Arctic waders and the capital‐income continuum: Further tests using isotopic contrasts of egg components

Birds migrating annually to high‐latitude breeding grounds may benefit from the transport of endogenous nutrient reserves that ultimately contribute to reproduction. Shorebirds represent a diverse group of Arctic breeders that typically arrive on the breeding grounds with body reserves enriched in ¹³C and ¹⁵N due to wintering and staging in marine or estuarine habitats. Such isotopic differences between endogenous macronutrient reserves and local foodwebs allow the use of stable isotopes to test for the source of nutrient allocations to eggs. We examined δ¹³C and δ¹⁵N values in lipid‐free yolk and albumen and δ¹³C values in yolk lipid of first clutches of ten species of sandpiper and plover breeding near Churchill, Manitoba, Canada in 2003. Most birds had egg isotope values indicating a C3 terrestrial biome, which fits primarily an income (exogenous) breeding strategy. Two exceptions were single sandpiper and plover with strong marine isotope values. Among species, strong positive relationships for each isotope between egg tissue components suggest that egg proteins and lipids tended to be derived from the same isotopic source. Correlations of egg δ¹³C values for lipids and proteins approached theoretical relationships expected for exogenous breeding strategies, based on captive studies. Significant positive correlations between clutch initiation date and δ¹³C values of egg lipids and albumen suggest some endogenous nutrient contributions to later laid eggs but the circumstances under which this may occur are unstudied. Where possible, we recommend that researchers use blood and fat biopsies from laying females as a means of anchoring endogenous and exogenous endpoints for modeling of each reproductive event. We encourage the isotopic analysis of egg albumen, yolk and yolk lipids among individuals and species and tests of correlations among these components as a means of inferring origins of nutrients to eggs.     

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.     

Intra‐clutch pattern of albumen δ13C and δ15N in yellow‐legged gulls Larus michahellis: female dietary shift or resource allocation strategy?

Energy or nutritional constraints associated to female dietary shifts during the clutch production period may play a role in generating intra‐clutch egg size variation in yellow‐legged gulls Larus michahellis. To explore this possibility, we determined albumen δ¹³C and δ¹⁵N values in three‐egg clutches (modal clutch size) from three different breeding episodes: Ebro Delta 2004 and 2006, and Columbretes Islands 2004. Rather than a shift in females’ diet, consistent intra‐clutch patterns of variation in egg size and albumen isotopic values (particularly in the case of albumen δ¹³C, which values held constant throughout the laying sequence) pointed to an intrinsic mechanism as the most feasible cause for the relatively smaller size of third/last‐laid eggs. However, diet “quality” for breeding females seemed to affect intra‐clutch egg size variation. In particular, a deficit of specific nutrients for egg formation associated to refuse scraps exploitation (as suggested by depleted albumen isotopic values) likely resulted in the more apparent intra‐clutch egg size profile for the Ebro Delta 2004. In the absence of dietary shifts, the observation of consistently higher δ¹⁵N values for third‐albumens suggested a greater contribution of endogenous resources to their synthesis, as conversion of stored reserves into egg proteins results in greater isotopic fractionation, thereby yielding enriched isotopic signatures (particularly for δ¹⁵N that shows greater isotopic fractionation with respect to that commonly assumed for δ¹³C). We point to reabsorbed material derived from the hormonally‐mediated regression of the female reproductive system (which is likely the intrinsic mechanisms resulting in the intra‐clutch pattern of egg size variation: the hormonal hypothesis) as the most feasible endogenous source of nutrients for the synthesis of last‐laid eggs, as optimize reproductive investment and maximize female fitness.     

