High annual variability in reproductive success and survival of an Antarctic seabird,the snow petrel Pagodroma nivea

Demographic parameters were estimated for snow petrels Pagodroma nivea nesting at Pointe Géologie Archipelago, Adélie Land, Antarctica between 1963 and 1990; 21 years of data on adult survival and 27 years of data on breeding success are available. The average age of first return and first breeding were 8.1 and 9.9 years respectively and there was no signifcant difference between the sexes. The overall breeding success averaged 51.3% and was very variable between years (21–80%). Breeding failure was mostly due to incubation failure and annual breeding success was negatively correlated with average snow falls in October–November and October–March. Breeding frequency was very low, averaging 52% of seasons during a reproductive lifetime. Good quality sites, with high occupancy rate and high breeding success were few in the study plots. Poor years in 1966–1967, 1976–1977 and 1983–1984, with low breeding success, very low proportions of nets with breeding attempts and high numbers of non-breeders, occurred 1 year after large-scale El Niño Southern Oscillation (ENSO) events. Snow petrels exhibited very low philopatry. Only 45 birds have been recovered in the study plots from a total of 1115 banded fledglings giving an estimated rate of return of 12.9% between fledging and 3 years old. Annual survival between 3 and 10 years was 91.4%. Annual adult survival (93.4%), though variable, was low during poor years of 1977–1978 and 1983–1984. Adult survival of males (94.7%) was not significantly different from that of females (93.9%). Over the study period, the population of Pointe Géologie was stable. Using the estimated parameters, a Leslie model gave a growth rate of 0.948%, which was probably compensated by immigration (5.7% per year). Restricted numbers of good-quality sites at the place of birth could have led young birds to prospect other colonies and could have selected low philopatry. High adult survival, strong site tenacity and capacity to spread breeding over a long lifetime are probably part of the adaptive strategy of this small fulmarine petrel facing highly variable environmental conditions.     

Sex-specific foraging behaviour in a monomorphic seabird

Sexual differences in the foraging behaviour of parents have been observed in a number of sexually sizedimorphic birds, particularly seabirds, and the usual inference has been that these sex-specific differences are mediated primarily by differences in body size. To test this explanation, we compared the foraging behaviour of parents in a monomorphic seabird species, the northern gannet Morus bassanus. Using specially designed instruments and radio telemetry we found that individuals of both sexes were consistent in the directions and durations of their foraging trips. However, there were significant differences in the foraging behaviour of males and females. Female gannets were not only more selective than males in the areas where they foraged, but they also made longer, deeper dives and spent more time on the sea surface than males. As the sexes are morphologically similar in this species, then these differences are unlikely to have been mediated by body size. Our work highlights the need to investigate sexual differences in the foraging behaviour of seabirds and other species more closely, in order to test alternative theories that do not rely on differences in body size.     

Fine-scale spatial age segregation in the limited foraging area of an inshore seabird species,the little penguin

Competition for food resources can result in spatial and dietary segregation among individuals from the same species. Few studies have looked at such segregations with the combined effect of sex and age in species with short foraging ranges. In this study we examined the 3D spatial use of the environment in a species with a limited foraging area. We equipped 26 little penguins (Eudyptula minor) of known age, sex, and breeding output with GPS (location) and accelerometer (body acceleration and dive depth) loggers. We obtained dietary niche information from the isotopic analysis of blood tissue. We controlled for confounding factors of foraging trip length and food availability by sampling adults at guard stage when parents usually make one-day trips. We observed a spatial segregation between old (>11 years old) and middle-aged penguins (between 5 and 11 years old) in the foraging area. Old penguins foraged closer to the shore, in shallower water. Despite observing age-specific spatial segregation, we found no differences in the diving effort and foraging efficiency between age classes and sexes. Birds appeared to target similar prey types, but showed age-specific variation in their isotopic niche width. We hypothesize that this age-specific segregation was primarily determined by a “cohort effect” that would lead individuals sharing a common life history (i.e. having fledged and dispersed around the same age) to forage preferentially together or to share similar foraging limitations.     

