Age-related shifts in the diet composition of southern elephant seals expand overall foraging niche

Southern elephant seals are important apex predators in a highly variable and unpredictable marine environment. In the presence of resource limitation, foraging behaviours evolve to reduce intra-specific competition increasing a species’ overall probability of successful foraging. We examined the diet of 141 (aged 1–3 years) juvenile southern elephant seals to test the hypotheses that differences between ages, sexes and seasons in diet structure occur. We described prey species composition for common squid and fish species and the mean size of cephalopod prey items for these age groups. Three cephalopod species dominated the stomach samples, Alluroteuthis antarcticus, Histioteuthis eltaninae and Slosarczykovia circumantarcticus. We found age-related differences in both species composition and size of larger prey species that probably relate to ontogenetic changes in diving ability and haul-out behaviour and prey availability. These changes in foraging behaviour and diet are hypothesised to reduce intra-specific food competition concomitant with the increase in foraging niche of growing juveniles.     

Shift in foraging grounds and diet broadening during ontogeny in southern elephant seals from Kerguelen Islands

Highly dimorphic species like southern elephant seals (Mirounga leonina, SES hereafter) frequently exhibit resource partitioning according to sex and/or age classes. We measured carbon and nitrogen stable isotopes (δ¹³C and δ¹⁵N) of 404 blood samples (136 males and 268 females from Kerguelen Islands, 49°21′S, 70°18′E) from 2004 to 2011. Assuming that the distribution of carbon isotopes (δ¹³C value) reflects the two main foraging grounds (Polar Frontal and Antarctic Zones), we quantified the proportion of SES foraging within each zone in relation with size, a proxy for their age. We found a clear shift from Polar Frontal to Antarctic waters as male SES aged, but no relation as far as females is concerned. We also observed a widening range of nitrogen isotopic (δ¹⁵N) values, suggesting that both males and females expanded their diet spectra with age. Whereas males increased their trophic level, females remained constant on average, with some adult females feeding both at lower and at higher trophic levels than juveniles.     

Harem choice and breeding experience of female southern elephant seals influence offspring survival

Female mammals can increase their lifetime fitness through modification of investment potential and by providing better rearing environments with improved breeding experience. We examined the relationships between reproductive fitness and the behavioural decisions that female southern elephant seals (Mirounga leonina) made during the breeding season. We examined whether mother age and breeding experience influenced reproductive success (measured as 1st-year survival probability), and whether there was a change in the choice of harem size with increasing age. Pups produced by young mothers had lower 1st-year survival probability than pups produced by older mothers. A significant increase in mean female mass with age required an analysis of both these effects on offspring survival. There was a significant positive effect of both female age and mass, and the interaction between the two, on 1st-year pup survival. The proportion of young mothers (<5 years old) decreased and the proportion of older mothers (>6 years old) increased with increasing harem size (harems surveyed from 1997 to 2001). Females chose larger harems in which to breed as they aged. Females demonstrated fidelity to breeding areas among successive breeding seasons, with older females displaying greater breeding-site fidelity than younger females. The mean number of previous breeding attempts per female within a harem (breeding experience) increased significantly with increasing harem size. Breeding females returned to breed later in the breeding season as they aged—we hypothesize that young, subordinate females gain a priority advantage by returning earlier. These results lend support to the hypothesis that there are fitness advantages, in terms of offspring survival, that are conferred to females that breed in successively larger harems with age. Potential mechanisms that select for females to improve their breeding conditions include improved mate selection and the avoidance of conspecific harassment in harems.Communicated by F. Trillmich     

Water column use and forage strategies of female southern elephant seals from Marion Island

The at-sea behaviour of marine top predators provides valuable insights into the distribution of prey species and strategies used by predators to exploit patchily distributed resources. We describe the water column usage and dive strategies of female southern elephant seals from Marion Island tracked between 2004 and 2008. Dives representing increases in forage effort were identified using a method that combines dive type analyses and the calculation of relative amounts of time that animals spend in the bottom phases of dives. Results from this analysis indicate that female elephant seals from Marion Island tend to display lower levels of forage effort closer to the island and display intensive opportunistic forage bouts that occur at a minimum distance of approximately 215 km from the island. Females from Marion Island dived deeper and for longer periods of time, compared to females from other populations. Most animals displayed positive diel vertical migration, evidently foraging pelagically on vertically migrating prey. A few animals displayed periods of reverse (negative) diel vertical migration, however, diving to deeper depths at night, compared to daytime. This behaviour is difficult to explain and prey species targeted during such periods unknown. Our results illustrate plasticity in foraging behaviour of southern elephant seals, as well as inter-population differences in forage strategies.     