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 reliance on distant resources for egg formation in high Arctic breeding barnacle geese Branta leucopsis

Breeding in the high Arctic is time constrained and animals should therefore start with their annual reproduction as early as possible. To allow for such early reproduction in migratory birds, females arrive at the breeding grounds either with body stores or they try to rapidly develop their eggs after arrival using local resources. Svalbard breeding barnacle geese Branta leucopsis have to fly non‐stop for about 1100 km from their last continental staging site to the archipelago making the transport of body stores costly. However, environmental conditions at the breeding grounds are highly unpredictable favouring residual body stores allowing for egg production after arrival on the breeding grounds. We estimated the reliance on southern continental resources, i.e. body stores for egg formation, in barnacle geese using stable isotope ratios in the geese's forage along the flyway and in their eggs. Females adopted mixed breeding strategies by using southern resources as well as local resources to varying extents for egg formation. Southern capital in lipid‐free yolk averaged 41% (range: 23–65%), early laid eggs containing more southern capital than eggs laid late in the season. Yolk lipids and albumen did not vary over time and averaged a southern capital proportion of 54% (range: 32–73%) and 47% (range: 25–88%), respectively. Our findings indicate that female geese vary the use of southern resources when synthesising their eggs and this allocation also varies among egg tissues. Their mixed and flexible use of distant and local resources potentially allows for adaptive adjustments to environmental conditions encountered at the archipelago just before breeding.     

Stable Isotopes (δD, δ13C, δ15N) Reveal Associations Among Geographic Location and Condition of Alaskan Northern Pintails

Abstract: Information on spring migration routes, geographic linkages among winter, spring, and breeding locations, and potential geographic effects on arrival body condition of northern pintails (Anas acuta) are currently unknown. Through a combination of stable-isotope measurements of tissues representing different periods of dietary integration and body composition analyses, we examined these linkages for pintails breeding in Alaska, USA. We collected 77 females at 4 locations upon spring arrival. We performed carbon (δ¹³C), nitrogen (δ¹⁵N), and hydrogen (δD) isotope measurements on flight feathers, breast feathers, and whole blood, and we conducted body composition analyses. Inference based on stable-isotope values in pintail tissues suggests that philopatry to Alaska was strong, as most of the collected females had stable-isotope values consistent with the boreal forest of Canada or western Alaska and most spring migrating females had whole-blood values indicating use of a food web in the boreal forest before collection. These patterns highlight the importance of the boreal forest for production and staging of pintails. Breast feather isotope values grown during prealternate molt were variable and covered the currently documented distribution of wintering pintails. Our results indicate associations among specific geographic areas, habitat use, and arrival condition of female pintails settling in Alaska. Females that wintered or staged in coastal habitat (as indicated by elevated δ¹³C values) arrived with less body fat compared to those that we inferred to have wintered or staged on inland freshwater habitat. Those females we inferred to use coastal areas appeared to rely more heavily on agricultural fields for nutrient acquisition (as indicated by elevated δ¹⁵N but low δ¹³C values). Our results provide the first link between low-condition females and inferred use of specific geographic areas before arrival. Conservation on wintering grounds should focus on restoration and protection of wetland complexes that provide adequate natural food resources in proximity to coastal systems that are heavily used by wintering pintails. Conservation efforts should also focus on the boreal forest, not only for pintail, but for other boreal-dependent species such as lesser scaup (Aythya affinis) (JOURNAL OF WILDLIFE MANAGEMENT 72(3):715–725; 2008)     

Examining carry‐over effects of winter habitat on breeding phenology and reproductive success in prairie warblers Setophaga discolor

Winter habitat quality can influence breeding phenology and reproductive success of migratory birds. Using stable isotope ratios of carbon (δ¹³C) from bird claws and red blood cells collected in Massachusetts, USA, we assessed if winter habitat occupancy carried over to affect prairie warbler Setophaga discolor breeding arrival dates, body condition upon arrival, pairing success, first‐egg dates and reproductive success. In two of three years (in 2011 and 2012, but not in 2013), after‐second‐year (ASY) males wintering in drier habitat, as indicated by enriched δ¹³C values, arrived later on the breeding grounds. Based on the North Atlantic Oscillation index, there was likely less rainfall in the Caribbean wintering grounds during the winters of 2011 and 2012 compared to the winter of 2013, suggesting increased winter rainfall in 2013 may have diminished the influence of winter habitat occupancy on arrival date. We did not find any effects of winter habitat on breeding season phenomena for second‐year (SY) males or females, but our sample sizes for these age/sex classes were relatively low. Although winter habitat quality influenced arrival dates of ASY males, there was no evidence that it affected reproductive performance, perhaps because of high rates of nest depredation in our system. Our study adds to a growing body of research that shows the influence of carry‐over effects can differ among species and within populations, and also can be modulated by other environmental conditions. This information enriches our understanding of the role of carry‐over effects in population limitation for migratory birds.     

