Eggs brought in from afar: Svalbard‐breeding pink‐footed geese can fly their eggs across the Barents Sea

Many Arctic‐breeding waterbirds are thought to bring nutrients for egg production from southern latitudes to allow early breeding. It has proved problematic to quantify the extent of such capital breeding and identify whether nutrients for egg production are brought in from nearby or from afar. Before reaching their breeding grounds on Svalbard, pink‐footed geese Anser brachyrhynchus fly ～ 1100 km across the Barents Sea from Norway. Using abdominal profile indexing (API) we scored body stores in individually marked geese just prior to migration from the northernmost staging area in Norway to Svalbard, followed by their breeding success on their non‐breeding grounds in autumn. In productive breeding years leading to a high (> 13.8%) proportion of juveniles in the autumn population, there was a positive relationship between female API and number of young produced, suggesting that the geese are at least partial capital breeders. Moreover, focusing on the geographic origin of proteins used in egg synthesis and measuring nitrogen stable isotope ratios in pink‐footed geese's eggs and food sources in Norway and Svalbard, we identified that capital breeding in this species is ～ 50% on average but may potentially amount to as much as 100%, notably in females laying early. About 60% of this protein capital is carried in well‐developed follicles across the Barents Sea, the remainder likely being stored in muscle tissues. Conditions on the wintering grounds and migratory stopover sites can have profound effects on an individual's fitness but the here presented link between the use of migratory stopover sites and breeding performance is particularly noteworthy. Apparently, some individuals accept the putative costs of carrying body stores over large distances to the breeding grounds. The data also highlights considerable variation in the reliance on capital for breeding, suggesting substantial individual scope to adjust breeding strategy to changing environmental conditions.     

Keeping up with early springs: rapid range expansion in an avian herbivore incurs a mismatch between reproductive timing and food supply

Within three decades, the barnacle goose population wintering on the European mainland has dramatically increased in numbers and extended its breeding range. The expansion has occurred both within the Arctic as well as by the colonization of temperate areas. Studies of performance of individuals in expanding populations provide information on how well species can adapt to novel environments and global warming. We, therefore, studied the availability of high quality food as well as timing of reproduction, wing moult, fledgling production and postfledging survival of individually marked geese in three recently established populations: one Arctic (Barents Sea) and two temperate (Baltic, North Sea). In the Barents Sea population, timing of hatching was synchronized with the peak in food availability and there was strong stabilizing selection. Although birds in the Baltic and North Sea populations bred 6–7 weeks earlier than Arctic birds, timing of hatching was late in relation to the peak in food availability, and there was moderate to strong directional selection for early breeding. In the Baltic, absolute timing of egg laying advanced considerably over the 20‐year study period, but advanced little relative to spring phenology, and directional selection on lay date increased over time. Wing moult of adults started only 2–4 weeks earlier in the temperate populations than in the Arctic. Synchronization between fledging of young and end of wing moult decreased in the temperate populations. Arctic‐breeding geese may gradually accumulate body stores from the food they encounter during spring migration, which allows them to breed relatively early and their young to use the peak of the Arctic food resources. By contrast, temperate‐breeding birds are not able to acquire adequate body stores from local resources early enough, that is before the quality of food for their young starts to decrease. When global temperatures continue to rise, Arctic‐breeding barnacle geese might encounter similar problems.     

