Non-breeding season habitat quality mediates the strength of density-dependence for a migratory bird

Our understanding of when natural populations are regulated during their annual cycle is limited, particularly for migratory species. This information is needed for parametrizing models that can inform management and conservation. Here, we use 14 years of data on colour-marked birds to investigate how conspecific density and habitat quality during the tropical non-breeding period interact to affect body condition and apparent annual survival of a long-distance migratory songbird, the American redstart (Setophaga ruticilla). Body condition and survival of birds in high-quality mangrove habitat declined as density increased. By contrast, body condition improved and survival did not vary as density increased in adjacent, lower quality scrub habitat, although mean condition and survival were almost always lower than in mangrove. High rainfall enhanced body condition in scrub but not in mangrove, suggesting factors such as food availability outweighed consequences of crowding in lower quality habitat. Thus, survival of overwintering redstarts in mangrove habitat, disproportionately males, appears to be regulated by a crowding mechanism based on density-dependent resource competition. Survival of individuals in scrub, mostly females, appears to be limited by density-independent environmental factors but not regulated by crowding. The contrasting effects of density and food limitation on individuals overwintering in adjacent habitats illustrate the complexity of processes operating during the non-breeding period for migratory animals, and emphasize the need for long-term studies of animals in multiple habitats and throughout their annual cycles.     

Does winter region affect spring arrival time and body mass of king eiders in northern Alaska?

Events during the non-breeding season may affect the body condition of migratory birds and influence performance during the following breeding season. Migratory birds nesting in the Arctic often rely on endogenous nutrients for reproductive efforts, and are thus potentially subject to such carry-over effects. We tested whether king eider (Somateria spectabilis) arrival time and body mass upon arrival at breeding grounds in northern Alaska were affected by their choice of a winter region in the Bering Sea. We captured birds shortly after arrival on breeding grounds in early June 2002–2006 at two sites in northern Alaska and determined the region in which individuals wintered using satellite telemetry or stable isotope ratios of head feathers. We used generalized linear models to assess whether winter region explained variation in arrival body mass among individuals by accounting for sex, site, annual variation, and the date a bird was captured. We found no support for our hypothesis that either arrival time or arrival body mass of king eiders differed among winter regions. We conclude that wintering in different regions in the Bering Sea is unlikely to have reproductive consequences for king eiders in our study areas.     

Density-mediated carry-over effects explain variation in breeding output across time in a seasonal population

In seasonal environments, where density dependence can operate throughout the annual cycle, vital rates are typically considered to be a function of the number of individuals at the beginning of each season. However, variation in density in the previous season could also cause surviving individuals to be in poor physiological condition, which could carry over to influence individual success in the following season. We examine this hypothesis using replicated populations of Drosophila melanogaster, the common fruitfly, over 23 non-overlapping generations with distinct breeding and non-breeding seasons. We found that the density at the beginning of the non-breeding season negatively affected the fresh weight of individuals that survived the non-breeding season and resulted in a 25% decrease in per capita breeding output among those that survived to the next season to breed. At the population level, per capita breeding output was best explained by a model that incorporated density at the beginning of the previous non-breeding season (carry-over effect, COE) and density at the beginning of the breeding season. Our results support the idea that density-mediated COEs are critical for understanding population dynamics in seasonal environments.     

The effects of rainfall on different components of seasonal fecundity in a tropical forest passerine

Seasonal fecundity is a composite metric that is determined by component parameters such as clutch size, nest survival and re‐nesting probability. Many of these component parameters are known to vary with environmental conditions, in particular rainfall prior to or during the breeding season. In some species, seasonal fecundity is positively related to rainfall, but little is known about which component parameters of seasonal fecundity respond most strongly to rainfall. We used intensive nest monitoring of a multi‐brooded tropical forest passerine, the Montserrat Oriole Icterus oberi, to examine the effects of rainfall during the pre‐breeding season on component parameters of annual fecundity. We monitored all nests of a total of 42 pairs over 5 years in which rainfall varied substantially. We then related clutch size, nest survival, onset and length of the breeding season, re‐nesting probability and re‐nesting interval to pre‐breeding season rainfall using generalized linear mixed models that accounted for random variation across sites and individual pairs, and incorporated other variables known to affect the response. Higher pre‐breeding season rainfall led to an increase in clutch size and a decrease in re‐nesting interval, but nest survival, re‐nesting probability and length of the breeding season were not affected by variation in rainfall. The onset of the breeding season was delayed in very dry years. We conclude that higher rainfall is likely to increase food availability and thus body condition of female Montserrat Orioles, leading to an increase in fecundity due to larger clutch sizes.     

