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

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

Experimentally induced clutch size enlargements affect reproductive success in the Pied Flycatcher

We performed a food provisioning experiment in a population of Pied Flycatchers Ficedula hypoleuca breeding at high altitude in central Spain to test if food availability before and during laying determines clutch size. Food was provided to one of two pairs with the same date of initiation of nest-building (15 dyads of subsequently reproducing pairs were thus created). Food provisioning began on the day of initiation of nest-building and ended on the day after the last egg was laid. Although laying date was unaffected by the experiment, clutch size in the experimental treatment was significantly larger. This result could indicate that food availability at laying (1) proximately constrained clutch size or (2) that females evaluated future conditions for incubating eggs and feeding nestlings based on food availability at laying. Reproductive success (proportion of eggs that resulted in fledged young) was significantly reduced in the experimental treatment. This effect suggest that supplemented females were tricked by the experiment into laying more eggs than the number of eggs they were able to incubate with success and the number of nestlings they were able to feed, a source of error in clutch size adjustment which could be common in non-experimental situations.     

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

Introduction

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

Results

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

Conclusions

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

     

Timing of current reproduction directly affects future reproductive output in European coots

Life-history theory suggests that the variation in the seasonal timing of reproduction within populations may be explained on the basis of individual optimization. Optimal breeding times would vary between individuals as a result of trade-offs between fitness components. The existence of such trade-offs has seldom been tested empirically. We experimentally investigated the consequences of altered timing of current reproduction for future reproductive output in the European coot (Fulica atra). First clutches of different laying date were cross-fostered between nests, and parents thereby experienced a delay or an advance in the hatching date. The probability and success of a second brood, adult survival until and reproduction in the next season were then compared to the natural variation among control pairs. Among control pairs the probability of a second brood declined with the progress of season. Delayed pairs were less likely and advanced pairs were more likely to produce a second brood. These changes were quantitatively as predicted from the natural seasonal decline. The number of eggs in the second clutch was positively related to egg number in the first clutch and negatively related to laying date. Compared to the natural variation, delayed females had more and advanced females had fewer eggs in their second clutch. The size of the second brood declined with season, but there was no significant effect of delay or advance. Local adult survival was higher following a delay and reduced following an advance. The effect of the experiment on adult survival was independent of sex. Laying date and clutch size of females breeding in the next year were not affected by treatment. The study demonstrates the existence of a trade-off between increased probability of a second brood and decreased parental survival for early breeding. Timing-dependent effects of current reproduction on future reproductive output may thus play an important role in the evolution of the seasonal timing of reproduction.     

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.     

Differential maternal investment counteracts for late breeding in magpies Pica pica: an experimental study

Reproductive success in many avian populations declines throughout the breeding season. Two hypotheses have gained attention to explain such a decline: the "timing" hypothesis proposes that deteriorating food availability causes the decline in reproductive success (causal effect of breeding time), whereas the "quality" hypothesis proposes that individuals of lower phenotypic quality reproduce at the end of the breeding season, causing the correlation between breeding time and breeding success. We tested both of these hypotheses in a monogamous single breeder, the magpie Pica pica , by experimentally inducing some pairs to lay a replacement clutch, after removal of the first clutch. The first clutch was left in the nest of another magpie pair (matched by laying date and clutch size), and incubated and raised by these foster parents. In this way we obtained two clutches from the same magpie pair with full siblings raised in conditions of the first and second reproductive attempts. High quality pairs (with laying dates in the first half of the breeding season) reached similar breeding success in replacement clutches as compared to first clutches of the same female. In addition, experimental pairs reared significantly more offspring of similar quality in their replacement clutches as compared to late-season first clutches, thereby suggesting that late season first clutches were produced by pairs of lower phenotypic quality. Results indicate that high quality pairs trade-off clutch size for larger eggs in replacement clutches, which could help magpie pairs to partly compensate for poorer environmental conditions associated with a delayed breeding attempt.     