Relative contribution of lipid sources to eggs of lesser scaup

Studies of how birds mobilize nutrients to eggs have traditionally considered a continuum of possible allocation strategies ranging from income breeding (rely on food sources found on the breeding grounds) to capital breeding (rely on body reserves stored prior to the breeding season). For capital breeding, stored body reserves can be acquired either on or away from the breeding grounds, but it has been difficult to quantify the relative contribution of each, precluding identification of key habitats for acquiring nutrients for clutch formation. During 2006–2009, we explored the importance of spring‐staging habitats versus breeding‐area habitats for egg‐lipid formation in female lesser scaup Aythya affinis using stable carbon (δ¹³C) isotope analyses. Although δ¹³C values for abdominal lipid reserves brought to the breeding grounds overlapped those of local amphipods, we were able to quantify the importance of local plant carbohydrates (seeds of emergent wetland plants) to the production of eggs. We compared the importance of local wetland seeds (overall δ¹³C: ?29.1 ± 0.9‰ SD) to combined lipid stores and lipids from local amphipods (overall δ¹³C: ?23.8 ± 2.2‰). Local seeds and stored body lipids contributed equally to egg lipid formation across years but we found evidence of annual variation in their relative importance. Wetland seeds contributed 39% (SE = 10%) to egg lipid production, and the importance of this source varied by year (90% CI = 47–75% in 2006, 13–42% in 2007, 29–65% in 2008, and 7–30% in 2009). In contrast to earlier studies that suggest lesser scaup predominantly employ a capital breeding strategy, our results suggest that in some years females may attain half of their energy for clutch formation from foods on the breeding grounds.     

Forage plants of an Arctic‐nesting herbivore show larger warming response in breeding than wintering grounds,potentially disrupting migration phenology

During spring migration, herbivorous waterfowl breeding in the Arctic depend on peaks in the supply of nitrogen‐rich forage plants, following a “green wave” of grass growth along their flyway to fuel migration and reproduction. The effects of climate warming on forage plant growth are expected to be larger at the Arctic breeding grounds than in temperate wintering grounds, potentially disrupting this green wave and causing waterfowl to mistime their arrival on the breeding grounds. We studied the potential effect of climate warming on timing of food peaks along the migratory flyway of the Russian population of barnacle geese using a warming experiment with open‐top chambers. We measured the effect of 1.0–1.7°C experimental warming on forage plant biomass and nitrogen concentration at three sites along the migratory flyway (temperate wintering site, temperate spring stopover site, and Arctic breeding site) during 2 months for two consecutive years. We found that experimental warming increased biomass accumulation and sped up the decline in nitrogen concentration of forage plants at the Arctic breeding site but not at temperate wintering and stop‐over sites. Increasing spring temperatures in the Arctic will thus shorten the food peak of nitrogen‐rich forage at the breeding grounds. Our results further suggest an advance of the local food peak in the Arctic under 1–2°C climate warming, which will likely cause migrating geese to mistime their arrival at the breeding grounds, particularly considering the Arctic warms faster than the temperate regions. The combination of a shorter food peak and mistimed arrival is likely to decrease goose reproductive success under climate warming by reducing growth and survival of goslings after hatching.     

Travel schedules to the high arctic: barnacle geese trade-off the timing of migration with accumulation of fat deposits

On their way from the wintering area to the breeding grounds in Spitsbergen, barnacle geese Branta leucopsis stage on islands off the coast of Norway. The aim of this study was to describe when the geese migrate in relation to the body stores deposited and explore questions related to the concept of optimal migration schedules and on the possible mechanisms involved. We estimated fat stores by repeated assessments of the abdominal profile index of individually marked females throughout staging. Reproductive success was derived from observations of the same individuals later in the annual cycle. Females arriving late, or with low fat stores at arrival, achieved higher fat deposition rates, probably by spending more time foraging. But they were unable to match final fat scores of birds that arrived earlier or with larger fat stores. Reproductive success was correlated with the timing of migration and individuals departing at intermediate dates achieved highest success. The exact date of peak reproductive success depended on the size of fat stores accumulated, such that low-quality birds (depositing less fat) benefited most from an early departure to the breeding grounds. Observations in the breeding colonies showed that these birds did not initiate a nest earlier but they spent a longer time in Spitsbergen before settling. The length of stay in Norway was close to the prediction derived from an optimisation model relating spring events to eventual breeding success. Poorest performing birds stayed longer than expected, perhaps depositing more fat to avoid the risk of starvation. Two possible mechanisms of the timing of migration were contrasted and it seemed that the geese departed for migration as soon as they were unable to accumulate any more fat stores.     