Seasonal changes in testosterone and corticosterone levels in four social classes of a desert dwelling sociable rodent

Animals have to adjust their physiology to seasonal changes, in response to variation in food availability, social tactics and reproduction. I compared basal corticosterone and testosterone levels in free ranging striped mouse from a desert habitat, comparing between the sexes, breeding and philopatric non-breeding individuals, and between the breeding and the non-breeding season. I expected differences between breeders and non-breeders and between seasons with high and low food availability. Basal serum corticosterone was measured from 132 different individuals and serum testosterone from 176 different individuals of free living striped mice. Corticosterone and testosterone levels were independent of age, body weight and not influenced by carrying a transmitter. The levels of corticosterone and testosterone declined by approximately 50% from the breeding to the non-breeding season in breeding females as well as non-breeding males and females. In contrast, breeding males showed much lower corticosterone levels during the breeding season than all other classes, and were the only class that showed an increase of corticosterone from the breeding to the non-breeding season. As a result, breeding males had similar corticosterone levels as other social classes during the non-breeding season. During the breeding season, breeding males had much higher testosterone levels than other classes, which decreased significantly from the breeding to the non-breeding season. My results support the prediction that corticosterone decreases during periods of low food abundance. Variation in the pattern of hormonal secretion in striped mice might assist them to cope with seasonal changes in energy demand in a desert habitat.     

Food limitation and demography of a migratory antelope,the white-eared kob

Summary Although it is commonly presumed that many populations of large-herbivores are limited by food availability, supporting evidence is scarce. This hypothesis was investigated in a population of over 800,000 white-eared kob in the Boma National Park region of the southern Sudan. Food availability, body condition, and mortality rates of adults and calves were measured during the dry seasons of 1982 and 1983. Sampled age distributions from both the live population and carcasses were used to calculate age-specific rates of mortality. In 1982, food supplies during the dry season were augmented by substantial rainfall, which produced regrowth of grass in areas that ordinarily had little green forage. As a result, fat reserves declined little, and rates of adult mortality showed no increasing trend. Total adult mortality was 5%. In 1983, there was no rainfall during the dry season and food intake was insufficient to meet the estimated energy requirements of kob. As a result, fat reserves declined and adult mortality rates increased fourfold. Total adult mortality was 10% (equivalent to the recruitment rate of yearling into the population). Calf mortality during the dry season was similar in both years (50%), based on field estimates of mortality rates and calf/female ratios. Lactation throughout the dry season possibly provided a buffer for calves against variations in food availability. The age structure of the live population in 1983 suggests that a drought in 1980 reduced kob numbers by 40%. These results suggest that adult survival is influenced strongly by the availability of food during the dry season. However, the duration of the dry season also plays an important role. During the dry season, declining fat reserves make an increasing proportion of the population vulnerable to mortality. As a result, even moderate droughts may lead to substantial changes in population numbers.     

Diet and habitat use of frillneck lizards in a seasonal tropical environment

A population of frillneck lizards, Chlamydosaurus kingii, was monitored by mark-recapture and telemetry over a 2 year period in Kakadu National Park. The aims of the study were to document changes in diet, growth, condition and habitat use between the wet and dry seasons of northern Australia, in light of recent research examining seasonal variation in the physiology of this species. Frillneck lizards feed on a diverse range of invertebrates in both seasons, even though there is a substantial reduction in food avail-ability in the dry season. Harvester termites from the genus Drepanotermes constitute a major component of the diet, and the prevalence of termites in the diet of sedentary foragers in a tropical environment is unusual. Adult male body condition remained relatively stable throughout the year, but females experienced considerable variation. These differences are attributed to different reproductive roles of the sexes. Growth in C. kingii was restricted to the wet season, when food availability was high, and growth was minimal in the dry season when food availability was low. The method used in catching lizards was an important factor in determining seasonal habitat use. Telemetered lizards selected a significantly different distribution of tree species than was randomly available, and they selected significantly larger tree species during the dry season. Lizards spotted along roadsides showed little seasonal variation in the selection of tree species or tree sizes. The results suggest a comprehensive change in the ecology of this species, in response to an annual cycle of low food and moisture availability, followed by a period with few resource restrictions.     