King eiders use an income strategy for egg production: a case study for incorporating individual dietary variation into nutrient allocation research

The use of stored nutrients for reproduction represents an important component of life-history variation. Recent studies from several species have used stable isotopes to estimate the reliance on stored body reserves in reproduction. Such approaches rely on population-level dietary endpoints to characterize stored reserves (“capital”) and current diet (“income”). Individual variation in diet choice has so far not been incorporated in such approaches, but is crucial for assessing variation in nutrient allocation strategies. We investigated nutrient allocation to egg production in a large-bodied sea duck in northern Alaska, the king eider (Somateria spectabilis). We first used Bayesian isotopic mixing models to quantify at the population level the amount of endogenous carbon and nitrogen invested into egg proteins based on carbon and nitrogen isotope ratios. We then defined the isotopic signature of the current diet of every nesting female based on isotope ratios of eggshell membranes, because diets varied isotopically among individual king eiders on breeding grounds. We used these individual-based dietary isotope signals to characterize nutrient allocation for each female in the study population. At the population level, the Bayesian and the individual-based approaches yielded identical results, and showed that king eiders used an income strategy for the synthesis of egg proteins. The majority of the carbon and nitrogen in albumen (C: 86 ± 18%, N: 99 ± 1%) and the nitrogen in lipid-free yolk (90 ± 15%) were derived from food consumed on breeding grounds. Carbon in lipid-free yolk derived evenly from endogenous sources and current diet (exogenous C: 54 ± 24%), but source contribution was highly variable among individual females. These results suggest that even large-bodied birds traditionally viewed as capital breeders use exogenous nutrients for reproduction. We recommend that investigations of nutrient allocation should incorporate individual variation into mixing models to reveal intraspecific variation in reproductive strategies.     

Two eggs,two different constraints: a potential explanation for the puzzling intraclutch egg size dimorphism in Eudyptes penguins

Phenotypic plasticity and phenotypic stability are major components of the adaptive evolution of organisms to environmental variation. The invariant two-egg clutch size of Eudyptes penguins has recently been proposed to be a unique example of a maladaptive phenotypic stability, while their egg mass is a plastic trait. We tested whether this phenotypic plasticity during reproduction might result from constraints imposed by migration (migratory carry-over effect) and breeding (due to the depletion of female body reserves). For the first time, we examined whether these constraints differ between eggs within clutches and between egg components (yolk and albumen). The interval between colony return and clutch initiation positively influenced the yolk mass, the albumen mass, and the subsequent total egg mass of first-laid eggs. This time interval had only a slight negative influence on the yolk mass of second-laid eggs and no influence on their albumen and subsequent total masses. For both eggs, female body mass at laying positively influenced albumen and total egg masses. Female investment into the entire clutch was not related to the time in the colony before laying but increased with female body mass. These novel results suggest that the unique intraclutch egg size dimorphism exhibited in Eudyptes penguins, with first-laid eggs being consistently smaller than second-laid eggs, might be due to a combination of constraints: a migratory carry-over effect on the first-laid egg and a body reserve depletion effect on the second-laid egg. Both these constraints might explain why the timing of reproduction, especially egg formation, is narrow in migratory capital breeders.     

