Optimal foraging strategies in the diet of the green monkey,Cercopithecus sabaeus,at Mt. Assirik,Senegal

The feeding behavior of one group of green monkeys (Cercopithecus sabaeus)was observed between October 1978 and December 1979 in the Parc National du Niokolo-Koba, Senegal. Details of the vegetational composition of the habitat and seasonal variation in food availability were also recorded. The green monkeys’ diet was omnivorous and diverse, including over 65 species of plants, many invertebrates, and some eggs and meat. Preference was given to fruits and flowers, although particular species were not selected; rather, these foods were eaten in proportion to their availability. Leaves, gum, seeds, and fungi were secondarily preferred foods, their consumption depending mostly on the availability of fruit or flowers. There was little overlap in the composition of the diet from month to month, reflecting the strong seasonality of the environment, although there was a consistent intake of invertebrates each month. Differences in diet between populations of the superspecies C. aethiopsare related to the floristic composition of the vegetation. Data on seasonal variation in the diet and changing patterns of resource availability are drawn together within the framework of optimal foraging theory to examine the adaptive strategies underlying the monkeys’ behavior. Their choice of diet was optimal in that they were more selective when profitable food items were common: higher proportions of the diet were given over to fruit and flowers when food availability was high. In parallel with this strategy, a nutritive balance was maintained by consistent inclusion of invertebrates and at least some foliage in the diet, regardless of the amount of fruit or flowers available.     

Fine-Scale Variability in Harbor Seal Foraging Behavior

Understanding the variability of foraging behavior within a population of predators is important for determining their role in the ecosystem and how they may respond to future ecosystem changes. However, such variability has seldom been studied in harbor seals on a fine spatial scale (<30 km). We used a combination of standard and Bayesian generalized linear mixed models to explore how environmental variables influenced the dive behavior of harbor seals. Time-depth recorders were deployed on harbor seals from two haul-out sites in the Salish Sea in 2007 (n = 18) and 2008 (n = 11). Three behavioral bout types were classified from six dive types within each bout; however, one of these bout types was related to haul-out activity and was excluded from analyses. Deep foraging bouts (Type I) were the predominant type used throughout the study; however, variation in the use of bout types was observed relative to haul-out site, season, sex, and light (day/night). The proportional use of Type I and Type II (shallow foraging/traveling) bouts differed dramatically between haul-out sites, seasons, sexes, and whether it was day or night; individual variability between seals also contributed to the observed differences. We hypothesize that this variation in dive behavior was related to habitat or prey specialization by seals from different haul-out sites, or individual variability between seals in the study area. The results highlight the potential influence of habitat and specialization on the foraging behavior of harbor seals, and may help explain the variability in diet that is observed between different haul-out site groups in this population.     

Spring fasting behavior in a marine apex predator provides an index of ecosystem productivity

The effects of declining Arctic sea ice on local ecosystem productivity are not well understood but have been shown to vary inter‐specifically, spatially, and temporally. Because marine mammals occupy upper trophic levels in Arctic food webs, they may be useful indicators for understanding variation in ecosystem productivity. Polar bears (Ursus maritimus) are apex predators that primarily consume benthic and pelagic‐feeding ice‐associated seals. As such, their productivity integrates sea ice conditions and the ecosystem supporting them. Declining sea ice availability has been linked to negative population effects for polar bears but does not fully explain observed population changes. We examined relationships between spring foraging success of polar bears and sea ice conditions, prey productivity, and general patterns of ecosystem productivity in the Beaufort and Chukchi Seas (CSs). Fasting status (≥7 days) was estimated using serum urea and creatinine levels of 1,448 samples collected from 1,177 adult and subadult bears across three subpopulations. Fasting increased in the Beaufort Sea between 1983–1999 and 2000–2016 and was related to an index of ringed seal body condition. This change was concurrent with declines in body condition of polar bears and observed changes in the diet, condition and/or reproduction of four other vertebrate consumers within the food chain. In contrast, fasting declined in CS polar bears between periods and was less common than in the two Beaufort Sea subpopulations consistent with studies demonstrating higher primary productivity and maintenance or improved body condition in polar bears, ringed seals, and bearded seals despite recent sea ice loss in this region. Consistency between regional and temporal variation in spring polar bear fasting and food web productivity suggests that polar bears may be a useful indicator species. Furthermore, our results suggest that spatial and temporal ecological variation is important in affecting upper trophic‐level productivity in these marine ecosystems.     