Space‐use strategies of wintering Ovenbirds in Belize: causes and consequences

For long‐distance migrants, territoriality and prey biomass during the non‐breeding season have been linked to body condition that can carry over to affect spring migration and breeding events. For Ovenbirds (Seiurus aurocapilla), studies in Jamaica showed that body condition in mid‐season depended on leaf litter prey biomass and declined seasonally as conditions became drier, and that individuals with sedentary (territorial) and wandering space‐use strategies did not differ in age or body condition. During October and November 2010–2011, we radio‐tracked Ovenbirds in Belize to determine if space‐use patterns differed with age and sex, if space‐use strategy influenced foraging behavior or body condition, and if areas used by wanderers and territory owners differed in food abundance or habitat characteristics. Most Ovenbirds (41 of 51, 80.4%) possessed small (1‐ha) territories with largely non‐overlapping cores, but 22.5% (10 of 51) of birds were wanderers (～7‐ha home range). Early season space use was predicted by age class, with most wanderers (90%, 9 of 10) being first‐year birds and most territory owners (63%) being older birds. Sex ratios of wanderers and territory owners did not differ. We found that wanderers may have been at a disadvantage because they had significantly lower foraging rates and lower relative body masses than territorial birds, although baseline corticosterone levels did not differ. Habitat characteristics of areas used by wanderers and territory owners did not differ nor did biomass of ground‐surface arthropods, likely because ranges of most wanderers overlapped those of territory owners. Using stable hydrogen isotopes in feathers, we found that first‐year Ovenbirds that were wanderers tended to have a more northern natal origins than sedentary birds, though the difference was not significant. Longer migration distances could delay arrival and reduce competitive ability. Our results suggest that wandering may not be an alternative and equally successful strategy, at least early in the season, and instead young birds may be competitively excluded from territory ownership.     

Variation in a sparrow's reproductive success with rainfall: food and predator-mediated processes

From 1997 to 1999, we monitored the reproductive success of individual rufous-crowned sparrows (Aimophila ruficeps) in coastal sage scrub habitat of southern California, USA. Annual reproductive output of this ground-nesting species varied strongly with annual variation in rainfall, attributed to the El Niño-Southern Oscillation. Birds fledged 3.0 young per breeding pair in 1997, when rainfall was near the long-term mean, 5.1 offspring per pair in 1998, a wet El Niño year, and 0.8 fledglings per pair in 1999, a dry La Niña year. Variation in many components of reproductive output was consistent with the hypothesis that food availability was positively correlated with rainfall. However, the factor most responsible for the high reproductive output in 1998 was low early season nest predation which, combined with favorable nesting conditions, enabled more pairs to multiple-brood. Cool, rainy El Niño conditions may have altered the activity of snakes, the main predator of these nests, in the early season of 1998. Overall, more of the annual variation in fecundity was attributable to variation in within-season components of reproductive output (mean number of nests fledged per pair) than to within-nest components (mean brood size). Annual variation in rufous-crowned sparrow fecundity appears to be driven primarily by food resource-mediated processes in La Niña years and by predator-mediated processes in El Niño years.     

European Swallows Hirundo rustica in Botswana during three non-breeding seasons: the effects of rainfall on moult

van den Brink, B., Bijlsma, R.G. & van der Have, T.M. 2000. European swallows Hirundo rustica in Botswana during three non-breeding seasons: the effects of rainfall on moult. Ostrich: 71 (1): 198–204.

The rate of moult of European Swallows spending the non-breeding season in Botswana was studied during December-January of 1992/93,1993/94 and 1994/95 to investigate the effects of variability in rainfall and roosting habitat availability. In January 1994, 2–3 million European Swallows were counted at a traditional roost along the Boteti River. The rate of moult was relatively slow, about one feather (primary, secondary or tail feather) was replaced every two weeks in both adults and juveniles. The speed of moult in juveniles was generally lower than in adults, in particular of secondaries and tail feathers. Moulting rate of both primaries and tail feathers was lowest in 1994/95 during a period of drought and coincided with the almost complete destruction of roosting habitat. In 1992/93, moulting rate was highest when rainfall was moderate and roosting habitat abundant. Moulting rate was intermediate in 1993/94 when rainfall was frequent but roosting habitat reduced because of the low water level in the Boteti River. The combined effect of reduced food availability during droughts and higher densities and longer foraging flights when roosting habitat is scarce might explain the annual variation in moulting rate. From the second week of January onwards many adults started moulting the outermost tail feather before the penultimate feathers. This phenomenon could indicate the importance of long tail streamers in aerial manoeuvring when foraging during the return migration to the breeding grounds.     