Rainfall at African wintering grounds predicts age‐specific probability of haemosporidian infection in a migratory passerine bird

In migratory species breeding in temperate zones and wintering in tropical areas, the prevalence of blood parasites may be affected by migratory strategies and winter habitat choice. We explored whether African winter habitat was linked to the probability of haemosporidian infection in the House Martin Delichon urbicum breeding in Spain, and tested for potential differences between age‐classes. As a proxy for winter habitat features, we analysed stable isotope (δ²H, δ¹³C and δ¹⁵N) values of winter‐grown feathers moulted in tropical Africa. Rainfall at the African winter grounds was related to the probability of being infected with haemosporidians and this effect differed among age‐classes. We found that haemosporidian prevalence was similar for young and experienced birds wintering in habitats of higher rainfall (²H‐depleted), whereas there were great differences in winter habitats of lower rainfall (²H‐enriched), with young having a much higher prevalence compared with experienced birds. Likewise, experienced birds wintering in habitats of higher rainfall had a higher probability of haemosporidian infection compared with experienced birds wintering in habitats of lower rainfall. By contrast, young birds wintering in habitats of lower rainfall had a higher probability of haemosporidian infection compared with young birds wintering in habitats of higher rainfall. These outcomes highlight the interaction of age with haemosporidian infection in the migratory ecology of the House Martin, which may drive carry‐over effects in this long‐distance aerial insectivore.     

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.     

Nutritional implications for sexual cannibalism in a sexually dimorphic orb web spider

Models predicting mechanisms driving sexual cannibalism in spiders with sexual size dimorphism (SSD) often assume that spiders use post‐copulatory food to channel nutrients into eggs and fecundity is altered through changes in clutch size or egg mass. I tested these assumptions using an orb web spider with extreme SSD, Argiope keyserlingi. I fed mated female spiders prey of either high protein‐low energy or low protein‐high energy composition. I measured egg energy density; a measure of the relative volumes of yolk and albumen. I predicted that if A. keyserlingi increase their egg energy density upon feeding on prey of a specific nutrient composition, they could enhance their fecundity by investing in more energy dense eggs. However, if the egg energy densities are dissimilar to their post‐copulatory prey they must be drawing energy from their somatic reserves to invest in eggs. In a further experiment I allowed female spiders to mate with and cannibalize males to determine if cannibalism induces similar effects on egg energy density. Male spider protein energy ratio was measured and found to resemble the high protein‐low energy prey. I found disagreement between the composition of post‐copulatory food and eggs in both experiments. Additionally, spiders fed high protein‐low energy prey lost weight indicating that they draw on their energy reserves to invest in eggs. I thus concluded that spiders that feed on high protein‐low energy prey or on males increase their egg energy density and, possibly, fecundity. However, the nutrient content of the prey or males cannot provide for investment in eggs. The energy invested in eggs is drawn from somatic reserves, probably induced by an as yet undescribed physiological trigger.     

Lesser Scaup Breeding Probability and Female Survival on the Yukon Flats,Alaska

ABSTRACT Information on the ecology of waterfowl breeding in the boreal forest is lacking, despite the boreal region's importance to continental waterfowl populations and to duck species that are currently declining, such as lesser scaup (Aythya affinis). We estimated breeding probability and breeding season survival of female lesser scaup on the Yukon Flats National Wildlife Refuge, Alaska, USA, in 2005 and 2006. We captured and marked 93 lesser scaup with radiotransmitters during prelaying and nesting periods. Although all marked lesser scaup females were paired throughout prelaying and incubation periods, we estimated breeding probability over both years as 0.12 (SE = 0.05, n = 67) using telemetry. Proportion of lesser scaup females undergoing rapid follicle growth at capture in 2006 was 0.46 (SE = 0.11, n = 37), based on concentration of yolk precursors in blood plasma. By combining methods based on telemetry, yolk precursors, and postovulatory follicles, we estimated maximum breeding probability as 0.68 (SE = 0.08, n = 37) in 2006. Notably, breeding probability was positively related to female body mass. Survival of female lesser scaup during the nesting and brood-rearing periods was 0.92 (SE = 0.05) in 2005 and 0.86 (SE = 0.08) in 2006. Our results suggest that breeding probability is lower than expected for lesser scaup. In addition, the implicit assumption of continental duck-monitoring programs that all paired females attempt to breed should be reevaluated. Recruitment estimates based on annual breeding-pair surveys may overestimate productivity of scaup pairs in the boreal forest.     

