to January–March temperature. We found that date of peak abundance of caterpillars and laying date of great tits advanced, while in blue tits clutch size decreased over the decades but laying date did not advance. The results suggest that weather conditions during the months that preceded the breeding season, as well as temporally more distant winter conditions, were connected to breeding date. Our results highlight that phenological synchronization to food availability was different between the two tit species, namely it was disrupted in blue tits only. Additionally, the results suggest that in order to find the climatic drivers of the phenological changes of organisms, we should analyze a broader range of global meteorological parameters.     

Reduced immunocompetence of nestlings in replacement clutches of the European magpie (Pica pica)

Laying date is one of the most important determinants of reproductive success and recruitment probability in birds. Late breeders usually fledge fewer chicks than individuals with earlier breeding dates, and fledglings produced late in the season have high mortality rates. Food availability and nestling mass have been evoked as the principal mechanistic links between laying date and offspring survival. Here we suggest that another factor may actually account for the difference in survival rate between early and late offspring: immunocompetence. We predicted that nestlings produced later in the season or in replacement clutches should have lower immune responses when challenged with an antigen, than early nestlings or nestlings produced in first clutches. This hypothesis was tested in a population of magpies (Pica pica), in which we experimentally induced breeding failure in a group of nests and compared the immune response of nestlings in replacement clutches with the immune response of first clutch nestlings. Cellular immune response, as measured by wing web swelling (a correlate of T-lymphocyte production after injection of phytohaemagglutinin-P), significantly decreased with hatching date and was significantly lower in nestlings of replacement clutches. Furthermore, coefficients of intraclutch variation in immune response were higher in nestlings of replacement clutches. This experiment demonstrates an inverse relationship between immune responsiveness and breeding date, and reduced recruitment probability of late nestlings may be a direct consequence of their inability to cope with parasites.     

Seasonal habitat-dependent change in nest box occupation by Tengmalm’s owl associated with a corresponding change in nest predation

We investigated the nest-site settlement of the forest-dwelling Tengmalm’s owl (Aegolius funereus) in the temperate zone of Central Europe (Czech Republic) over a 14-year period. We analysed the effect of changes in nest-site suitability (expressed as the proportion of deciduous forest) within the immediate surroundings of the nest boxes (a buffer radius of 25 m) in relation to nest box occupancy, the mean standardized laying date (further the laying date), occurrence of nest predation, and breeding density. In line with the predictions of the site dependence model, we found a positive relationship between the laying date and the proportion of deciduous trees in the nest box surrounding. Further, we found the interactive effect of the laying date and the proportion of deciduous forest surrounding a nest box on the occurrence of breeding attempts predated by pine martens. A higher proportion of deciduous forest was found in the surroundings of predated than non-predated breeding attempts, and a steeper positive relationship between the laying date and the proportion of deciduous forest was found in the surroundings of non-predated than predated breeding attempts. The change in the owls’ habitat utilization can be explained by the process of tree leafing which makes the deciduous forest a less suitable habitat with high risk of nest predation before tree foliage and a more suitable habitat with low risk of nest predation after tree foliage. We conclude that our results are in agreement with the site dependence model, and that the degree of foliage of deciduous forest in the nest surrounding of Tengmalm’s owl is a key factor determining site suitability due to predation risk from pine martens in Central Europe.     

Arrival date, age and breeding success in white stork Ciconia ciconia

Early arrival to breeding grounds is a life history trait in birds that can result in fitness benefits. We studied the relationship between arrival date and breeding success of individuals in a central Iberian population of white stork Ciconia ciconia , between 1999 and 2005, and the ways in which other potential factors, such as age or sex, affect this relationship. Our results showed that age was the factor most closely related to arrival date and breeding success. Older individuals returned earlier to the breeding grounds, achieved larger clutch sizes and produced more chicks than younger birds. After controlling statistically for age effect, breeding probability (laid eggs or not) and laying date were still significantly explained by arrival date. A higher probability of failure to reproduce (no eggs laid) was found in birds arriving later than in those arriving early. However, clutch size and nestling success (number of nestlings in the nest 40 days after hatching) were not correlated with arrival date. Food availability in the study area throughout the breeding cycle, due to a nearby rubbish dump, could be the factor mitigating differences in clutch size and nestling success related to individual arrival date.     