Size and seasonal influences on the foraging range of female grey seals in the northeast Atlantic

Evidence of segregation in foraging habitat has been demonstrated in some top marine predators, including cetaceans, pinnipeds and seabirds. However, most data are not adequate to assess differences relating to body size or seasonal influences. This has implications for quantitative modelling of population-level predator–prey interactions and ecosystem structure. We examined potential influence of body size and ‘fatness’ on the foraging trip characteristics of a top marine predator, the Atlantic grey seal (Halichoerus grypus), in southwest Ireland within the framework of optimal foraging theory to examine how female grey seals foraging behaviour varied with size, and across the period between moult and breeding. Larger seals undertook trips of greater duration and travelled further from haul-out sites than smaller seals. However, body fat was negatively associated with trip duration and extent. Seals spent more time at sea during the summer, but trips were shorter in extent, suggesting more localized foraging during this season.     

The cost of success: reproductive effort in male southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>)

Reproductive effort is a key parameter of life history because it measures the resources allocated to reproduction at the expense of growth and maintenance. Male reproductive effort always had a minor role with respect to female effort both in the development of theories and in field research. Elephant seals are an ideal subject for reproductive effort studies because they fast during the breeding season, splitting the phase of energy acquisition from the phase of energy use for breeding. In this paper, we present results on male reproductive effort (weight loss estimated by photogrammetry) in southern elephant seals (Mirounga leonina), the most dimorphic and polygynous of all mammal species. We show that total reproductive effort increases with age, with no sign of late decrease or senescence. Male reproductive effort in this species depends mostly on behavioral factors, i.e., the success in competition with other males, and the intensity of interaction with females. A large effort results in large gains in both mating success and fertilizations. The large reproductive success that a few males are able to achieve come at a big cost in terms of energy expenditure, but this cost does not seem to affect the likelihood of survival to the following breeding season.     

Contrast in the importance of arrow squid as prey of male New Zealand sea lions and New Zealand fur seals at The Snares,subantarctic New Zealand

New Zealand sea lions (Phocarctos hookeri) are threatened by incidental bycatch in the trawl fishery for southern arrow squid (Nototodarus sloanii). An overlap between the fishery and foraging sea lions has previously been interpreted as one piece of evidence supporting resource competition for squid. However, there is currently no consensus about the importance of squid in the diet of New Zealand sea lions. Therefore, we investigated this importance independently of spatial and temporal differences in squid availability through a simultaneous study with sympatric New Zealand fur seals (Arctocephalus forsteri), a species known to target arrow squid. Diet sampling at The Snares (48°01′S 166°32′E), subantarctic New Zealand, in February 2012 coincided with peak annual catch in the nearby squid fishery. Diets were deduced by analyses of diagnostic prey remains from scats (faeces) and casts (regurgitations). The contribution of each prey species to the diet was quantified using the per cent index of relative importance (% IRI) that combined frequency of occurrence, mass and number of prey items. Arrow squid was a minor component in sea lion scats (2 % IRI), and none was found in their casts. In contrast, arrow squid was the major component in fur seal scats and casts (93 and 99 % IRI, respectively). This study found that New Zealand sea lions ate minimal squid at a time when squid was clearly available as evidenced by the diet of New Zealand fur seals; hence, there was no indication of resource competition between sea lions and the squid fishery.     

The effects of predator abundance and habitat structural complexity on survival of juvenile sea urchins

We studied the effect of the abundance of predatory fishes and structural complexity of algal assemblages on the survival of juveniles of the sea urchin Paracentrotus lividus on Mediterranean infralittoral rocky bottoms. Post-settlement juveniles (2–10 mm) were placed on four distinct natural substrates with increasing structural complexity (coralline barren, algal turf, erect fleshy algal assemblages and small crevices) inside and outside the Medes Islands Marine Reserve. Predation on these sea urchins increased at greater abundance of predatory fishes, and decreased with greater structural complexity. The refuge provided by structural complexity, however, decreased with increasing size of sea urchin recruits. Predation on the smallest post-settlers was carried out almost exclusively by small fishes (<20 cm), mainly the labrid Coris julis, while the dominant predator of larger juveniles was the sparid Diplodus sargus. Our results demonstrate the cascading effects caused by the prohibition of fishing in marine reserves, and highlight the potential role of small predatory fishes in the control of sea urchin populations.Communicated by O. Kinne, Oldendorf/Luhe     