Female reproductive success in gynodioecious Thymus vulgaris: pollen versus nutrient limitation and pollinator foraging behaviour

Gynodioecy is a dimorphic breeding system in which female individuals coexist with hermaphroditic individuals in the same population. Females only contribute to the next generation via ovules, and many studies have shown that they are usually less attractive than hermaphrodites to pollinators. Several mechanisms have been proposed to explain how females manage to persist in populations despite these disadvantages. The ‘resource reallocation hypothesis’ (RRH) states that females channel resources not invested in pollen production and floral advertisement towards the production of more and/or larger seeds. We investigated pollination patterns and tested the RRH in a population of Thymus vulgaris. We measured flower display, flower size, nectar production, visitation rates, pollinator constancy and flower lifespan in the two morphs. In addition, we measured experimentally the effects of pollen and resource addition on female reproductive success (fruit set, seed set, seed weight) of the two morphs. Despite lower investment in floral advertisement, female individuals were no less attractive to pollinators than hermaphrodites on a per flower basis. Other measures of pollinator behaviour (number of flowers visited per plant, morph preference and morph constancy) also showed that pollinators did not discriminate against female flowers. In addition, stigma receptivity was longer in female flowers. Accordingly, and contrary to most studies on gynodioecious species, reproductive success of females was not pollen limited. Instead, seed production was pollen limited in hermaphrodites, suggesting low levels of cross‐pollination in hermaphrodites. Seed production was resource limited in hermaphrodites, but not in females, thus providing support for the RRH. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 395–408.     

Sex-specific provisioning behaviour in a monomorphic seabird with a bimodal foraging strategy

Sexual differences in foraging and provisioning behaviour have been observed in several size-dimorphic seabird species. These differences are usually thought to be driven by size-related mechanisms such as the ability to compete for food or defend the nest. However, recent studies on monomorphic species suggest that sexual differences in foraging may arise independently of size. Selective forces driving sex-specific patterns are poorly known but essential to understand parental strategies. In this study, we examine sex differences in the provisioning behaviour of a monomorphic species, the Little Auk Alle alle . Using automated recording systems during two consecutive seasons at two colonies, we found that both sexes used a bimodal foraging strategy in which they regularly alternated single foraging trips of long duration with a cycle of several short-trips. The duration of long-trips was substantially longer in females than in males, and the sexes differed in the number of short-trips they performed in between long-trips, resulting in male-biased provisioning rates in both years. In species with a bimodal foraging strategy, long-trips have been interpreted as self-feeding trips to replenish body reserves. Our results therefore suggest that female Little Auks allocate more time to self-maintenance at the cost of chick provisioning, possibly due to different energetic constraints of the sexes prior and/or subsequent to chick-rearing. Our findings contribute to accumulating evidence that sex-specific foraging patterns may be widespread in sexually size-monomorphic seabird species.     

Capture and blood sampling do not affect foraging behaviour, breeding success and return rate of a large seabird: the black-browed albatross

During the last decades, eco-physiological studies have usually relied on the collection of blood from wild organisms in order to obtain relevant physiological measures. However, accurate estimates of the impact of capture and blood collection on performances of Polar seabird species have rarely been conducted. We investigated for the first time the effects of a blood sampling process on subsequent foraging behaviour, reproductive performance and return rate of black-browed albatrosses (Thalassarche melanophris) at Kerguelen Islands. We did not find any evidence that the blood sampling process as conducted in our study had detrimental effects on the breeding or foraging strategies or performance of black-browed albatrosses. Because blood collection can be performed in several different ways, we recommend that eco-physiologists conduct pilot studies to test whether their blood sampling process affects the performances of their study species.     

Sex-specific foraging behaviour in a seabird with reversed sexual dimorphism: the red-footed booby

Most hypotheses attempting to explain the evolution of reversed sexual dimorphism (RSD) assume that size-related differences in foraging ability are of prime importance, but the studies on sex-specific differences in foraging behaviour remain scarce. We compare the foraging behaviour of males and females in a seabird species with a RSD by using several miniaturised activity and telemetry loggers. In red-footed boobies males are 5% smaller and 15% lighter than females, but have a longer tail than females. Both sexes spend similar time on the nest while incubating or brooding. When foraging at sea, males and females spend similar time foraging in oceanic waters, forage in similar areas, spend similar proportion of their foraging trip in flight, and feed on similar prey—flying fishes and flying squids—of similar size. However, compared to males, females range farther during incubation (85 km vs. 50 km), and furthermore feed mostly at the extremity of their foraging trip, whereas males actively forage throughout the trip. Males are much more active than females, landing and diving more often. During the study period, males lost mass, whereas females showed no significant changes. These results indicate that males and females of the red-footed boobies differ in several aspects in their foraging behaviour. Although some differences found in the study may be the direct result of the larger size of females, that is, the slightly higher speeds and deeper depths attained by females, others indicate clearly different foraging strategies between the sexes. The smaller size and longer tail of males confer them a higher agility, and could allow them to occupy a foraging niche different from that of females. The higher foraging effort of males related to its different foraging strategy is probably at the origin of the rapid mass loss of males during the breeding period. These results suggest that foraging differences are probably the reason for the differential breeding investment observed in boobies, and are likely to be involved in the evolution and maintenance of RSD.     

