Intraseasonal Variation in the Development of Sexual Size Dimorphism in a Precocial Bird: Evidence from the Lesser Snow Goose

1. Variation in offspring sex ratio at the end of parental care may reflect passive sexual differences in offspring survival, or a facultative manipulation by parents to balance trade-offs between sex-specific offspring growth and survival and parental fitness. In contrast, among species with limited size dimorphism, there may be little reason to expect significant sexual differences in offspring survival, and the presence of offspring sex-ratio variation in such species may indicate sex-specific differences in parental investment at some stage. However, sex-ratio variation in monomorphic species with praecocial young may be more difficult to explain, since the decreased parental investment, which is largely shared between offspring of either sex, may reduce the opportunity for sex-ratio manipulation. 2. We examined this question by analysing annual variation in fledging sex ratio of an avian herbivore with strong female natal philopatry and praecocial young, the lesser snow goose Anser caerulescens caerulescens, using data from a 23-year observational study during a period of marked population growth. Although snow geese show limited sexual size dimorphism, with males being 2-6% larger at all ages posthatching, growth and survival of snow goose goslings has been shown previously to be extremely sensitive to variation in food supply. 3. The proportion of males at fledging has decreased from 51% in the early years, when food at the colony was abundant, to c. 48% in recent years, when food supply has been reduced. Concurrent with this change, the mass and condition of goslings have declined significantly over the years, with males showing a proportionately greater decline than females. Hatching sex-ratio data showed no systematic significant differences overall, and thus the most likely explanation for the change in sex ratio is increased mortality of male goslings between hatching and fledging. 4. The change in fledging sex ratio occurred only among broods from younger adults (age < 5 years). These age-specific differences were evident both among and within individual females. Younger adults may be less able to accommodate long-term changes in foraging conditions successfully, resulting in proportionately greater gosling mortality overall, and increasingly biased mortality of male goslings, which may be more susceptible to food restriction. 5. Despite potential differences in indirect costs of rearing male or female offspring, it is suggested that these results are more consistent with increased mortality of male goslings under increasing food restriction, rather than active parental manipulation.

VARIATION IN BLUE TIT BREEDING PERFORMANCE ACROSS GRADIENTS IN HABITAT RICHNESS

The hypothesis that migratory bird populations are limited during the non-breeding season is increasingly supported by empirical studies that also suggest consequences that carry-over into subsequent seasons. Although variation in food supply is the best supported explanation for non-breeding season limitation, the ecological mechanisms and physiological consequences are not well understood. We both supplemented and reduced Ovenbird (Seiurus aurocapilla) food availability on replicate plots in Jamaica in each of 3 years to determine the direct role of food in limiting the physical condition of Ovenbirds. Annual variation in rainfall and food supply created a natural experiment in parallel with manipulations. Sex and age-classes of Ovenbirds did not respond differently in terms of body condition to either food manipulation or natural variation in environmental conditions, suggesting that this population is not structured by strong dominance relationships. Ovenbird body mass, fat, and pectoralis muscle shape were positively and predictably related to manipulated food availability. Feather regrowth rate also responded positively to food supplementation and negatively to food reduction in the drier of 2 years. Prior to manipulation, annual variation in body mass corresponded to annual variation in food supply and rainfall, providing additional, correlational evidence of food limitation. Since multiple intercorrelated body condition indices of Ovenbirds responded directly to food supply, and since food supply influenced body condition independently of other habitat features, we argue that food is a primary driver of non-breeding season population limitation. Moreover, since these effects were observed during the late non-breeding period, when individuals are preparing to migrate, we infer that food availability likely initiates carry-over effects.