Resource use for reproduction depends on spring arrival time and wintering area in an arctic breeding shorebird

Resources for egg production may come from body reserves stored before breeding (“capital breeders”) or from food acquired at the breeding site (“income breeders”). Arctic migrants were long thought to be capital breeders, because they often arrive at a time when local food availability is still limited. However, later evidence suggested that arctic breeding shorebirds are primarily income breeders, or that they use a mixed strategy depending on laying date. We explored the relationship between laying date and resource use for reproduction in the pectoral sandpiper Calidris melanotos breeding in the Alaskan arctic by contrasting carbon isotope (δ¹³C) values of the local diet and of maternal plasma, cellular blood, feather and claw with those of the eggs produced. Our results revealed that early breeding females utilize resources for egg production that were acquired recently at staging areas, whereas later breeding females mostly relied on nutrients derived from local food sources. These findings suggest that the resource allocation strategy used for reproduction differs among females, and varies depending on the timing of arrival and the start of reproduction. The arrival date at the breeding ground and laying date may critically depend on non‐breeding season events such as winter habitat choice, staging areas or migration routes. By comparing maternal feather δ¹³C, claw δ¹³C and feather δD, we examined whether non‐breeding season events influenced the use of resources for egg production through variation in capture date or clutch initiation date. Female pectoral sandpipers originating from moulting areas characterized by higher (more positive) δD signatures were caught earlier and started laying earlier, and they used stored resources for reproduction. Using regional maps of δD values for precipitation in the wintering sites in South America, we compared the spatial variation in the observed feather δD signatures. This analysis indicated that female pectoral sandpipers with higher δD signatures, presumably coming from more north‐easterly wintering sites in southern America, started laying earlier and used mostly stored resources for egg production, compared to females that wintered (or at least moulted) further south. Our results thus show that winter moulting habitat is linked to breeding resource allocation strategy in this high‐arctic breeding shorebird.     

The adaptive value of energy storage and capital breeding in seasonal environments

Timing of reproduction in a seasonal cycle is a life history trait with important fitness consequences. Capital breeders produce offspring from stored resources and, by decoupling feeding and reproduction, may bend the constraints caused by seasonality in food or predation. Income breeders, on the other hand, produce offspring from concurrent food intake, with the disadvantage of less flexibility, but with high efficiency and no inventory costs of carrying stores. Here, we assess relative profitability of capital and income breeding in herbivorous zooplankton inhabiting seasonal, high-latitude environments. We apply a state-dependent life history model where reproductive values are used to optimise energy allocation and diapause strategies over the year. Three environmental scenarios were modelled: an early, an intermediate, and a late feeding season. We found that capital breeding was most important in the early season. Capital breeding ranged from 7–9% of the eggs produced but, because of the high reproductive value of early eggs, capital breeding ranged from 9–30% when measured in terms of reproductive value. The main benefit of capital breeding was reproduction prior to the feeding season – when the reproductive value of an egg peaked. In addition, capital breeding was also used to increase egg production rates at times of income breeding. For individuals born late in the season the model predicted a two-year cycle instead of the typical annual life cycle. These individuals could then reap the benefits of early reproduction and capital breeding in their second year instead of income breeding late in the first year. We emphasize the importance of evaluating reproductive strategies such as capital and income breeding from a complete life cycle perspective. In particular, knowing the seasonality in offspring fitness is essential to appreciate evolutionary and population-level consequences of capital breeding.     