Range‐wide patterns of migratory connectivity in the western sandpiper Calidris mauri

Understanding the population dynamics of migratory animals and predicting the consequences of environmental change requires knowing how populations are spatially connected between different periods of the annual cycle. We used stable isotopes to examine patterns of migratory connectivity across the range of the western sandpiper Calidris mauri. First, we developed a winter isotope basemap from stable‐hydrogen (δD), ‐carbon (δ¹³C), and ‐nitrogen (δ¹⁵N) isotopes of feathers grown in wintering areas. δD and δ¹⁵N values from wintering individuals varied with the latitude and longitude of capture location, while δ¹³C varied with longitude only. We then tested the ability of the basemap to assign known‐origin individuals. Sixty percent of wintering individuals were correctly assigned to their region of origin out of seven possible regions. Finally, we estimated the winter origins of breeding and migrant individuals and compared the resulting empirical distribution against the distribution that would be expected based on patterns of winter relative abundance. For breeding birds, the distribution of winter origins differed from expected only among males in the Yukon‐Kuskokwim (Y‐K) Delta and Nome, Alaska. Males in the Y‐K Delta originated overwhelmingly from western Mexico, while in Nome, there were fewer males from western North America and more from the Baja Peninsula than expected. An unexpectedly high proportion of migrants captured at a stopover site in the interior United States originated from eastern and southern wintering areas, while none originated from western North America. In general, we document substantial mixing between the breeding and wintering populations of both sexes, which will buffer the global population of western sandpipers from the effects of local habitat loss on both breeding and wintering grounds.     

Winter habitat use does not influence spring arrival dates or the reproductive success of Yellow Warblers breeding in the arctic

Winter habitat use can influence the breeding success of migratory songbirds in temperate regions due to its impact on bird condition and breeding phenology. How such carry-over effects vary with latitude is unknown. To address this question, we examined how winter habitat use, inferred from δ¹³C and δ¹⁵N signatures in winter-grown feathers, influenced the breeding phenology and productivity of Yellow Warblers (Setophaga petechia) at the extreme north of their range in the Canadian arctic (68°N) and compared this population with midlatitude Yellow Warbler (51°N) and American Redstart (Setophaga ruticilla; 44°N) populations reported in previous studies. In the arctic, we examined male arrival dates, female clutch initiation dates and the relationship between these timing variables and the number and quality of offspring produced within the season. In contrast to warblers breeding at midlatitudes, we find no support for an impact of winter habitat use on breeding phenology or productivity. Male arrival dates and female clutch initiation dates in both young and older individuals were not correlated with isotopic signatures acquired on the wintering grounds. Males with enriched δ¹⁵N signatures paired more rapidly after arrival, indicating a possible relationship between winter habitat use and condition. This relationship did not enhance annual productivity for these individuals, however, as the negative relationship between breeding phenology and reproductive success in our arctic population was significantly weaker than among Yellow Warblers breeding further south. This reduction or absence of timing effects on productivity in the north effectively removes one pathway through which carry-over effects can act.     

Stable‐hydrogen isotope turnover in red blood cells of two migratory thrushes: application to studies of connectivity and carry‐over effects

ABSTRACT Understanding turnover rates of stable isotopes in metabolically active tissues is critical for making spatial connections for migratory birds because samples provide information about pre‐migratory location only until the tissue turns over to reflect local values. We calculated stable‐hydrogen isotope (δ²H) turnover rate in the red blood cells of two long‐distance migratory songbirds, Bicknell's Thrushes (Catharus bicknelli) and Swainson's Thrushes (Catharus ustulatus), using samples collected at a breeding site in New Brunswick, Canada. Blood from both species captured early in the breeding site was more positive in δ²H than blood sampled later in the summer, but did not match blood values for wintering Bicknell's Thrushes. An asymptotic exponential model was used to estimate turnover of red blood cell δ²H and yielded a half‐life estimate of 21 days and 14 days for Bicknell's and Swainson's thrushes, respectively. Red blood cells of both species approached the local breeding site value one month after the first individuals were detected at the site. For Bicknell's Thrushes, estimated δ²H in blood at arrival (?72‰) was closer to blood collected at wintering sites (mean ?61‰) than to expected breeding site δ²H (?120‰). Discrimination values calculated for red blood cells collected at the breeding site for both species were greater than expected based on studies using keratin. Turnover during migration currently limits the use of blood sampled early in the breeding season for connectivity/carry‐over effect studies. However, direct tracking technology such as geolocators can provide information about migration duration, timing, and stopovers that can be used to improve isotopic turnover equations for metabolically active tissues.     