Bird species adapted to variable environments tend to have slow lean tissue growth rates and high fat deposition rates, allowing survival during food shortages. This emphasis on fat deposition may be a fixed physiological trait. Alternatively, tissue allocation may be adjusted facultatively according to the proximate food supply. We consider two models of facultative adjustment that could account for the emphasis on fat deposition: (1) the fat‐priority model, in which no lean growth occurs when food is scarce, and (2) the lean‐priority model, in which a minimal level of lean growth always occurs but nutrients are otherwise allocated to fat deposition. We tested these two models using Welcome Swallows Hirundo neoxena, a species we show to have a variable food supply that is influenced by weather. We reduced food supply to chicks experimentally, by enlarging broods or excluding parents from chicks, and tested for reduction in wing growth (an indicator of lean growth) and mass growth (an indicator of fat deposition). Mass growth was retarded by both manipulations, but not wing growth, corroborating the lean‐priority model. This growth strategy may function not to cope with violent variation in food supply, but to maintain development and symmetry of wings and feathers in the face of moderate variation in food supply. Our results contrast with those of a similar experiment on the Black Noddy Anous minutus, a species with more severe variation in food supply.

Earlier reports suggested that seasonal variation in food-caching behavior (caching intensity and cache retrieval accuracy) might correlate with morphological changes in the hippocampal formation, a brain structure thought to play a role in remembering cache locations. We demonstrated that changes in cache retrieval accuracy can also be triggered by experimental variation in food supply: captive mountain chickadees (Poecile gambeli) maintained on limited and unpredictable food supply were more accurate at recovering their caches and performed better on spatial memory tests than birds maintained on ad libitum food. In this study, we investigated whether these two treatment groups also differed in the volume and neuron number of the hippocampal formation. If variation in memory for food caches correlates with hippocampal size, then our birds with enhanced cache recovery and spatial memory performance should have larger hippocampal volumes and total neuron numbers. Contrary to this prediction we found no significant differences in volume or total neuron number of the hippocampal formation between the two treatment groups. Our results therefore indicate that changes in food-caching behavior and spatial memory performance, as mediated by experimental variations in food supply, are not necessarily accompanied by morphological changes in volume or neuron number of the hippocampal formation in fully developed, experienced food-caching birds.

We studied recruitment age (i.e. the age that a gull bred for the first time) and the probability that an adult reproduced during its life in three cohorts of Western Gulls Larus occidentalis breeding on southeast Farallon Island. We examined the effects of sex, hatching date, hatching order, brood size and annual variation in food supply during the recruitment period. Modal recruitment ages of males and females were 4 and 5 years, respectively, a difference that was significant. None of the factors examined, including hatching date, hatching order and brood size, had a significant effect on recruitment age. The probability of breeding in males who experienced low food supply in their fourth and fifth years was lower than that of males who experienced favourable food supply in their fourth and fifth years. Recruitment probability (the proportion of the sample group that recruited among those available to recruit during a given year) of 4‐ and 5‐year‐old males was lower in the food‐poor 1983 El Nino than in years of favourable food supply. The only variable affecting recruitment probability of females was age: recruitment probability was greater among 5‐year‐olds than 4‐year‐olds. Recruitment age in males was more sensitive to annual variation in food supply, probably because females have more difficulty recruiting due to an excess of adult females in the population and because of the different roles in reproduction in which males are primarily responsible for provisioning chicks and maintaining the territory.

Carbon, nitrogen, and phosphorus (C, N, P) stoichiometry influences the growth of plants and nutrient cycling within ecosystems. Indeed, elemental ratios are used as an index for functional differences between plants and their responses to natural or anthropogenic variations in nutrient supply. We investigated the variation in growth and elemental content of the rootless terrestrial bromeliad Tillandsia landbeckii, which obtains its moisture, and likely its nutrients, from coastal fogs in the Atacama Desert. We assessed (1) how fog nutrient supply influences plant growth and stoichiometry and (2) the response of plant growth and stoichiometry to variations in nutrient supply by using reciprocal transplants. We hypothesized that T. landbeckii should exhibit physiological and biochemical plastic responses commensurate with nutrient supply from atmospheric deposition. In the case of the Atacama Desert, nutrient supply from fog is variable over space and time, which suggests a relatively high variation in the growth and elemental content of atmospheric bromeliads. We found that the nutrient content of T. landbeckii showed high spatio-temporal variability, driven partially by fog nutrient deposition but also by plant growth rates. Reciprocal transplant experiments showed that transplanted individuals converged to similar nutrient content, growth rates, and leaf production of resident plants at each site, reflecting local nutrient availability. Although plant nutrient content did not exactly match the relative supply of N and P, our results suggest that atmospheric nutrient supply is a dominant driver of plant growth and stoichiometry. In fact, our results indicate that N uptake by T. landbeckii plants depends more on N supplied by fog, whereas P uptake is mainly regulated by within-plant nutrient demand for growth. Overall, these findings indicate that variation in fog nutrient supply exerts a strong control over growth and nutrient dynamics of atmospheric plants, which are ubiquitous across fog-dominated ecosystems.