Analysis of the laying rhythm and reproductive traits of geese

The aim of the performed investigations was to analyse the laying rhythm and reproductive traits of Ko?uda white geese from the W11 reproduction strain and to determine the heritability of these traits as well as correlations between the laying rhythm traits and reproductive traits. The total number of geese participating in the experiment included 383 one-year old layers from the control flock (the first year of reproductive utilisation). The following traits characterizing the laying rhythm were assessed individually for each layer: the number of 2 and 3-egg clutches or more, length (in days) of 2- or more egg clutches as well as the length of intervals between the laid eggs during the entire laying period. The following reproductive traits were also assessed individually for each bird: age at sexual maturity, initial number of eggs (eggs laid during the period from January, 1st to April, 30th), number of eggs during the whole laying period, laying intensity (the total number of eggs x 100/length of the laying period in days) as well as the length of the reproductive period. It was found that Ko?uda white geese laid most of their eggs (on average 70.2%) singly and not in clutches. With regard to egg clutches, it was found that 2-egg clutches constituted 85.3% of eggs laid in clutches. Moderate or high variability of traits associated with the laying rhythm and reproduction were demonstrated. The observed moderate heritability of the laying rhythm traits indicate that they may be utilised in the selection programs for geese. On the other hand, the reported high, positive genetic correlation coefficients between the number of egg clutches and the initial and total egg number as well as laying intensity confirm the existence of interactions between these traits. This fact may be helpful in breeding programs for determining the optimal selection systems for geese.     

Travelling to breed

Traditionally, investigation of the dynamics of avian migration has been heavily biased towards the autumn return trip to the wintering quarters. Since the migratory prelude to breeding has direct fitness consequences, the European Science Foundation recently redressed the balance and sponsored a workshop on spring travels. We here survey the findings elucidating the complications arising during migration directed towards the breeding quarters. The evidence that early nesting confers advantage is overwhelming, hence demands of reproduction pose a constraint on both time and energy resources during spring. Both during migration and upon arrival there must therefore be strong selection in favour of saving time. Experimental results (e.g. using supplementary feeding) show that the date of laying is generally proximately constrained by the inability of the female to find sufficient nutrients. A key question thus concerns the implementation of the ‘capital’ strategy for breeding, the female accumulating nutrient stores along the way to bridge periods of shortage upon arrival on the breeding grounds. Eight studies on waterfowl (geese and eiders) and shorebirds (turnstones and knots) nesting in the arctic combine tracking of individuals (satellite telemetry) with direct observation (marked birds) and reconstruction of the origin of nutrients laid down in the eggs or in the form of body stores of the female parent (stable isotope signatures of tissues compared to potential food sources). The consensus emerges that in most cases a mixed strategy prevails, with nutrients garnered locally supplementing ‘imported’ body stores. The species concerned face a shortage of feeding time during incubation and suffer an energy deficit. Successful breeding thus depends on adequate fat depots that form part of the ‘capital’ the parents bring with them. Some headway has been made in predicting the extent of dependence on body stores for breeding in relation to body mass and length of migration from rates of fat deposition during stopover and fat consumption during flight for waterfowl. This work poses a challenge to refine field studies accordingly. The pressure to save time en route highlights the need to effectively exploit rich food resources. Several case studies underline the crucial significance of a very limited set of stopover sites, each with a narrow time window of optimal harvest. The influence of man at such sites often verges on the disastrous, and ongoing climate change may unhinge the finely tuned timing long-distance migrants depend on. There is a real need to extend this work to provide the scientific basis to underpin adequate conservation actions.     

Body condition of shorebirds upon arrival at their Siberian breeding grounds

Arctic breeding shorebirds carry substantial body stores on their long-distance migrations from their non-breeding grounds. Upon arrival at the breeding area the remains of these stores can be used for egg formation, insurance against poor feeding conditions or rebuilding organs. We quantified body condition (body mass, total body water, lean body mass and fat mass estimated using the deuterium dilution method) in seven shorebird species caught upon arrival in the Siberian Arctic. Arrival condition was compared with incubation condition in a subset of species. After correction for structural size, body mass was significantly lower at arrival than during incubation in most of the species (but 3–18% above lean mass). Fat index (fat mass/lean mass) varied between 5.1 and 13.2%. Fat stores were estimated to enable survival for 0.6 days for the smallest and 2.5 days for the largest species. We discuss possible functions of arrival stores: insurance, egg-formation or rebuilding organs. G. H. Visser: Deceased.     