CONTRASTING PATTERNS OF PHENOTYPIC PLASTICITY IN REPRODUCTIVE TRAITS IN TWO GREAT TIT (PARUS MAJOR) POPULATIONS

Phenotypic plasticity is an important mechanism via which populations can respond to changing environmental conditions, but we know very little about how natural populations vary with respect to plasticity. Here we use random‐regression animal models to understand the multivariate phenotypic and genetic patterns of plasticity variation in two key life‐history traits, laying date and clutch size, using data from long‐term studies of great tits in The Netherlands (Hoge Veluwe [HV]) and UK (Wytham Woods [WW]). We show that, while population‐level responses of laying date and clutch size to temperature were similar in the two populations, between‐individual variation in plasticity differed markedly. Both populations showed significant variation in phenotypic plasticity (IxE) for laying date, but IxE was significantly higher in HV than in WW. There were no significant genotype‐by‐environment interactions (GxE) for laying date, yet differences in GxE were marginally nonsignificant between HV and WW. For clutch size, we only found significant IxE and GxE in WW but no significant difference between populations. From a multivariate perspective, plasticity in laying date was not correlated with plasticity in clutch size in either population. Our results suggest that generalizations about the form and cause of any response to changing environmental conditions across populations may be difficult.     

Selection on laying date is connected to breeding density in the pied flycatcher

Timing of reproduction and clutch size are important determinants of breeding success, especially in seasonal environments. Several recent bird population studies have shown changes in breeding time and in natural selection on it. These changes have often been linked with climate change, but few studies have investigated how the traits or natural selection are actually connected with climatic factors. Furthermore, the effect of population density on selection has been rarely considered, despite the potential importance of density in demographic processes. We studied variation in natural selection on laying date and on clutch size in relation to measures of spring phenology and population density in a long-term study of pied flycatchers in SW Finland. The phenological stage of the environment at mean egg-laying did not affect the direction of selection on either laying date or on clutch size. There was, however, stronger selection for earlier laying date when the breeding density of the population was high, suggesting that early breeding is not necessarily beneficial as such, but that its importance is emphasized when high population density increases competition. In addition, early breeding was favoured when the pre-breeding period was cool, which may indicate an increased advantage for the fittest individuals in harsher conditions. In the middle of the twentieth century, there was selection for large clutch size, which subsequently ceased, along with an overall decrease in recruit production. Our results indicate that attention should be paid to demographic factors such as breeding density when studying natural selection and temporal changes in it.     

吉林白城地区草原栗斑腹(巫鸟)窝卵数、营巢成功率和繁殖成功率的研究

对分布于吉林白城地区草原生境中栗斑腹巫鸟的窝卵数、营巢成功率和繁殖成功率的初步研究结果表明 ,繁殖期栗斑腹巫鸟种群的平均窝卵数为 5 .0 9± 0 .5 8枚 /巢 ;窝卵数与产卵期、出巢数与产卵期、窝卵数与卵大小之间呈负相关 ,产卵期与孵化率之间存在极显著的负相关关系 ,巢外径与窝卵数之间存在显著的正相关关系 ,巢的其余指标均与窝卵数呈正相关 ;平均孵化期为 12± 0 .4 9d ,孵化率为 36 .3% ,繁殖成功率为 11.11% ;7日龄以上的雏鸟群体大小为 2 .5 6± 1.5 3只 ,栗斑腹巫鸟的雏鸟存活率为 2 7.6 9% .     

Natural selection and inheritance of breeding time and clutch size in the collared flycatcher

Many characteristics of organisms in free-living populations appear to be under directional selection, possess additive genetic variance, and yet show no evolutionary response to selection. Avian breeding time and clutch size are often-cited examples of such characters. We report analyses of inheritance of, and selection on, these traits in a long-term study of a wild population of the collared flycatcher Ficedula albicollis. We used mixed model analysis with REML estimation ("animal models") to make full use of the information in complex multigenerational pedigrees. Heritability of laying date, but not clutch size, was lower than that estimated previously using parent-offspring regressions, although for both traits there was evidence of substantial additive genetic variance (h2 = 0.19 and 0.29, respectively). Laying date and clutch size were negatively genetically correlated (rA = -0.41 +/- 0.09), implying that selection on one of the traits would cause a correlated response in the other, but there was little evidence to suggest that evolution of either trait would be constrained by correlations with other phenotypic characters. Analysis of selection on these traits in females revealed consistent strong directional fecundity selection for earlier breeding at the level of the phenotype (beta = -0.28 +/- 0.03), but little evidence for stabilising selection on breeding time. We found no evidence that clutch size was independently under selection. Analysis of fecundity selection on breeding values for laying date, estimated from an animal model, indicated that selection acts directly on additive genetic variance underlying breeding time (beta = -0.20 +/- 0.04), but not on clutch size (beta = 0.03 +/- 0.05). In contrast, selection on laying date via adult female survival fluctuated in sign between years, and was opposite in sign for selection on phenotypes (negative) and breeding values (positive). Our data thus suggest that any evolutionary response to selection on laying date is partially constrained by underlying life-history trade-offs, and illustrate the difficulties in using purely phenotypic measures and incomplete fitness estimates to assess evolution of life-history trade-offs. We discuss some of the difficulties associated with understanding the evolution of laying date and clutch size in natural populations.     