Corticosterone and stable isotopes in feathers predict egg size in Atlantic Puffins Fratercula arctica

Examining factors that operate outside the breeding season may provide new insights into life‐history traits such as egg size, in which individual variation has not been fully explained. We measured corticosterone (CORT) levels and δ¹⁵N values (trophic level) in feathers grown several months before egg‐laying to test the prediction that a female's physiological state and feeding behaviour prior to the breeding season can influence egg mass in Atlantic Puffins Fratercula arctica. As predicted, egg mass increased with both CORT and δ¹⁵N values in feathers, suggesting that the ability of female Puffins to meet the nutritional costs of egg production is related to CORT promoting increased foraging effort during moult and to consumption of a higher trophic‐level diet.     

Complex trade-offs in the pigeon (Columba livia): egg antioxidant capacity and female serum oxidative status in relation to diet quality

Despite the great deal of ecological research interest in modulators of offspring quality and consequences of reproduction on female status, we still know little about the relationships among diet quality, antioxidant capacity of egg components (yolk and albumen) and oxidative status of female birds. In this study, I compared the egg quality (egg size, albumen and yolk antioxidant capacity) and serum oxidative status (oxidative damage, total serum antioxidant capacity, and serum thiols) of female pigeons (Columba livia) fed with foods of different quality (standard quality and decreased quality). I also analysed the patterns of covariation among egg and female traits. The study focussed on the first clutch laid by the female in the breeding season and on the short-term effects of a decrease in diet quality. The treatment did not affect the egg volume, the lipophilic and hydrophilic components of antioxidant capacity (lipOXY and hydrOXY, respectively) or the antioxidant capacity of the albumen (albumOXY). However, females fed a higher quality diet were fatter and had marginally higher values of serum hydroperoxides (oxidative damage) than females fed a lower quality diet. Moreover, females that showed an increase in yolk hydrOXY and serum hydroperoxides also showed a decrease in yolk lipOXY, albumOXY and serum antioxidant capacity. These results show that the female’s oxidative status can be correlated with the antioxidant content of her eggs, but the nature of these correlations is complex, depending on the molecular component measured. The results also suggest that in the pigeon the deposition of hydrophilic and lipophilic antioxidants in the egg may trade off against each other.     

The effect of maternal state on the steroid and macronutrient content of lesser black-backed gull eggs

It has been proposed that female birds can influence the phenotype of their offspring by provisioning eggs with variable amounts of nutrients and maternal hormones. Egg quality is strongly influenced by maternal body reserves and the amount of food available at the time of egg formation. This study investigated the effects of maternal state and food availability on the capacity of female lesser black-backed gulls Larus fuscus to provision their eggs with macronutrients and steroid hormones. Maternal state was reduced by increasing egg-production effort, whereas extra food was provided to reverse this effect. Compared with eggs of first clutches, eggs of experimentally induced replacement clutches exhibited a lower yolk/albumen ratio and contained more yolk testosterone. During one of the three years in which the study was performed, replacement eggs also contained more 17β-estradiol. Food provisioning during the relaying interval did not affect changes in yolk/albumen ratio or steroid concentrations, but fed females produced bigger eggs in their replacement clutch. This study demonstrates significant within-female consistency in egg size, macronutrient content, and yolk steroid concentration, and it shows that these egg characteristics are influenced by maternal state, food availability, and the timing of breeding.     

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.