Happy to breed in the city? Urban food resources limit reproductive output in Western Jackdaws

Urban areas expand worldwide, transforming landscapes and creating new challenging habitats. Some bird species, mainly omnivorous feeding on human waste and cavity nesters, commonly breed in these habitats and are, therefore, regarded as urban‐adapted. Although urban areas may provide new nesting sites and abundant human waste, the low breeding success found in some of these species suggests that the poor protein content in human waste might limit breeding parameters. We investigated whether the breeding success of a cavity nester and omnivorous species commonly breeding in urban areas, the Western Jackdaw (Corvus monedula), depended on the availability of good‐quality non‐urban food. We approached the objective by combining a literature review and experiments in the field. With the literature review, we compared jackdaw populations in different habitats across Europe and found that clutch size and number of fledglings per pair decreased with distance to non‐urban foraging grounds, even after controlling for the effect of colony size, latitude, and climate. In two experiments, we tested whether the breeding success of urban pairs could be increased by supplementing high‐quality food, first only during egg formation and second also until chick fledging. Food supplementation during egg formation led to larger eggs and higher hatching success than in urban control nests, but this did not result in higher chick survival. However, when food supplementation was prolonged until fledging in the second experiment, we observed a significant increase of nestling survival. These findings highlight that research and management actions should not only focus on species displaced by urbanization, but also on “urban‐adapted” species, as they might be suffering from a mismatch between availability of nesting sites in buildings and adequate non‐urban food resources. In these cases, nest sites should be provided in or close to adequate food resources.     

Effect of food abundance on laying date and clutch size in the White Stork Ciconia ciconia

CapsuleFood independently affects both laying date and clutch size, suggesting that seasonal decline in clutch size is related to a decrease in food availability.

Aim To test the effect of food abundance on laying date and clutch size of the White Stork and identify the cause of seasonal decline in the number of eggs laid.

Methods During 1991 and 1996 we recorded clutch size and laying date of pairs breeding next to rubbish dumps (food abundant and constant throughout the breeding season) and birds breeding far from rubbish dumps (using natural food sources).

Results In 1991 there was no difference in mean laying date between pairs nesting at rubbish dumps and control pairs. Clutch size was significantly larger at rubbish dump nests. In contrast, mean laying date was earlier in control pairs in 1996 and there was no significant differences in clutch sizes, even when controlling for laying date effect.

Conclusion The results support the hypothesis that food availability independently affects both laying date and clutch size. The seasonal decline in clutch size close to rubbish dumps was negligible (1991) or much smaller than in the control zone (1996) suggesting that a progressive deterioration of natural food sources is the most probable reason for a decline in clutch size as the season advances.     

Seasonal variation in reproductive measures of tropical Roseate Terns Sterna dougallih previously undescribed breeding patterns in a seabird

Seasonal variation in egg-laying, egg size, hatching success, hatchling mass, fledging success and chick growth of Roseate Terms Sterna dougallii breeding on Aride Island (Seychelles), Indian Ocean, were studied in 1997 and 1998. I investigated to what extent two patterns, common in a range of species, were followed by tropical Roseate Terns: (a) seasonal decrease in clutch size, egg size and breeding success and (b) an increase in breeding success with increasing egg weight. In 1997 (a poor year), the earliest nesting birds laid significantly smaller eggs, and chicks were lighter at hatching than those of peak nesting birds. The mean clutch size, of 1.04 eggs, showed no seasonal variation and no 'b'-eggs hatched. In 1998 (a good year) the earliest nesting birds laid eggs of similar size and their chicks were of similar weight to those of peak nesting birds. Mean clutch size, of 1.25 eggs, increased significantly through the season and about 60% of the 'b'-eggs hatched. In 1997, hatching success was 57% whereas in 1998 it was 80%. In both years, breeding success declined significantly through the season. The fact that the earliest breeding birds laid smaller eggs in a poor year and smaller clutches in a good year is in marked contrast to a range of other species, and to temperate-nesting Roseate Terns. Egg volume explained about half of the variance in hatchling mass in both years, but only 15% of the variation in linear growth rate. Hatching date was the only variable with a significant effect on fledging success. Roseate Terns on Aride seemed to sacrifice egg size and clutch size for earliness of laying. Presumably it is a strategy of older birds to lay as early as possible and may be regarded as a response of tropical Roseate Terns to breeding under relatively poor, and seasonally declining, food conditions.     