Do purely capital layers exist among flying birds? Evidence of exogenous contribution to arctic-nesting common eider eggs

The strategy of relying extensively on stored resources for reproduction has been termed capital breeding and is in contrast to income breeding, where needs of reproduction are satisfied by exogenous (dietary) resources. Most species likely fall somewhere between these two extremes, and the position of an organism along this gradient can influence several key life-history traits. Common eiders (Somateria mollissima) are the only flying birds that are still typically considered pure capital breeders, suggesting that they depend exclusively on endogenous reserves to form their eggs and incubate. We investigated the annual and seasonal variation in contributions of endogenous and exogenous resources to egg formation in eiders breeding at the East Bay colony in the Canadian Arctic. We collected prey items along with females and their eggs during various stages of breeding and used two complementary analytical approaches: body reserve dynamics and stable isotope [δ(13)C, δ(15)N] mixing models. Indices of protein reserves remained stable from pre-laying to post-laying stages, while lipid reserves declined significantly during laying. Similarly, stable isotope analyses indicated that (1) exogenous nutrients derived from marine invertebrates strongly contributed to the formation of lipid-free egg constituents, and (2) yolk lipids were constituted mostly from endogenous lipids. We also found evidence of seasonal variation in the use of body reserves, with early breeders using proportionally more exogenous proteins to form each egg than late breeders. Based on these results, we reject the hypothesis that eiders are pure capital layers. In these flying birds, the fitness costs of a strict capital breeding strategy, such as temporary loss of flight capability and limitation of clutch and egg size, may outweigh benefits such as a reduction in egg predation rate.     

Potential for an Arctic‐breeding migratory bird to adjust spring migration phenology to Arctic amplification

Arctic amplification, the accelerated climate warming in the polar regions, is causing a more rapid advancement of the onset of spring in the Arctic than in temperate regions. Consequently, the arrival of many migratory birds in the Arctic is thought to become increasingly mismatched with the onset of local spring, consequently reducing individual fitness and potentially even population levels. We used a dynamic state variable model to study whether Arctic long‐distance migrants can advance their migratory schedules under climate warming scenarios which include Arctic amplification, and whether such an advancement is constrained by fuel accumulation or the ability to anticipate climatic changes. Our model predicts that barnacle geese Branta leucopsis suffer from considerably reduced reproductive success with increasing Arctic amplification through mistimed arrival, when they cannot anticipate a more rapid progress of Arctic spring from their wintering grounds. When geese are able to anticipate a more rapid progress of Arctic spring, they are predicted to advance their spring arrival under Arctic amplification up to 44 days without any reproductive costs in terms of optimal condition or timing of breeding. Negative effects of mistimed arrival on reproduction are predicted to be somewhat mitigated by increasing summer length under warming in the Arctic, as late arriving geese can still breed successfully. We conclude that adaptation to Arctic amplification may rather be constrained by the (un)predictability of changes in the Arctic spring than by the time available for fuel accumulation. Social migrants like geese tend to have a high behavioural plasticity regarding stopover site choice and migration schedule, giving them the potential to adapt to future climate changes on their flyway.     

Capital breeding and allocation to life-history demands are highly plastic in lizards

The use of stored resources to fuel reproduction, growth, and self-maintenance in the face of uncertain nutrient availability is a tactic common to many organisms. The degree to which organisms rely on stored resources in response to varied nutrients, however, is not well quantified. In this study, we used stable isotope methods to quantify the use of stored versus incoming nutrients to fuel growth and egg and fat body development in lizards under differing nutrient regimes. We found that the degree of capital breeding is a function of an individual's body condition. Furthermore, given sufficient income, lizards in poor condition can allocate simultaneously to storage, growth, and reproduction and "catch up" in body size and reproductive allocation to better-conditioned animals. Using natural variation in the δ(13)C of environmental nutrient pulses, we also found a high degree of variation in capital breeding in a lizard community. These findings demonstrate that capital breeding in lizards is not simply a one-way flow of endogenous stores to eggs but is a function of the condition state of individuals and seasonal nutrient availability. We use our findings to comment on capital breeding in lizards and the utility of the capital-income concept in general.     