Migrants in tropical bird communities: the balanced breeding limitation hypothesis

Explanations for the ecological integration of migratory and non-migratory (resident) insectivorous birds in the tropics have been complicated by the paradox that arthropod abundances are low when bird abundances reach their annual peak. The breeding currency hypothesis and the nest predation hypothesis both account for this paradox by postulating that residents are held below the non-breeding season carrying capacity, which frees resources available for migratory insectivores. The breeding currency hypothesis suggests residents are limited by food suitable for nestlings, whereas the nest predation hypothesis emphasizes the primacy of high rates of nest predation. However, theoretical arguments suggest that food availability and predation risk interact strongly to limit breeding birds. We use graphical analyses to extend the breeding currency hypothesis to incorporate effects of nest predation. This yields a more synthetic and realistic model for the integration of migrant and resident insectivores in the tropics – the balanced breeding limitation hypothesis.     

Supplementary fed Ural owls increase their reproductive output with a one year time lag

Life-history components may be food-limited. We supplemented food to 18 Ural owl, Strix uralensis, nests during the nestling period. Food supplementation led to a higher somatic condition in the female parent, but effects in males were moderate. Parents delivered less food to fed nests than to control nests. Offspring survival and fledging condition did not differ between control and fed nests. In the season following food supplementation, fed pairs bred 1 week earlier than control pairs and, coupled to this advance in laying date, fed pairs produced 0.6 eggs more than control pairs. This is the first evidence that food limitation in the current season may constrain next seasons reproduction. Such carry-over effects of food-limitation may have important consequences for population dynamics.     

Influences of landscape composition on hunter-harvested mallard body mass and condition in eastern Arkansas

Waterfowl with more body mass and a greater body condition during the non-breeding season are thought to be more likely to survive and have increased productivity during the following breeding season. Body mass and body condition in waterfowl should reflect the resources available to them locally. We analyzed the relationship of landscape composition on mallard (Anas platyrhynchos) body mass and body condition (mass-wing length index) among age and sex groups. We calculated these variables from hunter-harvested mallards during the 2019–2020 and 2020–2021 duck hunting seasons in the Lower Mississippi Alluvial Valley of Arkansas, USA. We used linear mixed-effects models to analyze changes in body mass and body condition with changes in the percent landscape composition of water cover, woody wetlands, herbaceous wetlands, rice, soybeans, and disturbance. We found that body mass and condition of harvested mallards were positively associated with greater proportions of water cover and woody wetlands but negatively associated with greater proportions of herbaceous wetlands and human disturbance from human infrastructure. Management actions focused on providing flooded and woody wetland areas on the landscape that allow waterfowl to access food resources, while decreasing the disturbance around wetlands in the form of road density and human infrastructure, should increase body mass and body condition in mallards spending the non-breeding season in the Lower Mississippi Alluvial Valley.     

Contributions of juvenile and adult diet to the lifetime reproductive success and lifespan of a spider

Food availability can vary widely for animals in nature and can have large effects on growth, reproduction and survival. While the consequences of food limitation for animals have been extensively studied, significant questions still remain including how ontogenetic variation in food availability contributes to lifetime reproductive success. We tested the effects of juvenile and adult food limitation on the lifetime reproductive success and lifespan of bridge spiders, Larinioides sclopetarius. Food availability was manipulated (low or high) over the entire juvenile and adult stage in a full‐factorial design and reproductive output and lifespan were measured. Juvenile and adult food limitation both reduced lifetime egg and hatchling production with effect sizes that were not significantly different from each other. Unlike some other arthropods, where juvenile food limitation reduces fecundity by reducing adult body size, body size was not affected by juvenile diet in bridge spiders. Clutch size was also significantly reduced by both juvenile and adult food limitation. The effect of adult diet on clutch size was stronger than that of juvenile diet. Juvenile and adult food limitation both extended total lifespan, and adult food limitation extended adult longevity (i.e. time from maturation to death). However, juvenile food limitation decreased adult longevity, in contrast to what would be predicted by dietary or caloric restriction. Compensatory feeding and growth are widely recognized mechanisms through which animals can ameliorate some of the negative effects of periods of food limitation. Yet our results combined with studies of a range of other species suggest that there may be lasting consequences of juvenile food limitation on lifetime reproductive success that cannot be compensated for by adult feeding in some species.     