Population size,breeding performance and territory quality of Bonelli's Eagle Hieraaetus fasciatus in eastern Spain

Capsule A five-year monitoring study is described of overall population stability and differential reproductive success in relation to habitat heterogeneity.

Aims To assess the effect of the altitude and orientation on laying date and breeding performance, and analyse the effect of territory quality as a likely factor that could be regulating the population.

Methods We monitored a population of 28–33 pairs, from 2002 to 2006, counting a total of 131 breeding attempts.

Results Territories located at lower altitude showed higher mean fecundity than those located at higher altitude. The mean laying date was February 18 ± 16 days. Laying date was positively correlated with nest altitude, the coastal pairs laying earlier than those in mountainous regions. Pairs located at lower altitudes showed higher mean fecundity than those located at higher altitudes. There was no preference in mean orientation either in breeding performance or in relationship to nest altitude. We did not find a difference in breeding performance between territories classed as being at high density and those classed as being at low density.

Conclusion The population has remained stable since the first national census was conducted 17 years ago. Our results could be explained in the light of the Habitat Heterogeneity Hypothesis. We suggest a differential reproductive success in relation to habitat heterogeneity.     

Sex ratio adjustments in common terns: influence of mate condition and maternal experience

Adaptive sex allocation has frequently been studied in sexually size dimorphic species, but far less is known about patterns of sex allocation in species without pronounced sexual size dimorphism. Parental optimal investment can be predicted under circumstances in which sons and daughters differ in costs and/or fitness returns. In common terns Sterna hirundo, previous studies suggest that sons are the more costly sex to produce and rear. We investigated whether hatching and fledging sex ratio and sex‐specific chick mortality correlated with the ecological environment (laying date, clutch size, hatching order and year quality) and parental traits (condition, arrival date, experience and breeding success), over seven consecutive years. Population‐wide sex ratios and sex‐specific mortality did not differ from parity, but clutch size, mass of the father, maternal breeding experience and to some extent year quality correlated with hatching sex ratio. The proportion of sons tended to increase in productive years and when the father was heavier, suggesting the possibility that females invest more in sons when the environmental and the partner conditions are good. The proportion of daughters increased with clutch size and maternal breeding experience, suggesting a decline in breeding performance or a resources balance solved by producing more of the cheaper sex. No clear patterns of sex‐specific mortality were found, neither global nor related to parental traits. Our results suggest lines for future studies on adaptive sex allocation in sexually nearly monomorphic species, where adjustment of sex ratio related to parental factors and differential allocation between the offspring may also occur.     

Social and demographic effects of anthropogenic mortality: a test of the compensatory mortality hypothesis in the red wolf

Whether anthropogenic mortality is additive or compensatory to natural mortality in animal populations has long been a question of theoretical and practical importance. Theoretically, under density-dependent conditions populations compensate for anthropogenic mortality through decreases in natural mortality and/or increases in productivity, but recent studies of large carnivores suggest that anthropogenic mortality can be fully additive to natural mortality and thereby constrain annual survival and population growth rate. Nevertheless, mechanisms underlying either compensatory or additive effects continue to be poorly understood. Using long-term data on a reintroduced population of the red wolf, we tested for evidence of additive vs. compensatory effects of anthropogenic mortality on annual survival and population growth rates, and the preservation and reproductive success of breeding pairs. We found that anthropogenic mortality had a strong additive effect on annual survival and population growth rate at low population density, though there was evidence for compensation in population growth at high density. When involving the death of a breeder, anthropogenic mortality was also additive to natural rates of breeding pair dissolution, resulting in a net decrease in the annual preservation of existing breeding pairs. However, though the disbanding of a pack following death of a breeder resulted in fewer recruits per litter relative to stable packs, there was no relationship between natural rates of pair dissolution and population growth rate at either high or low density. Thus we propose that short-term additive effects of anthropogenic mortality on population growth in the red wolf population at low density were primarily a result of direct mortality of adults rather than indirect socially-mediated effects resulting in reduced recruitment. Finally, we also demonstrate that per capita recruitment and the proportion of adults that became reproductive declined steeply with increasing population density, suggesting that there is potential for density-dependent compensation of anthropogenically-mediated population regulation.     