The finding of a previously undescribed cold-water coral reef (Banana Reef) in the Scottish Mingulay reef complex, with denser coverage of living Lophelia pertusa than the principal Mingulay 1 Reef, was the incentive for a comparative study of the food supply to the 2 reefs. Suspended particulate matter (SPM) samples from the surface and bottom water covering a tidal cycle were compared with respect to lipids, pigments, and delta C-13 and delta N-15. Lipid profiles and stable isotope signatures of SPM were compared with those of coral tissue samples. Concurrently, hydrographic measurements were conducted to track the movement of the water masses across both reefs. Between-reef differences in SPM lipid concentrations were small compared to those in coral tissue. Corals at Banana Reef had lower lipid concentrations, pointing to less favourable food conditions than at Mingulay 1. Stable isotopes signatures and lipid profiles showed that corals on both reefs feed primarily on surface algal matter, within the timeframe of our study. At Mingulay 1, fresh microalgae are supplied to the coral reef by local downwelling. This downwelling pulse is tidally advected to Banana Reef. Food conditions observed during this study at both reefs do not explain the between-reef difference in coral coverage. A speculative explanation for the denser coral coverage at the deeper Banana Reef encompasses the slightly lower temperature that exhibits lower metabolic stress on corals, in combination with a higher current speed and particle encounter rate.

Food availability is likely to influence body condition and, in turn, fitness. The intensity of this response may vary between populations of the same species on a small spatial and temporal scale. We used 5 yr of data from 6 Eurasian red squirrel Sciurus vulgaris populations from the southern Alps to explore differences in body size and body mass among neighbouring populations, in relation to habitat type and variation in food supply. We also investigated sexual dimorphism in these traits and whether phenotypic variation affects local survival and female reproductive success. Mean hind foot length, a measure of body size, did not differ between sexes but differed between areas. Seasonal variation in body mass was small with no evidence for fattening in autumn. Females were slightly heavier than males, but this difference was largely explained by mass gain of females during reproduction. The size of conifer seed crops, the major food supply, varied strongly over years and between habitats, but this variation corresponded only weakly with autumn body mass. Differences in size and mass between populations were partially explained by habitat-related differences in body size and variability of seed-crops, suggesting differential selection for smaller squirrels in spruce-larch forests against selection for larger and heavier animals in mixed broadleaves and conifer forests and in Scots pine forests with more stable seed production. The probability of reproduction by females increased with body mass, but varied strongly between habitats and years, with more females reproducing in years with rich seed-crops. In both sexes, body mass positively affected probability of settlement and length of residency. Our results suggest that in temporally variable environments that differ in overall amount of food resources, individual variation in body mass is related to habitat type, and that having a relatively high body mass, within each population, positively affects male and female settlement success and local survival, and female reproductive success.

Among the range of determinants of post‐fledging survival in altricial birds, the energy supply to the growing juveniles is likely to play a central role. However, the exact mechanisms shaping post‐fledging survival are poorly understood. Using a food supplementation experiment, we determined the effect of variation in food supply on the survival of juvenile Little Owls Athene noctua from hatching to 2 months post‐fledging. Experimental broods were food‐supplemented for 36 days during the nestling and the early post‐fledging period. The fate of 307 juveniles (95 of them provided with extra food) was determined by nest monitoring and radiotelemetry. In unsupplemented birds, the rates of survival measured at 5‐day intervals were lowest during the nestling stage, remained low during the early post‐fledging stage and steadily increased after about 2 weeks post‐fledging. Food supplementation substantially increased nestling survival, but we detected no direct treatment effect on post‐fledging survival. Instead, we found a strong indirect effect of food supplementation, in that fledglings of good physical condition had markedly higher chances of surviving the post‐fledging period compared with those in poor condition. Experimental food supplementation increased survival over the first 3 months from 45% to 64.6%. This suggests that energy reserves built up during the nestling stage influence post‐fledging survival and ultimately parental reproductive output. The low nestling and post‐fledging survival shows that the early life‐history stages constitute a crucial bottleneck of reproductive ecology in Little Owls. The strong treatment effects on the number of independent offspring indicate that natural variation in food supply is an important determinant of spatio‐temporal patterns in Little Owl demography.