The role of behavioral responses to predators in modifying urchins' (Strongylocentrotus droebachiensis) destructive grazing and seasonal foraging patterns

We documented spatial and temporal patterns of urchins (Strongylocentrotus droebachiensis) and periwinkles (Littorina littorea) in three habitats: a persistent Laminaria longicruris and L. digitata bed; an urchin dominated barrens, and the edge of the kelp bed that formed a boundary between the two. Urchins were rare in the kelp and, when present, always large and well hidden, a pattern we interpret as a response to crab and lobster predation. Urchins were abundant in the barrens, and, in the summer when predaceous fish were active during the day, foraged only at nigh. We observed the formation of a dense urchin feeding front along the kelp bed edge, and these urchins remained exposed and feeding even during the summer. Laboratory experiments demonstrated that aggregations are an effective defense against some predators, and that the presence of crabs increases the tendency of large urchins to aggregate. We hypothesize that healthy Laminaria spp. beds persist because kelp bed associated predators keep urchins at low densities and in hiding. A reduction in predation pressure permits urchin densities to increase to the point where they form aggregations, which provide better defense than hiding. These aggregations then graze destructively on Laminaria spp., forming barrens. These barrens seem to be a new, stable configuration of the system.     

Stable isotopes document winter trophic ecology and maternal investment of adult female southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>) breeding at the Kerguelen Islands

Individual specialisation is widespread and can affect a population’s ecological and evolutionary dynamics. Whether intra-specific niche differences can influence reproductive investment was examined in a marine mammal, the southern elephant seal (Mirounga leonina), whose females were known to forage in two different areas during the austral winter. The study was conducted at Kerguelen Islands (49°21′S, 70°18′E), southern Indian Ocean, in late winter–early spring 2006. Pups were used as proxies of their mothers’ biology and combined information on their weaning mass (a proxy of females’ foraging success and short-term fitness) together with their blood δ¹³C value (a proxy of female foraging zone). First, the use of isotopic signature of pups was validated to study the female foraging ecology during their pre-breeding trip by demonstrating that δ¹³C and δ¹⁵N values of pups and their mothers were positively and linearly correlated. Then, blood samples were taken from a large number of newly-weaned pups, which were also weighed, to provide information at the population level. Estimated δ¹³C values of female seals encompassed a large range of values (from −23.7 to −19.1‰) with an unimodal frequency distribution, suggesting no contrasted foraging areas within the population. No significant relationship was found between pup weaning mass and their carbon signature, indicating no link between female foraging areas and maternal foraging success and investment. Finally, blood δ¹³C and δ¹⁵N values gave new insights into southern elephant seal ecology, suggesting that females mainly foraged north of the Polar Front where they preyed upon myctophid fish in late winter.     

Estimates of sex- and age-class-specific survival probabilities for a southern Great Barrier Reef green sea turtle population

Sex- and age-class-specific survival probabilities of a southern Great Barrier Reef green sea turtle population were estimated using a capture–mark–recapture (CMR) study and a Cormack–Jolly–Seber (CJS) modelling approach. The CMR history profiles for 954 individual turtles tagged over a 9-year period (1984–1992) were classified into three age classes (adult, subadult, juvenile) based on somatic growth and reproductive traits. Reduced-parameter CJS models, accounting for constant survival and time-specific recapture, fitted best for all age classes. There were no significant sex-specific differences in either survival or recapture probabilities for any age class. Mean annual adult survival was estimated at 0.9482 (95% CI: 0.92–0.98) and was significantly higher than survival for either subadults or juveniles. Mean annual subadult survival was 0.8474 (95% CI: 0.79–0.91), which was not significantly different from mean annual juvenile survival estimated at 0.8804 (95% CI: 0.84–0.93). The time-specific adult recapture probabilities were a function of sampling effort but this was not the case for either juveniles or subadults. The sampling effort effect was accounted for explicitly in the estimation of adult survival and recapture probabilities. These are the first comprehensive sex- and age-class-specific survival and recapture probability estimates for a green sea turtle population derived from a long-term CMR program.Communicated by M.S. Johnson, Crawley     

The importance of an infrequently flooded intertidal marsh surface as an energy source for the mummichog Fundulus heteroclitus: An experimental approach

Fundulus heteroclitus is known to ascend onto the marsh surface to feed. Our study investigated whether the marsh surface food items are a necessary source of caloric intake for the Canary Creek, Delaware, USA population of this species. Enclosure techniques were used to restrict mummichogs from the marsh surface and the growth rates of these fish were compared to those having access to the marsh surface. Growth rates were significantly higher for mummichogs allowed access to the marsh surface. Food addition and density reduction experiments showed that food availability per fish, rather than behavioral responses due to fish crowding, was responsible for the increased growth. Although food was available in the subtidal portion of the habitat, it was of insufficient quantity for fish at natural density to grow at a normal rate, and mummichogs must utilize the marsh surface for at least a portion of their energy intake.     