Nutritional quality of prebreeding diet influences breeding performance of the Florida scrub-jay

Food supplementation studies of breeding birds have traditionally concentrated on energetic constraints on breeding performance. It is only recently that the nutritional quality of the prebreeding diet has also been considered influential. We examined the importance of specific nutrients in the prebreeding diet of the Florida scrub-jay ( Aphelocoma coerulescens). Birds were provided with one of two supplements (rich in protein and fat or rich in fat only) prior to breeding in 2000 and 2001 and their breeding performance, in relation to unsupplemented (control) birds, was examined. Birds receiving both supplements significantly advanced laying in both years, and increased clutch size in 2000 but not in 2001. Laying date explained variation in clutch size in birds on dietary supplements. Egg mass and volume declined with laying order, irrespective of dietary treatment, but birds on the high fat, high protein diet laid heavier third eggs than controls and this was independent of laying date. Laboratory analysis of 14 abandoned and unhatched eggs revealed that as egg mass increased so did the absolute amount of protein and water while fat content remained relatively fixed. Using these relationships between the masses of egg components and fresh egg mass, we calculated that heavier third eggs laid by birds on high fat and high protein, compared with those laid by controls, contained more water that may be fundamental to chick growth and survival. This is the first demonstration for an avian species that nutritional quality of prebreeding diet can simultaneously influence laying date, clutch size, and egg size and composition.     

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.     

Variation and trade-off in reproductive output of the Andean frog<Emphasis Type="Italic"> Hyla labialis</Emphasis>

I analysed variation in number and size of eggs in association with variation in body size of Hyla labialis females from three altitudinally widely separated populations in the Colombian Andes. Females at 3,500 m were significantly larger and heavier than those at 2,700 m, which in turn were larger and heavier than those at 2,000 m. After accounting for interpopulational differences in body size, clutch size (i.e. number of eggs) and egg size differed significantly between populations, but clutch mass did not. At 3,500 m, large relative clutch mass tended to be associated with larger eggs, whereas at lower altitudes it was associated with relatively larger clutch size. Within each population, females showed a significant trade-off between clutch size and egg size, even though relative clutch mass varied substantially and a clutch rarely used up the carrying capacity of their body cavity. At 3,500 m, comparisons between the early and late breeding season within a year suggested that climate fluctuations may play a role in reproductive-output variation within a population. Laboratory-kept females in a high somatic condition after spawning laid larger clutches in the next breeding event, but the trade-off persisted despite ad libitum food supply. Comparisons between field and laboratory suggest that limits to reproductive output were due to physiological, but not morphological restrictions. Therefore, the trade-off is better explained by energy limitation for egg growth depending on clutch size, than by the body cavity limitation hypothesis.     

Breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in high-arctic northeastern Greenland, 1993

The breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in the high Arctic was studied in relation to the occurrence of the northeast water polynya in northeasternmost Greenland (80̀N). Mean laying dates were 31 May in the Fulmar and 18 June in the Kittiwake; the total nesting season for the Fulmar just matched the time window of the polynya opening period. Fulmar colony attendance fluctuated within a period of 11.6 days because of variation in nonbreeding prospectors but showed no clear diurnal variation. Fulmar incubation shifts, on average, lasted 6.1 days (range 1–13 days), which is significantly longer than elsewhere, and the average chick-guard period of 10.9 days (range 1–17 days) was significantly shorter than in other studies. Egg neglect occurred in 18% of Fulmar nests or 0.7% of nests per day. Overall breeding success (chicks fledged per egg laid) was 0.56 in the Fulmar and 0.67 in the Kittiwake; the latter produced 1.4 young per active nest or 1.2 per completed nest. Mean Kittiwake clutch size was 2.03; larger clutches were laid early. Nest site characteristics (presumably reflecting nest predation risk) and breeding behaviour affected breeding success. in the Fulmar, hatching success was negatively correlated with laying date and the proportion of egg neglect, while overall breeding success was correlated negatively with distance to nearest neighbouring site and positively with the length of the chick-guard period. Kittiwake breeding success was negatively correlated with laying date. Using seabirds as indicators of marine food supply, breeding success in both species suggested moderate to good food supply in the northeast water polynya in 1993, although at least in the Fulmar the high reproductive output appeared partly maintained by behavioural buffering; long incubation shifts, egg neglect and short chick-guard periods were symptoms of foraging constraints.     