Trophic level and fatty acids in harp seals compared with common minke whales in the Barents Sea

The objectives of this study were to explore trophic levels and possible diet overlap between harp seals (Pagophilus groenlandicus) and common minke whales (Balaenoptera acutoroostrata) in the Barents Sea using stable isotopes of nitrogen (δ¹⁵N) and carbon (δ¹³C) and fatty acid analyses, and to explore the energy pathways from the plankton to the top predators. Blubber and muscle samples from 93 harp seals and 20 minke whales were collected in the southern Barents Sea in May 2011. The study showed that harp seals were at a higher trophic level than minke whales during spring. This supported previous diet studies suggesting a more fish-dominant diet for seals, as compared with the whales, at this time of the year. The stable isotopes and fatty acids indicated niche separation between the seals and the whales, and between different age groups of the harp seals. Older seals had fatty acid profiles more equal to minke whales as compared with younger seals. Furthermore, while the fatty acid profiles suggested that krill were of particular importance for the young seals, the profiles from older seals and whales suggested that fish dominated their diets.     

Diet composition of polar bears in Svalbard and the western Barents Sea

We estimated both the numerical and biomass composition of the prey of polar bears (Ursus maritimus) from 135 opportunistic observations of kills in Svalbard and the western Barents Sea collected from March to October 1984-2001. By number, the prey composition was dominated by ringed seals (Phoca hispida) (63%), followed by bearded seals (Erignathus barbatus) (13%), harp seals (P. groenlandica) (8%) and unknown species (16%). However, when known prey were converted to biomass, the composition was dominated by bearded seals (55%), followed by ringed seals (30%) and harp seals (15%). Results indicated that bearded seals are an important dietary item for polar bears in the western Barents Sea. We believe that different patterns of space use by different bears may result in geographic variation of diet within the same population.     

Changes in haematological parameters in relation to prey switching in a wild population of harbour seals

1. Previous studies have found marked inter-annual variation in winter food availability, diet composition and body condition in a population of harbour seals ( Phoca vitulina L.) in Northeast Scotland. This study aimed to determine whether there were other physiological consequences of prey switching by comparing haematological parameters in years when the clupeids herring and sprat dominated the diet and in years when seals switched to alternative prey.
2. There were significant differences in leukocyte and erythrocyte parameters in relation to diet composition. In contrast, indices of body condition did not explain the variation in haematological parameters, suggesting that the observed changes did not result from differences in the energetic content of the prey.
3. Leukocyte counts were significantly higher after 'good' clupeid years, although the differences in mean counts were small. Such differences could have resulted either from immuno-suppression, for example because of differences in prey nutrient or contaminant levels, or from differences in the pathogen challenge resulting from geographical variations in water quality.
4. The differences in erythrocyte parameters were more marked, and there was evidence of widespread macrocytic anaemia when seals switched from clupeids to alternative prey. Such differences could result either from acclimation, as a result of prey-specific foraging strategies, or from differences in the nutritional quality of prey.
5. These results indicate that generalist predators such as the harbour seal may exhibit physiological responses to changes in the composition of their diet. These data highlight the need to consider the long-term physiological effects of variations in food availability on the population dynamics of generalist marine top predators. In particular, it is hypothesized that fish-induced anaemia may be responsible for declines in certain pinniped populations.     

Diets of hooded seals (Cystophora cristata) in coastal waters and drift ice waters along the east coast of Greenland

To provide data on the diets of hooded seals (Cystophora cristata) in the Greenland Sea, seals were collected for scientific purposes on expeditions conducted in the pack ice belt east of Greenland in September/October 1999, 2002 and 2003 (autumn), July/August in 2000 (summer), and February/March in 2001 and 2002 (winter). The results from analyses of stomach and intestinal contents from captured seals revealed that their diet was comprised of relatively few prey taxa. The squid Gonatus fabricii and polar cod (Boreogadus saida) were particularly important, whereas capelin (Mallotus villosus) and sand eels (Ammodytes spp.) occasionally contributed more. These four prey items constituted 60-97% of the diet biomass. Gonatus fabricii was the most important food item in autumn and winter, whereas polar cod dominated the summer diet, with important contributions from G. fabricii and sand eels. The latter was only observed on the hooded seal menu during the summer period, whereas polar cod, which was an important component during the autumn survey, was almost absent from the winter samples. During the latter survey, capelin also contributed to the hooded seal diet. Samples obtained from hooded seals in more coastal waters indicated a more varied diet based on fish such as polar cod, redfish (Sebastes sp.) and Greenland halibut (Reinhardtius hippoglossoides).     