The role of horsetails (Equisetaceae) in the nutrition of northern-breeding geese

Summary The nutrient content and balance of the horsetail Equisetum fluviatile was measured in samples taken from southern James Bay during early spring, summer and fall, and related to the nutrient requirements of geese at these times. Young emergent stems contained in excess of 20% protein of well balanced amino acid composition, apart from deficiencies of cystine and methionine. The ash fraction contained high levels of calcium, magnesium, potassium and phosphorus, which would satisfy the mineral needs of breeding and growing geese. The aerial parts of E. fluviatile did not contain much energy as neutral fats and soluble carbohydrate. The growth pattern of E. fluviatile, with young aerial stems arising throughout the summer, would provide a source of succulent, high-protein food. This could be used by geese migrating to the breeding grounds, during egg formation, to facilitate growth of goslings, and in the reacquisition of body protein lost during reproduction. The nutritive value of E. fluviatile was not high during the fall.     

Egg fertility among vasectomized and non‐vasectomized male resident Canada geese at the Wildlife Conservation Park/Bronx Zoo

In 1991, a vasectomy program was initiated at the Wildlife Conservation Park/Bronx Zoo to minimize the population growth of resident Canada geese (Branta canadensis). Sixty‐nine males were vasectomized over a 6‐year period. From 1994 to 1998, egg fertility was examined in females paired with vasectomized and non‐vasectomized geese. Of the 340 eggs laid by females paired with vasectomized males, 12% were fertile. In comparsion, of the 526 eggs laid by females paired with non‐vasectomized males, 90% were fertile. The vasectomy procedure was easier to perform on males captured during the breeding season than on those captured during molt owing to the enlarged size of the vas deferens. Where resident Canada goose populations are small and resources available, the implementation of a vasectomy program can be effective in reducing population growth. Zoo Biol 19:35–40, 2000. © 2000 Wiley‐Liss, Inc.     

Reproductive performance in arctic-nesting geese is influenced by environmental conditions during the wintering, breeding and migration seasons

Arctic regions are expected to experience pronounced changes in climate during the current century. Large numbers of waterfowl breed in these regions, and any climate induced changes are likely to have consequences for their demographics. Moreover, environmental changes experienced during migration and on the wintering grounds may also have impacts but remain poorly understood.
We investigate the role of climate variation during breeding, migration and wintering, while controlling for possible effects of mammalian predation and density dependence on the reproduction of Svalbard breeding barnacle geese Branta leucopsis using 40 years of observations.
Breeding success was significantly positively correlated with temperature on both the wintering grounds (Scotland) and breeding grounds (Svalbard), but negatively correlated with the number of days of strong cross-winds during the northward migration period. These factors remained significant when controlling for a strong negative effect of population size.
Goose reproduction on Svalbard was also linked to fluctuations in arctic fox Alopex lagopus populations occurring elsewhere in the arctic. This reveals the importance of mammalian predation, which may vary as a non-linear function of conditions within the wider arctic region.
Climate predictions were used to project barnacle goose reproduction and hence the population until 2050. These simulations suggest the population will grow at between 1% and 2.7% per year, in response to increasing temperatures. However, it is harder to predict how changes in other factors, such as reductions in sea ice, may impact on arctic breeding birds.     

Relationship between arrival timing and breeding success of intra‐tropical migratory Fork‐tailed Flycatchers (Tyrannus savana)