The breeding colouration of male three-spined sticklebacks (Gasterosteus aculeatus) as an indicator of energy investment in vigour

Summary A necessary condition of most models of intersexual selection requires that secondary sexual traits are costly so that cheating is prevented. If the conspicuous breeding colouration of male three-spined sticklebacks (Gasterosteus aculeatus L.) is such a handicap, it must involve costs. I examined the energetic costs of the breeding colouration by varying the energy contents of the daily food supply among five groups of sticklebacks over a 10 week period. The nutritional carotenoid level, i.e. the colour pigment used in the breeding colouration, was constant for all fish. Both the increase of their condition factor and the condition level they finally achieved correlated positively with the food ration of the groups. Individuals whose condition increased during the experiment developed a more intensive red colouration. However, a direct correlation between food quantity and the red breeding colouration reached at the end of the experiment did not exist. Nevertheless, given the limitation of pigment availability, there was still variation in the breeding colouration and the costs for the metabolism of the colouration were sufficient to render it an honest signal: a female stickleback can assess a male's condition and condition change over the past few weeks by the intensity of the colour of his blue eyes (which is not based on carotenoids and whose pigments were therefore not controlled in the food) and his red jaw, respectively. How much an individual male fish invests in increase of length and increase of condition (which correlate negatively with each other) seems to be, at least partly, his own strategic decision, which could have important consequences in the competition for female mates. It is eventually this decision that a male stickleback seems to signal with his red jaw.     

Context-dependent specialisation drives temporal dynamics in intra- and inter-individual variation in foraging behaviour within a generalist bird population

Individual niche variation is common within animal populations, and has significant implications for a wide range of ecological and evolutionary processes. However, individual niche differences may also temporally vary as a result of behavioural plasticity. While it is well understood how niche variation is affected by changes in resource availability, comparatively little is known about the extent to which individual niche differences may vary within the annual cycle due to internal drivers. Here, we assess how time- and energy-constraints imposed by incubating and brood rearing affect inter- and intra-individual variation in the foraging behaviour of lesser black-backed gulls, a generalist seabird with strong individual niche variation. To this end, we compared daily foraging trips of 22 breeding and 23 non-breeding GPS-tracked adult gulls from two colonies in the Southern Bight of the North Sea over the course of the breeding season. We find that breeding birds, unlike non-breeding ones, did indeed alter their foraging behaviour during the breeding season. Both sexes reduced their searching effort by increasingly revisiting earlier foraging locations, allowing for shorter and more frequent foraging trips. Breeding females also showed pronounced shifts in their habitat use and strongly specialised on urbanised foraging habitats throughout the breeding season. Hence, while individual variation in habitat use remained largely consistent within non-breeders and in breeding males, individual variation among breeding females almost completely disappeared. Female lesser black-backed gulls are on average smaller, and therefore often outcompeted by males for the most profitable food sources. The temporal specialisation on spatially reliable anthropogenic food sources during breeding hence suggests a complex interplay between intrinsic competitive constraints, resource reliability and shifting time- and energy budges in shaping temporal dynamics in individual niche variation within our study population.     

Birth seasonality in cotton-top tamarins (saguinus oedipus) despite constant food supply and body weight

Goldizen et al. (1988) reported that wild saddle-back tamarins (Saguinus fuscicollis, Callitrichidae) show birth seasonality that is correlated with food supply and body weight. They suggested a sequence of ultimate causality in which shortage of food leads to reduced body weight which leads to timing of weaning and lactation when resources are more abundant. Cotton-top tamarins in captivity show birth seasonality despite constant food supply and body weight. Although natural availability of fruit and insects (which are key foods for tamarins) is related to rainfall, birth seasonality and body weight in captive cotton-top tamarins are unrelated to rainfall. The most likely proximate mechanism for seasonality of births in tamarins is photo-period, given existing data on populations living in natural and artificial lighting.     