The importance of agricultural lands for Himalayan birds in winter

The impacts of land‐use change on biodiversity in the Himalayas are poorly known, notwithstanding widespread deforestation and agricultural intensification in this highly biodiverse region. Although intact primary forests harbor many Himalayan birds during breeding, a large number of bird species use agricultural lands during winter. We assessed how Himalayan bird species richness, abundance, and composition during winter are affected by forest loss stemming from agriculture and grazing. Bird surveys along 12 elevational transects within primary forest, low‐intensity agriculture, mixed subsistence agriculture, and intensively grazed pastures in winter revealed that bird species richness and abundance were greatest in low‐intensity and mixed agriculture, intermediate in grazed pastures, and lowest in primary forest at both local and landscape scales; over twice as many species and individuals were recorded in low‐intensity agriculture than in primary forest. Bird communities in primary forests were distinct from those in all other land‐use classes, but only 4 species were unique to primary forests. Low‐, medium‐, and high‐intensity agriculture harbored 32 unique species. Of the species observed in primary forest, 80% had equal or greater abundance in low‐intensity agricultural lands, underscoring the value of these lands in retaining diverse community assemblages at high densities in winter. Among disturbed landscapes, bird species richness and abundance declined as land‐use intensity increased, especially in high‐intensity pastures. Our results suggest that agricultural landscapes are important for most Himalayan bird species in winter. But agricultural intensification—especially increased grazing—will likely result in biodiversity losses. Given that forest reserves alone may inadequately conserve Himalayan birds in winter, comprehensive conservation strategies in the region must go beyond protecting intact primary forests and ensure that low‐intensity agricultural lands are not extensively converted to high‐intensity pastures.     

The importance of handling time for the flight directionality in bees

Summary Individual workers of the honeybee (Apis mellifera), trained to collect food from a patch of artificial flowers, were used to test the following hypotheses about the mechanism leading to a decrease in flight directionality when moving from one flower to the next. Directionality decreases with (1) an increasing amount of energy taken from a flower, (2) an increasing amount of time spent on a flower, or (3) an increasing profitability of the flower, i.e. rate of nectar uptake. The bees were tested singly on an arrangement with equidistant flowers. They could move from a decision point forward, right, left, or backwards to the next flower. The rewards at the decision point were altered. When rewards at this point were compared that offered the same quality of nectar (50% sugar solution), the time hypothesis (2) was consistent with the observed behaviour of the bees. When conditions were compared that involved two different reward qualities (50% and 25% sugar solation), none of the original hypotheses could entirely account for the results.It is suggested that a slight modification of the time hypothesis would be consistent with the observations: The bees might possibly forget their arrival direction on the flower with increasing time, but forgetting the direction of the last flower visited is intensified with more concentrated rewards.     

The importance of nestling location for obtaining food in open cup-nests

Observations and experiments were carried out at 34 four-nestling nests of Arabian babblers (Turdoides squamiceps) and feeding events were analyzed according to hatching order and nestling location in the nest. Nestlings were fed in a negative correlation with hatching order. Nestling locations in relation to the provisioners were defined as Near, Far, Right, Left, and Center. Feeders fed closer nestlings more often than those further away, and straight ahead rather than sideways. In the open circular nest, the feeding adults' positions were unpredictable and equally distributed around the nest. Each peripheral position therefore served equally as Near, Far, Right and Left positions. The Center was independent of the arrival direction of the feeders and was advantageous relative to the peripheral positions. The importance of the central position was demonstrated by three experiments. (a) Food deprivation for 1 h caused the first-hatched nestling to occupy the center position and gain more feedings than its siblings. (b) Introduction of a Perspex barrier into the nest, thus eliminating the central position and preventing the nestlings from changing places, led to a loss of importance of hatching order and equal provisioning of the nestlings. (c) Fencing the nest so that entry was possible from only one permanent direction, resulted in the first-hatched nestling occupying the near position more often than its nestmates and obtaining 52% of the feedings provisioned to the whole brood. These findings imply that the architecture of the nest has an important role in food distribution among the nestlings, and contributes to reducing inequalities in the siblings' abilities to obtain food.     