For migratory birds, early arrival at breeding areas has many benefits, such as acquisition of better territories and mates. This strategy has been found in numerous species breeding at north‐temperate latitudes, but has not been yet reported for intra‐tropical migratory species. We evaluated the relationship between arrival date, initiation of breeding, and breeding success of Fork‐tailed Flycatchers (Tyrannus savana) breeding in southeastern Brazil and overwintering in northern South America. We color‐banded adult flycatchers during three breeding seasons and searched for them during the following breeding seasons. We also monitored nests from construction until either failure or fledging of young. We found that: (1) male Fork‐tailed Flycatchers arrived at the breeding site earlier than females, (2) males that arrived earlier had greater breeding success, and (3) nests where eggs were laid earlier in the breeding season were more likely to be successful than those where eggs were laid later. Male Fork‐tailed Flycatchers appeared to benefit from early arrival at a tropical breeding site, potentially mediated by their ability to acquire a high‐quality territory and mate as early as possible, and by the ability of their mate to begin breeding as early as possible. Breeding success for female Fork‐tailed Flycatchers may be determined primarily by a combination of the arrival date of their mate and how quickly they can begin breeding. Our results suggest that protandry occurs in an intra‐tropical migratory bird and that early arrival of males and early initiation of reproduction by females results in greater reproductive success. A better understanding of the underlying mechanisms that control the timing of migration and reproduction of this and other intra‐tropical migratory species is important for evaluating the challenges they face in light of current and future rapid environmental changes.     

Energetics of reproduction in female and male greater snow geese

Arctic-nesting geese are classified as capital breeders (i.e., birds that rely largely on endogenous reserves to meet the high nutrient requirement of clutch formation) as opposed to income breeders (those that rely directly on ingested food). However, some evidence has suggested that energy reserves of greater snow geese (Chen caerulescens atlantica) are not sufficient to meet these costs. We tested whether greater snow geese were capital breeders and examined the consequences on their reproductive strategy. We collected 48 females and 47 males from arrival at the breeding colony on Bylot Island (NWT) to the postlaying stage, and determined fat and protein content in somatic and reproductive tissues. Rapid follicular growth was initiated only after arrival on the breeding ground. Somatic fat of females tended to increase during the prelaying and laying stages at the same time that most of the fat was deposited in developing follicles. Decrease in somatic fat in postlaying females was mostly related to its use for meeting metabolic requirements during early incubation. Hence, almost all fat invested in the clutch came from food intake, not endogenous reserves. Somatic protein was maintained during prelaying but decreased during laying, suggesting that some protein reserves were deposited in the eggs. There was no relationship between somatic fat and the number of developing follicles (incubating females excluded) but a relationship was found with somatic protein. Clutch size was not related to body size. In males, somatic fat, which was similar to females at arrival, was almost completely depleted by the postlaying stage. Male somatic protein remained stable. Male somatic reserves were not related to the somatic reserves, clutch size or nutrient investment in reproduction of their mate. We conclude that, in female greater snow geese, little fat reserves (if any) were used for egg formation though some protein reserves were used, whereas males relied heavily on their fat reserves during the same period. This is probably a consequence of the long and costly migration between the last staging area and the breeding grounds.     

Reproductive behavior and spawning success of female Amblyglyphidodon leucogaster (Pisces: Pomacentridae) from the Red Sea

The reproductive behavior of female whitebelly damselfish, Amblyglyphidodon leucogaster, was investigated in the Gulf of Aqaba, Red Sea over two breeding seasons. Females were promiscuous, mating with 7–10 different males throughout the season. Females lay eggs in distinct batches, defined as the total number of eggs laid in a day. Generally females deposit a batch of eggs with one male (87.2%) and are capable of laying a new batch every other day. Egg batch size averaged 4009 eggs and females laid from 2 to 22 egg batches per season. The variation in spawning success was not correlated to body size. Females preferred to deposit eggs in nests that already contained early stage eggs (0–2 days old). Within a nest, females chose to lay eggs contiguous to the youngest egg batch, regardless if the nest contained either a single batch or multiple batches of different ages. Female within-nest spawning patterns appear to be a consequence of between nest preferences for nests with young eggs. It is proposed that the strong within-nest preference is a consequence of mate selection where females may use new egg batches as a visual cue as part of a copying style. Such a style may reduce the risk of predation and increase feeding opportunities, because less time is expended in mate selection, which would provide additional resources for egg production and ultimately increase female spawning success over the breeding season. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.