Effects of floral herbivory on foraging behaviour of bumblebees and female reproductive success in Pedicularis gruina (Orobanchaceae)

The effects of floral herbivores on floral traits may result in alterations in pollinator foraging behaviour and subsequently influence plant reproductive success. Fed-upon plants may have evolved mechanisms to compensate for herbivore-related decreased fecundity. We conducted a series of field experiments to determine the relative contribution of floral herbivores and pollinators to female reproductive success in an alpine herb, Pedicularis gruina, in two natural populations over two consecutive years. Experimental manipulations included bagging, hand supplemental, geitonogamous pollination, and simulated floral herbivory. Bumblebees not only avoided damaged flowers and plants but also decreased successive visits of flowers in damaged plants, and the latter may reduce the level of geitonogamy. Although seed set per fruit within damaged plants was higher than that in intact plants, total seed number in damaged plants was less than that in intact plants, since floral herbivory-mediated pollinator limitation led to a sharp reduction of fruit set. Overall, the results suggest that resource reallocation within inflorescences of damaged plants may partially compensate for a reduction in seed production. Additionally, a novel finding was the decrease in successive within-plant bumblebee visits following floral herbivory. This may increase seed quantity and quality of P. gruina since self-compatible species exhibit inbreeding depression. The patterns of compensation of herbivory and its consequences reported in this study give an insight into the combined effects of interactions between floral herbivory and pollination on plant reproductive fitness.     

Paternal reproductive success drives sex allocation in a wild mammal

Parents should bias sex allocation toward offspring of the sex most likely to provide higher fitness returns. Trivers and Willard proposed that for polygynous mammals, females should adjust sex‐ratio at conception or bias allocation of resources toward the most profitable sex, according to their own body condition. However, the possibility that mammalian fathers may influence sex allocation has seldom been considered. Here, we show that the probability of having a son increased from 0.31 to 0.60 with sire reproductive success in wild bighorn sheep (Ovis canadensis). Furthermore, our results suggest that females fertilized by relatively unsuccessful sires allocated more energy during lactation to daughters than to sons, while the opposite occurred for females fertilized by successful sires. The pattern of sex‐biased offspring production appears adaptive because paternal reproductive success reduced the fitness of daughters and increased the average annual weaning success of sons, independently of maternal allocation to the offspring. Our results illustrate that sex allocation can be driven by paternal phenotype, with profound influences on the strength of sexual selection and on conflicts of interest between parents.     

Landfast ice: a major driver of reproductive success in a polar seabird

In a fast-changing world, polar ecosystems are threatened by climate variability. Understanding the roles of fine-scale processes, and linear and nonlinear effects of climate factors on the demography of polar species is crucial for anticipating the future state of these fragile ecosystems. While the effects of sea ice on polar marine top predators are increasingly being studied, little is known about the impacts of landfast ice (LFI) on this species community. Based on a unique 39-year time series of satellite imagery and in situ meteorological conditions and on the world''s longest dataset of emperor penguin (Aptenodytes forsteri) breeding parameters, we studied the effects of fine-scale variability of LFI and weather conditions on this species'' reproductive success. We found that longer distances to the LFI edge (i.e. foraging areas) negatively affected the overall breeding success but also the fledging success. Climate window analyses suggested that chick mortality was particularly sensitive to LFI variability between August and November. Snowfall in May also affected hatching success. Given the sensitivity of LFI to storms and changes in wind direction, important future repercussions on the breeding habitat of emperor penguins are to be expected in the context of climate change.     