Life-history variation within a three-spined stickleback population in the Camargue

Among individuals of female three-spined sticklebacks Gasterosteus aculeatus from a population in the Camargue, southern France, studied in 12 successive years, adult L _T ranged from 31–64 mm, clutch size ranged from 33–660 eggs, and mean egg diameter per clutch ranged from 1.15–1.67 mm. Because the population was strictly annual, inter-annual variation corresponded to variation among generations having experienced different environmental conditions. Body mass varied significantly among years, suggesting an effect of varying environmental conditions. Gonad mass and clutch size increased with body mass, but mean egg diameter was not correlated to body mass. Body mass-adjusted gonad mass, interpreted as reproductive effort per clutch, did not vary significantly among years, suggesting that this trait was not influenced by environmental conditions. Body mass-adjusted clutch size and egg size varied significantly among years. Inter-annual variation in body length at breeding, clutch size and egg size was of the same order of magnitude as inter-population variation reported by other authors for this species. During the breeding season, reproductive effort and clutch size tended to increase. Egg size tended to decrease during the breeding season but this seasonal pattern varied among years. Observed life-history variation is discussed both in terms of its evolutionary significance and methodological implications in the study of life-history variation among populations.     

Pre-incubation feeding activities and energy budgets of Snow Geese: can food on the breeding grounds influence fecundity?

The potential contribution of early spring feeding conditions in the Arctic to clutch size variation was examined in a population of Lesser Snow Geese Anser caerulescens caerulescens. Behavioural observations were combined with energetic analyses of food material to derive an estimate of the energy budgets of pre-laying and laying females. Food intake of females between arrival on the breeding grounds and incubation was considerable; estimated energy gains in this period were in the same magnitude as the cost of one or several eggs. The pre-laying period spent on the breeding grounds can thus be energetically beneficial rather than costly. Accumulation of resources for reproduction in Snow Geese is a continual process including the breeding grounds, and nutrient limitation after arrival in the Arctic cannot sufficiently explain the environmental component of clutch size variation. The timing of migration and follicle development is such that clutch size decisions are sometimes made during the late stages of migration and some-times after arrival. In the latter case food conditions on the breeding grounds may greatly influence clutch size; in the former case they may still influence readjustments of clutch size after the initial decision. The universal negative correlation between clutch size and laying date in Snow Geese can be explained by negative fitness consequences of late hatching, which outweigh the benefits of delayed laying and further nutrient accumulation. Food shortage on the breeding grounds may sometimes be a secondary factor contributing to seasonal clutch size decline.     

Do marine birds use environmental cues to optimize egg production? An experimental test based on relaying propensity

According to the environmental cues hypothesis, female birds use information available to them in the early‐season environment to fine‐tune egg production annually. However, support for the hypothesis derives largely from correlational studies. In each year from 2002 to 2006, which spanned a period of extreme variation in environmental conditions, we removed eggs from early‐laying rhinoceros auklets Cerorhinca monocerata, burrow‐nesting seabirds that lay a single‐egg clutch. We then measured their relaying rates, relaying intervals, and breeding success. We also monitored the timing and success of breeding in control pairs, and control chick diets. If the experimental females base their relaying decision on early‐season cues, then we predict that few will relay in years in which early‐laying control birds breed unsuccessfully, and in which a preferred prey species, Pacific sandlance Ammodytes hexapterus, is in short supply in nestling diets. Results matched neither prediction. In each year, almost all (88–90%) of the experimental females relaid, despite that the control pairs’ breeding success (32–87% fledged chicks), and their chicks’ diets (twofold variation in proportion of sandlance), varied markedly. We conclude that female rhinoceros auklets did not modify their relaying decision in response to variation in environmental conditions, although relaying intervals and their own breeding success (0–78%) covaried negatively. Our results may have important implications related to using seabirds as monitors of the marine environment.     

Clutch size variation in the Nazca booby: a test of the egg quality hypothesis

In obligately siblicidal bird species, aggressive behavior bya dominant chick results in a fixed brood size of one, yetthese species usually show clutch size variation between individuals.Simmons proposed that variation in clutch size in obligatelysiblicidal species is related to a trade-off between egg qualityand egg quantity: some individuals produce a single highly hatchable egg, while others produce two small, lower qualityeggs. We tested the egg quality hypothesis as an explanationfor observed clutch size variation in the Nazca booby (Sulagranti), an obligately siblicidal seabird. We tested the assumptionthat egg volume is positively correlated with hatchabilityand the prediction that eggs from one-egg clutches are largerthan eggs from two-egg clutches. We did not find a positive relationship between egg volume and hatchability in this species.Eggs from two-egg clutches were either equivalent in volumeor larger than eggs from one-egg clutches. Thus, the egg qualityhypothesis was rejected as an explanation for clutch size variationin the Nazca booby. Instead, two-egg clutches appear to befavored because of the insurance value of the second-laid egg,while one-egg clutches result from food limitation.     