Food availability is likely to influence body condition and, in turn, fitness. The intensity of this response may vary between populations of the same species on a small spatial and temporal scale. We used 5 yr of data from 6 Eurasian red squirrel Sciurus vulgaris populations from the southern Alps to explore differences in body size and body mass among neighbouring populations, in relation to habitat type and variation in food supply. We also investigated sexual dimorphism in these traits and whether phenotypic variation affects local survival and female reproductive success. Mean hind foot length, a measure of body size, did not differ between sexes but differed between areas. Seasonal variation in body mass was small with no evidence for fattening in autumn. Females were slightly heavier than males, but this difference was largely explained by mass gain of females during reproduction. The size of conifer seed crops, the major food supply, varied strongly over years and between habitats, but this variation corresponded only weakly with autumn body mass. Differences in size and mass between populations were partially explained by habitat‐related differences in body size and variability of seed‐crops, suggesting differential selection for smaller squirrels in spruce‐larch forests against selection for larger and heavier animals in mixed broadleaves and conifer forests and in Scots pine forests with more stable seed production. The probability of reproduction by females increased with body mass, but varied strongly between habitats and years, with more females reproducing in years with rich seed‐crops. In both sexes, body mass positively affected probability of settlement and length of residency. Our results suggest that in temporally variable environments that differ in overall amount of food resources, individual variation in body mass is related to habitat type, and that having a relatively high body mass, within each population, positively affects male and female settlement success and local survival, and female reproductive success.

Colonies of Common Puffins (Fratercula arctica), differing in numbers of breeding Herring Gulls (Larus argentatus) present, also differed in puffin numbers, burrow placement, and burrow activity rate. All differences indicated that puffins avoided gulls. Sites differing in gull numbers did not differ, however, in puffin fledging success or weights of fledged chicks. Chick weights and fledging success were low. The pattern of weight gain by chicks was irregular over the season, although consistent among sites for individual periods. These results imply that food usually was hard to find, although occasionally locally abundant. Pressures on puffin reproductive efforts due to variation in food supply and levels of kleptoparasitism do not appear to be additive; rather, low food availability decreases the effect of gull kleptoparasitism as well.

Food supply fluctuations constrain group sizes of kangaroos and in turn shape their vigilance and feeding strategies

Hydrodynamic Control of the Supply of Suspended Chlorophyllato Infaunal Estuarine Bivalves

Seasonal variations in food supply for wild primates Seasonality and human population biology Food, seasonality, and human evolution Food as a function of seasonal environment and social history Efe pygmies of north-east Zaire - subsistence strategies in the Ituri Food supply uncertainty among the Aka Pygmies (Lobaye, Central African Republic) Tradition and change - problems and persistence in the Inuit diet Food storage among hunter-gatherers - more or less security in the way of life? Coping with uncertainty in food supply among the Oto and the Twa living in the equatorial flooded forest near Lake Tumba, Zaire Coping with seasonal fluctuations in food among savanna populations - the Massa and Mussey of Chad and Cameroon Coping with the market - uncertainty and food security among Hausa peasants Health and energy requirements of nomadic Turkana pastoralists Seasonality, climatic fluctuations, and food supplies (Sahelian nomadic pastoral societies) Nutrition and socio-economic strategies in pastoral societies in the Middle East with special reference to west Afghan Pashtuns How non-food producing nomads obtain their food - peripatetic strategies in Afghanistan Coping with uncertainty in food supply - case studies among the pastoral and non-pastoral nomads of western India Central Indian tribal societies under stress of modern socio-economic pressures - strategies to face the challenge Uncertainties in food supply, and nutritional deficiencies, in relation to economic conditions, in a village population of southern West Bengal, India Food allocation within the family - response to fluctuating food supply and food needs A framework for analysing uncertainty in highland areas Discussion and conclusions.