Yolk carotenoids and stable isotopes reveal links among environment, foraging behavior and seabird breeding success

Nutrients that are limited in availability, such as carotenoids, are potentially involved in trade-offs between homeostasis and reproduction. Despite their importance, factors that affect the capacity of female birds to meet their carotenoid requirements are poorly understood. We used δ¹⁵N stable isotope analysis to relate foraging behavior to yolk carotenoid deposition in two seabirds, Cassin’s auklet (Ptychoramphus aleuticus) and rhinoceros auklet (Cerorhinca monocerata), during each of five years. As expected from their narrower trophic range, Cassin’s auklets produced yolks with fewer carotenoid types than did rhinoceros auklets (one vs. three). Cassin’s auklets also fed on a lower trophic level diet richer in carotenoids, yet had lower total yolk carotenoid levels, which suggests a role for species-specific adaptations for carotenoid uptake and utilization. Within both species, lower trophic-level feeding was linked to higher yolk carotenoid levels, but through different mechanisms. In Cassin’s auklets, it was due to a population-wide response to environmental variation: in warm-water years, all females fed at a low trophic level and produced carotenoid-rich yolks. In rhinoceros auklets, it was due to individual differences similarly expressed in all years: females fed across a wide trophic range, and those that fed at a low trophic level produced carotenoid-rich yolks. Rhinoceros auklets bred more successfully in years when their yolks were rich in carotenoids, probably due to a correlated response to stronger marine primary production. Our results are novel because they link avian yolk carotenoid deposition to behavioral and environmental variations.     

Berry production drives bottom–up effects on body mass and reproductive success in an omnivore

Obligate herbivores dominate studies of the effects of climate change on mammals, however there is limited empirical evidence for how changes in the abundance or quality of plant food affect mammalian omnivores. Omnivores can exploit a range of different food resources over the course of a year, but they often rely on seasonally restricted highly nutritious fruiting bodies during critical life stages. Brown bears Ursus arctos in Sweden are dependent on berries for fattening before entering hibernation. We used a ten‐year time series to evaluate the effect of temperature and snow on annual variation in berry abundance and how this variation affected bears. We found marked interannual variation in berry production of bilberry Vaccinium myrtillus and lingonberry V. vitis‐idaea, that we could attribute in part to temperature during plant dormancy and flowering and precipitation during fruit ripening. Both, autumn weights of female bears and spring weights of yearling bears increased linearly with bilberry abundance. When bilberry abundance was low, lightweight female bears had a lower reproductive success than females in better condition. This effect vanished when food abundance was above average, indicating that lightweight females could compensate for their initial weight during good bilberry years. Our study highlights the importance of considering individuals’ dynamic responses to variation in food availability, which leave some more vulnerable to food shortage than others. Individual life‐history heterogeneity in response to resource variation likely affects long‐term population recruitment. Our findings emphasize that Scandinavian bears can be dependent on a single food resource during a critical period of the year and are therefore less resilient to environmental change than expected for an omnivore. Future climate scenarios predict ambiguous trends for weather covariates that affected crucial stages of berry phenology, preventing a clear prognosis of how climate change may affect long‐term bilberry production.     

Corticosterone and foraging behavior in a pelagic seabird

Because endocrine mechanisms are thought to mediate behavioral responses to changes in the environment, examining these mechanisms is essential for understanding how long-lived seabirds adjust their foraging decisions to contrasting environmental conditions in order to maximize their fitness. In this context, the hormone corticosterone (CORT) deserves specific attention because of its major connections with locomotor activities. We examined for the first time the relationships between individual CORT levels and measurements of foraging success and behavior using satellite tracking and blood sampling from wandering albatrosses (Diomedea exulans) before (pretrip CORT levels) and after (posttrip CORT levels) foraging trips during the incubation period. Plasma CORT levels decreased after a foraging trip, and the level of posttrip CORT was negatively correlated with individual foraging success, calculated as total mass gain over a foraging trip. Pretrip CORT levels were not linked to time spent at sea but were positively correlated with daily distance traveled and maximum range at sea. In this study, we were able to highlight the sensitivity of CORT levels to variation in energy intake, and we showed for the first time that individual CORT levels can be explained by variation in foraging success. Relationships between pretrip CORT levels and daily distance traveled and maximum range were independent of pretrip body mass, suggesting that slight elevations in pretrip CORT levels might facilitate locomotor activity. However, because both foraging behavior and pretrip CORT levels could be affected by individual quality, future experimental studies including manipulation of CORT levels are needed to test whether CORT can mediate foraging decisions according to foraging conditions.     