The impact of climate fluctuation on food availability and reproductive performance of the planktivorous red-billed gull Larus novaehollandiae scopulinus

1. Using 41 years of data, we examined annual variations in the reproductive performance of the red-billed gull at the Kaikoura Peninsula, New Zealand and related these to changes in climate, oceanography and the availability of the planktonic euphausiid Nyctiphanes australis, the bird's principal food. 2. In 1994 the population began to decline, and between 1983 and 2003 it was reduced by 51%. Adult non-breeders comprised an average of 32% of the population between 1983 and 1994, but following the decline they averaged only 18%. The age at recruitment changed markedly following the population decline: 27% of 2-year-old males bred for the first time prior to the decline, whereas the corresponding figure after the decline was 38%. The proportion of females commencing to breed as a 3-year-old was not significantly different before or after the decline. 3. An increase in the availability of euphausiids increased the likelihood of breeding and the recruitment of young individuals, caused earlier laying and resulted in an increase in the condition of adults, egg volume of gulls laying two egg clutches, clutch size and fledging success. 4. The relationship between food availability and the number of pairs that bred, laying date, clutch size and fledging success was significantly different prior to and after the population decline. The underlying cause appears to be a compensatory density-dependent mechanism that reduced interspecific competition for food. 5. The relative abundance of N. australis was correlated positively with the Southern Oscillation Index (SOI) and the frequency of occurrence of NE winds. The proportion of non-breeders and mean laying date were correlated negatively with the SOI, and mean egg volume of two egg clutches correlated positively with the SOI. 6. These results emphasize that availability of adult euphausiids is critical for red-billed gulls. We hypothesize that high primary productivity of inshore water near Kaikoura in winter, linked to a stable water column of coastal water and upwelling of additional dissolved inorganic nutrients, increases the availability of adult euphausiids to the red-billed gull as long as the coastal water is not replaced by offshore subtropical water intrusions of warmer, low-nutrient water.     

Seasonal change in Red-backed Shrike Lanius collurio territory quality - the role of nest predation

This paper examines the relationships between territory preference, reproductive performance and nest predation in a Red-backed Shrike Lanius collurio population breeding in rapidly changing farmland. Territory preference was measured by (a) the order of laying dates in territories within years and (b) territory occupation frequency over a five-year period. Seven environmental variables were tested but only the density of Sloe (Blackthorn) Prunus spinosa in territories was correlated positively with territory occupation frequency. Within years, Red-backed Shrikes started to breed earlier in territories with more Sloe, but reproductive performance was not related consistently to territory differences. A high proportion of Sloe-rich territories was vacant each year. Thus, the observed negative association between settlement order and amount of Sloe may reflect a seasonal change in habitat preference. Early breeders prefer Sloe-rich territories whereas late birds prefer Sloe-poor territories. Supporting evidence is that the proportion of occupied territories that was Sloe-rich was higher for early than for average or late first breeding attempts. Moreover, in contrast to early in the season, the probability of a replacement clutch following nest failure was not associated with the amount of Sloe, but increased with increasing densities of Juniper Juniperus communis. Late in the nesting season, pairs breeding in Sloe-rich territories suffered higher nest predation than those breeding in Sloe-poor territories. In early nesting attempts, however, the relationship was reversed. Nest predation is probably an important selective agent on habitat selection since it was the principal factor explaining variation in the numbers of fledglings produced among territories. Thus, the Red-backed Shrike may shift territorial preferences adaptively as the season progresses to match within-season changes in predation pressure in different types of territories.     

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

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

Intrinsic determinants of a population trend in timing of breeding in the wandering albatross

Numerous studies of wild animal species have documented that population level responses to environmental change are underpinned by individual level phenotypic plasticity. However, where the relationship between an individual trait and a climate variable occurs when both show a trend over time, phenotypic plasticity may be confounded by ageing. We investigated between and within individual change in laying date in the wandering albatross Diomedea exulans, a long‐lived species experiencing a dramatic decline in population size. Laying date has advanced over the last three decades. A mean‐centering analysis demonstrated that this pattern was driven by within‐individual changes as opposed to appearance or disappearance of phenotypes. Furthermore, a lack of between individual effect suggested the change resulted from ageing as opposed to phenotypic plasticity. Females varied significantly in rate of advance, such that those with low past reproductive rates exhibited a negative temporal trend in laying date, whereas birds with moderate to high past reproductive performance showed little change. The population trend was therefore driven by a subset with low past breeding success. An analysis of effects of timing of breeding on breeding success revealed stabilizing selection for relative laying date. Furthermore, current breeding success was positively related to past success rate, which suggests that there may be indirect selection against plasticity in this population. Our results show that population trends can arise from individual level change unrelated to prevailing environmental conditions, thus demonstrating the importance of longitudinal analyses in the interpretation of climate change effects.     

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.     

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.