Supplementary feeding increases Common Buzzard Buteo buteo productivity but only in poor‐quality habitat

Temporal heterogeneity in the effects of food supply during the breeding season on the productivity of the Common Buzzard Buteo buteo was investigated in a supplementary feeding experiment. Pairs were fed artificially (1) before egg‐laying, (2) after chicks hatched and (3) continuously throughout the season, and compared with (4) unfed controls. Pairs fed before egg‐laying had marginally larger clutches than those not fed, but lay date, egg volume and weight, brood size and hatching success were unaffected. Territorial quality had far greater effects, with pairs nesting in low‐quality habitats (bog, scrub and semi‐natural grassland) laying later and having lower hatching success, smaller broods and fewer fledglings than those in more productive agricultural landscapes. Supplementary feeding after egg hatching neutralized the negative effect of poor habitat, resulting in fed birds having significantly more fledglings. This study emphasizes the importance of food availability when provisioning chicks in suboptimal habitats and has implications for the success of diversionary feeding in reducing game losses to Buzzards.     

Components of productivity in black-legged kittiwakes Rissa tridactyla: response to supplemental feeding

In contrast to the high productivity of black-legged kittiwakes in Britain, kittiwakes at many colonies in Alaska have failed chronically to reproduce since the mid 1970s. To determine if food is limiting productivity and, if so, at what stages of nesting food shortages are most severe, in 1996 and 1997 we supplementally fed kittiwakes nesting on an abandoned building. The effects of feeding were stronger in 1997 than in 1996, possibly because naturally occurring prey were of poorer quality in 1997. Consumption of supplemental herring declined as egg laying approached then increased slowly during incubation and more rapidly after hatching. All of the six components of productivity we studied were improved by supplemental feeding to some degree. Supplemental food did not significantly alter laying success in either year, although fed pairs bred at slightly higher rates than unfed pairs in 1997, the poorer food year. In 1996 and 1997, extra food noticeably increased clutch size and hatching success, but significantly so only in 1997. Fledging success and productivity were substantially augmented by feeding in both years. Fed pairs fledged twice as many chicks per nest as did unfed pairs in 1996 and three times as many in 1997. Fed and unfed pairs lost most of their potential productivity through the inability to hatch eggs, and secondarily because of their poor success at raising chicks. The benefits of supplemental feeding did not carry over from one stage of breeding to another. Pairs cut off from supplemental food after laying or hatching performed similarly to pairs that had not been previously fed. This study provides benchmark values of breeding performance attainable by kittiwakes in Alaska under optimal conditions. These values are comparable to highly productive colonies in Britain and suggest that differences in life-history characteristics between Pacific and Atlantic breeding populations are primarily controlled by food supply.     

Reproductive parameters in relation to food supply in the whiskered tern ( Chlidonias hybrida )

It is generally accepted that breeding terns are sensitive to food supply and that their reproductive effort could be substantially affected by the availability and access to resources. In this study we examined reproductive parameters in the whiskered tern Chlidonias hybrida in relation to food supply during the courtship feeding period (food brought by males to females) over a 2-year period (2004–2005). We also studied whether the condition of the nestlings [body condition index (BCI)] was related to a proxy of the reproductive investment of the adults (the clutch size) during a season of food shortage. Behavioural observations showed a decrease in the intensity of male courtship feeding between years (2004 > 2005), and a strong shift in the relative abundance of the two prey groups (invertebrates/fish; invertebrate prey dropped from 88.0 to 49.3%) brought by males. This change in food delivery rates did not result in a delay in laying, but there was a significant difference in mean clutch size between years (2.71 ± 0.49 eggs in 2004 and 2.05 ± 0.78 eggs in 2005) without any within-year variation in relation to the laying dates. The egg size (volume and length) was related to the year (2004 < 2005), suggesting a trade-off in the quantity and the quality of eggs between the two seasons. We also found no evidence that the investment in a large clutch affected nestling BCI in the course of the food shortage season. Since many pairs (about 60%) interrupted breeding during the incubation stage, we assumed that parents that succeeded in rearing nestlings in these conditions were probably ‘high-quality’ individuals. Our results therefore showed that whiskered terns are sensitive to the varying food conditions they experienced throughout the courtship period. The diversity of prey types could be a key factor in the reproductive investment of this tern species.