The presence of co-flowering species facilitates reproductive success of Pedicularis monbeigiana (Orobanchaceae) through variation in bumble-bee foraging behaviour

Background and Aims The presence of co-flowering species can alter pollinator foraging behaviour and, in turn, positively or negatively affect the reproductive success of the focal species. Such interactions were investigated between a focal species, Pedicularis monbeigiana, and a co-flowering species, Vicia dichroantha, which was mediated by behaviour alteration of the shared bumble-bee pollinator. Methods Floral display size and floral colour change of P. monbeigiana were compared between pure (P. monbeigiana only) and mixed (P. monbeigiana and V. dichroantha) plots in two populations. Pollinator visitation rates, interspecific floral switching and successive within-plant pollinator visits were recorded. In addition, supplemental pollination at plant level was performed, and the fruit set and seed set were analysed in pure and mixed plots with different densities of P. monbeigiana. Key Results Pollinator visitation rates were dramatically higher in mixed plots than in pure plots. The higher pollinator visitation rates were recorded in both low- and high-density plots. In particular, successive flower visits within an individual plant were significantly lower in mixed plots. Supplemental pollination significantly increased fruit set and seed set of individuals in pure plots, while it only marginally increased seed set per fruit of plants in mixed plots. Conclusions The presence of V. dichroantha can facilitate pollination and increase female reproductive success of P. monbeigiana via both quantity (mitigating pollinator limitation) and quality (reducing geitonogamy) effects. This study suggests that successive pollinator movements among flowers within a plant, as well as pollinator visitation rates and interspecific flower switching, may be important determinants of the direction and mechanisms of interaction between species.     

Fishing in the dark: a pursuit-diving seabird modifies foraging behaviour in response to nocturnal light levels

Visual predators tend not to hunt during periods when efficiency is compromised by low light levels. Yet common murres, a species considered a diurnal visual predator, frequently dive at night. To study foraging of murres under different light conditions, we used a combination of archival tagging methods and astronomical models to assess relationships between diving behaviour and light availability. During diurnal and crepuscular periods, murres used a wide range of the water column (2-177 m), foraging across light intensities that spanned several orders of magnitude (10(3)-10(-10) Wm(-2)). Through these periods, they readily dived under conditions equivalent to ambient moonlight (~10(-4) Wm(-2)) but rarely under conditions equivalent to starlight (~10(-8) Wm(-2)). At night, murres readily foraged during both moonlit and starlit periods, and diving depth and efficiency increased with nocturnal light intensity, suggesting that night diving is at least partially visually guided. Whether visually guided foraging is possible during starlit periods is less clear. Given the dense prey landscape available, random-walk simulations suggest that murres could benefit from random prey encounters. We hypothesise that murres foraging through starlit periods rely either on close-range visual or possibly nonvisual cues to acquire randomly encountered prey. This research highlights the flexibility of breeding common murres and raises questions about the strategies and mechanisms birds use to find prey under very low light conditions.     

Effects of recruiting age on senescence, lifespan and lifetime reproductive success in a long-lived seabird

Theories of ageing predict that early reproduction should be associated with accelerated reproductive senescence and reduced longevity. Here, the influence of age of first reproduction on reproductive senescence and lifespan, and consequences for lifetime reproductive success (LRS), were examined using longitudinal reproductive records of male and female blue-footed boobies (Sula nebouxii) from two cohorts (1989 and 1991). The two sexes showed different relationships between age of first reproduction and rate of senescent decline: the earlier males recruited, the faster they experienced senescence in brood size and breeding success, whereas in females, recruiting age was unrelated to age-specific patterns of reproductive performance. Effects of recruiting age on lifespan, number of reproductive events and LRS were cohort- and/or sex-specific. Late-recruiting males of the 1989 cohort lived longer but performed as well over the lifetime as early recruits, suggesting the existence of a trade-off between early recruitment and long lifespan. In males of the 1991 cohort and females of both cohorts, recruiting age was apparently unrelated to lifespan, but early recruits reproduced more frequently and fledged more chicks over their lifetime than late recruits. Male boobies may be more likely than females to incur long-term costs of early reproduction, such as early reproductive senescence and diminished lifespan, because they probably invest more heavily than females. In the 1991 cohort, which faced the severe environmental challenge of an El Ni?o event in the first year of life, life-history trade-offs of males may have been masked by effects of individual quality.