The importance of groundwater discharge for plant species number in riparian zones

Riparian zones are hotspots of plant species richness in temperate and boreal biomes. The phenomenon is believed to be caused primarily by river-related processes, and upland influences on riparian zones have received relatively little attention. We investigated the importance of discharge of groundwater derived from uplands on riparian patterns in vascular plant species composition. We found that groundwater discharge areas in riparian zones were 36-209% more species rich than non-discharge areas, depending on spatial scale (1-50 m wide transects from annual high-water levels to summer low-water levels) and river (one free-flowing and one regulated). Higher nitrogen availability and less drought stress during low river stages are suggested as the major causes for the higher species diversity in discharge areas. Riparian zones lacking groundwater discharge lost more species following water-level regulation than did discharge areas. This indicates that groundwater discharge areas are more resistant to regulation because both individual plants and plant populations may grow larger in discharge areas. These results demonstrate that riparian zones are controlled by water and nutrient input from upland parts of catchments in ways that have been overlooked despite more than three decades of research into linkages between stream ecosystems and their valleys.     

The regulation of pollen foraging by honey bees: how foragers assess the colony's need for pollen

Summary The honey bee colony presents a challenging paradox. Like an organism, it functions as a coherent unit, carefully regulating its internal milieu. But the colony consists of thousands of loosely assembled individuals each functioning rather autonomously. How, then, does the colony acquire the necessary information to organize its work force? And how do individuals acquire information about specific colony needs, and thus know what tasks need be performed? I address these questions through experiments that analyze how honey bees acquire information about the colony's need for pollen and how they regulate its collection. The results demonstrate features of the colony's system for regulating pollen foraging: (1) Pollen foragers quickly acquire new information about the colony's need for pollen. (2) When colony pollen stores are supplemented, many pollen foragers respond by switching to nectar foraging or by remaining in the hive and ceasing to forage at all. (3) Pollen foragers do not need direct contact with pollen to sense the colony's change of state, nor do they use the odor of pollen as a cue to assess the colony's need for pollen. (4) Pollen foragers appear to obtain their information about colony pollen need indirectly from other bees in the hive. (5) The information takes the form of an inhibitory cue. The proposed mechanism for the regulation of pollen foraging involves a hierarchical system of information acquisition and a negative feedback loop. By taking advantage of the vast processing capacity of large numbers of individuals working in parallel, such a system of information acquisition and dissemination may be ideally suited to promote efficient regulation of labor within the colony. Although each individual relies on only limited, local information, the colony as a whole achieves a finely-tuned response to the changing conditions it experiences.     

The importance of considering multiple interacting species for conservation of species at risk

Conservation of species at risk of extinction is complex and multifaceted. However, mitigation strategies are typically narrow in scope, an artifact of conservation research that is often limited to a single species or stressor. Knowledge of an entire community of strongly interacting species would greatly enhance the comprehensiveness and effectiveness of conservation decisions. We investigated how camera trapping and spatial count models, an extension of spatial-recapture models for unmarked populations, can accomplish this through a case study of threatened boreal woodland caribou (Rangifer tarandus caribou). Population declines in caribou are precipitous and well documented, but recovery strategies focus heavily on control of wolves (Canis lupus) and pay less attention to other known predators and apparent competitors. Obtaining necessary data on multispecies densities has been difficult. We used spatial count models to concurrently estimate densities of caribou, their predators (wolf, black bear [Ursus americanus], and coyote [Canis latrans]), and alternative prey (moose [Alces alces] and white-tailed deer [Odocoileus virginianus]) from a camera-trap array in a highly disturbed landscape within northern Alberta's Oil Sands Region. Median densities were 0.22 caribous (95% Bayesian credible interval [BCI] = 0.08–0.65), 0.77 wolves (95% BCI = 0.26–2.67), 2.39 moose (95% BCI = 0.56–7.00), 2.64 coyotes (95% BCI = 0.45–6.68), and 3.63 black bears (95% BCI = 1.25–8.52) per 100 km². (The white-tailed deer model did not converge.) Although wolf densities were higher than densities recommended for caribou conservation, we suggest the markedly higher black bear and coyote densities may be of greater concern, especially if government wolf control further releases these species. Caribou conservation with a singular focus on wolf control may leave caribou vulnerable to other predators. We recommend a broader focus on the interacting species within a community when conserving species.