Large‐scale molecular barcoding of prey DNA reveals predictors of intrapopulation feeding diversity in a marine predator

Predator–prey interactions are critical in understanding how communities function. However, we need to describe intraspecific variation in diet to accurately depict those interactions. Harbor seals (Phoca vitulina) are an abundant marine predator that prey on species of conservation concern. We estimated intrapopulation feeding diversity (variation in feeding habits between individuals of the same species) of harbor seals in the Salish Sea. Estimates of feeding diversity were examined relative to sex, month, and location using a novel approach that combined molecular techniques, repeated cross‐sectional sampling of scat, and a specialization metric (within‐individual consistency in diet measured by the Proportional Similarity Index ()). Based on 1,083 scat samples collected from five haul‐out sites during four nonsequential years, we quantified diet using metabarcoding techniques and determined the sex of the scat depositor using a molecular assay. Results suggest that intrapopulation feeding diversity was present. Specialization was high over short periods (24–48 hr,  = 0.392, 95% CI = 0.013, R = 100,000) and variable in time and space. Females showed more specialization than males, particularly during summer and fall. Additionally, demersal and benthic prey species were correlated with more specialized diets. The latter finding suggests that this type of prey likely requires specific foraging strategies and that there are trade‐offs between pelagic and benthic foraging styles for harbor seals. This differential feeding on prey species, as well as between sexes of harbor seals, indicates that predator–prey interactions in harbor seals are complex and that each sex may have a different impact on species of conservation concern. As such, describing intrapopulation feeding diversity may unravel hitherto unknown complex predator–prey interactions in the community.     

Changes in diets of individual Baltic ringed seals (Phoca hispida botnica) during their breeding season inferred from stable isotope analysis of multiple tissues

The stable isotope ratios (δ¹³C and δ¹⁵N) of three tissues with different metabolic rates (plasma, liver, and muscle) were used to investigate temporal variation in diet among nine individual Baltic ringed seals (Phoca hispida botnica Gmelin) from the Bothnian Bay, northeast Baltic Sea. The isotope values from plasma should reflect the most recent diet, values from liver the diet of the past weeks prior to sampling, and values from muscle should integrate diet over almost the entire breeding season of the ringed seals. In general, δ¹³C values of liver were more enriched in ¹³C than were those of either muscle or plasma, suggesting that the diet of the seals may have included a higher proportion of ¹³C‐enriched benthic prey in April. Females showed more variable δ¹³C values than males, suggesting possible gender differences in diet or in foraging locations. The differences that were apparent between females possibly reflect individual variation in the onset and duration of parturition and lactation, both of which likely restrict female foraging. Previous data from parasite infections and from alimentary tract contents of the same seals were linked to the isotope data to assist in drawing inferences about changes in the diets of individual seals.     

First documentation of leopard seal predation of South Georgia pintail duck

Leopard seals are regular winter visitors to Bird Island, South Georgia, where they mostly prey on fur seals and penguins, and to a lesser extent on Antarctic krill and fish. Leopard seals can exploit many different species, but there are no records of predation on flying shorebirds in the wild. On 4 October 2008, an individually identified juvenile leopard seal female was observed killing and eating a South Georgia Pintail duck. It also preyed on Antarctic fur seals and gentoo and macaroni penguins during its 2-month temporary residency around the island. The varied diet of this seal exemplifies the generalist prey utilization typical of its species. Long-term diet studies at Bird Island and the published record suggest that predation on ducks is a rather exceptional finding; individual ducks are more likely to escape leopard seal attacks than penguins and provide a far less substantial ration. This note documents the first observation of this species of duck in the diet of leopard seals.     