Carry-over effects,sequential density dependence and the dynamics of populations in a seasonal environment

Most animal populations have distinct breeding and non-breeding periods, yet the implications of seasonality on population dynamics are not well understood. Here, we introduce an experimental model system to study the population dynamics of two important consequences of seasonality: sequential density dependence and carry-over effects (COEs). Using a replicated seasonal population of Drosophila, we placed individuals at four densities in the non-breeding season and then, among those that survived, placed them to breed at three different densities. We show that COEs arising from variation in non-breeding density negatively impacts individual performance by reducing per capita breeding output by 29–77%, implying that non-lethal COEs can have a strong influence on population abundance. We then parametrized a bi-seasonal population model from the experimental results, and show that both sequential density dependence and COEs can stabilize long-term population dynamics and that COEs can reduce population size at low intrinsic rates of growth. Our results have important implications for predicting the successful colonization of new habitats, and for understanding the long-term persistence of seasonal populations in a wide range of taxa, including migratory organisms.     

Bridging Developmental Boundaries: Lifelong Dietary Patterns Modulate Life Histories in a Parthenogenetic Insect

Determining the effects of lifelong intake patterns on performance is challenging for many species, primarily because of methodological constraints. Here, we used a parthenogenetic insect (Carausius morosus) to determine the effects of limited and unlimited food availability across multiple life-history stages. Using a parthenogen allowed us to quantify intake by juvenile and adult females and to evaluate the morphological, physiological, and life-history responses to intake, all without the confounding influences of pair-housing, mating, and male behavior. In our study, growth rate prior to reproductive maturity was positively correlated with both adult and reproductive lifespans but negatively correlated with total lifespan. Food limitation had opposing effects on lifespan depending on when it was imposed, as it protracted development in juveniles but hastened death in adults. Food limitation also constrained reproduction regardless of when food was limited, although decreased fecundity was especially pronounced in individuals that were food-limited as late juveniles and adults. Additional carry-over effects of juvenile food limitation included smaller adult size and decreased body condition at the adult molt, but these effects were largely mitigated in insects that were switched to ad libitum feeding as late juveniles. Our data provide little support for the existence of a trade-off between longevity and fecundity, perhaps because these functions were fueled by different nutrient pools. However, insects that experienced a switch to the limited diet at reproductive maturity seem to have fueled egg production by drawing down body stores, thus providing some evidence for a life-history trade-off. Our results provide important insights into the effects of food limitation and indicate that performance is modulated by intake both within and across life-history stages.     

Benefits of extra food to reproduction depend on maternal condition

The amount of food resources available to upper‐level consumers can show marked variations in time and space, potentially resulting in food limitation. The availability of food resources during reproduction is a key factor modulating variation in reproductive success and life‐history tradeoffs, including patterns of resource allocation to reproduction versus self‐maintenance, ultimately impacting on population dynamics. Food provisioning experiments constitute a popular approach to assess the importance of food limitation for vertebrate reproduction. In this study of a mesopredatory avian species, the lesser kestrel Falco naumanni, we provided extra food to breeding individuals from egg laying to early nestling rearing. Extra food did not significantly affect adult body condition or oxidative status. However, it increased the allocation of resources to flight feathers moult and induced females to lay heavier eggs. Concomitantly, it alleviated the costs of laying heavier eggs for females in poor body condition, and reduced their chances of nest desertion (implying complete reproductive failure). Extra food provisioning improved early nestling growth (body mass and feather development). Moreover, extra food significantly reduced the negative effects of ectoparasites on nestling body mass, while fostering forearm (a flight apparatus trait) growth among highly parasitized nestlings. Our results indicate that lesser kestrels invested the extra food mainly to improve current reproduction, suggesting that population growth in this species can be limited by food availability during the breeding season. In addition, extra food provisioning reduced the costs of the moult–breeding overlap and affected early growth tradeoffs by mitigating detrimental ectoparasite effects on growth and enhancing development of the flight apparatus with high levels of parasitism. Importantly, our findings suggest that maternal condition is a major trait modulating the benefits of extra food to reproduction, whereby such benefits mostly accrue to low‐quality females with poor body condition.