Analysis of growth and stable isotopes in teeth of male Australian fur seals reveals interannual variability in prey resources

To detect and monitor long‐term ecosystem responses to environmental variability, managers must utilize reliable and quantitative techniques to predict future ecosystem responses. Canine teeth from 67 male Australian fur seals (aged 2–19 yr), collected at Seal Rocks, between 1967 and 1976, were measured for relative growth within the dentine growth layer groups (GLGs), as an index of body growth. Fluctuations in relative growth were apparent during 1956–1971, suggesting interannual variation in prey resources within Bass Strait. These were positively correlated with the Southern Oscillation Index and negatively with the Indian Ocean Subtropical Dipole, both on a 2 yr lag. The observed delay may reflect the time required for the nutrient cascade to filter through to the predominantly benthic prey of Australian fur seals. Stable isotope analysis (δ¹⁵N/δ¹³C) was also used to investigate whether fluctuations in growth were associated with differences in diet. Relative growth was found to be negatively correlated with δ¹⁵N, suggesting years of greater resource availability may be associated with individuals consuming proportionally more prey biomass of lower isotopic value. This study demonstrates that fluctuations in the dentine GLGs of male Australian fur seals are related to environmental parameters, suggesting variation in body growth is mediated by changes in prey resources.     

The influence of diet on foraging habitat models: a case study using nursing Antarctic fur seals

Predictable sources of food underpin lifetime reproductive output in long lived animals. The most important foraging areas of top marine predators are therefore likely to be related to environmental features that enhance productivity in predictable spatial and temporal patterns. Even so, although productive areas within the marine environment are distributed patchily in space and time, most studies assess the relationships between feeding activity and proximate, not long term, environmental characteristics. In addition, individuals within a population may exploit different prey types, and these are often associated with different hydrographic features. Until now, models attempting to associate core foraging areas (CFAs) of marine predators with the environmental characteristics of those areas have not considered the diet of individual animals, despite the influence this could have on these relationships. We used bathymetry and multi‐year (n=24) mean sea surface temperature and variability as predictors of CFAs of lactating Antarctic fur seals Arctocephalus gazella at Heard Island. The effect of prey types on the predictability of these models was explored by matching diet and foraging trip data of individual seals (n=40 seals, n=1 trip each). Differences in diet between seals were mirrored by their spatial behaviour. Foraging strategies differed both between and within groups of seals consuming different diets. Long‐term environmental parameters were useful for predicting the foraging activity of seals that consumed a single prey type with relatively specific habitat preferences, but not for those that consumed single or multiple prey types associated with more varied habitats. Ignoring individual variation in predator diet probably contributes to the poor performance of foraging habitat models. These findings highlight the importance of incorporating individual specialization in foraging behaviour into ecological models and management of predator populations.     

Ecology of the ringed seal (Pusa hispida) from the Bering Strait in the late Holocene

The combined use of analysis of stable isotopes and archeozoological methods with respect to osteological material from an ancient Eskimo settlement (2370-810 yr BP) made it possible to reveal the main distinguishing features of ringed seal in the past. On the basis of the data obtained, we suppose that in the late Holocene in the Bering Strait, there were seals of two different ecotopes: those reproducing on fast ice and those on pack ice differing not only in habitats, but also in the proportion of various food items in the diet. The ringed seals caught by ancient Eskimos during their life had a mixed diet consisting of crustaceans and fish, but in most individuals, a shift towards fish eating was observed. No significant changes in the feeding related to individual age were recorded. The influence of feeding type on body size was shown: bigger, mature individuals occupied higher trophic positions.     

The diet of Antarctic fur sealsArctocephalus gazella during the breeding season at South Georgia

 The diet of lactating female Antarctic fur seals Arctocephalus gazella at South Georgia was investigated during the pup-rearing period (January/March) of 1991–1994. Antarctic krill Euphausia superba was the main prey item, occurring in 88% of all scats (n=497), whereas fish occurred in 47% and squid in 5% of all scats. There was considerable intra- and inter-annual variation in the characteristics of krill taken by fur seals. The distribution of krill sizes taken suggests that fur seals are not actively selecting particular sizes of krill and, therefore, that the krill in the diet reflects the krill available around South Georgia. The absence of group 3 krill (44–48 mm in length) in the South Georgia area, as indicated by their absence in the diet of seals, is suggested as a possible reason for low availability of krill and the subsequent reproductive failure among krill predators. The frequency of occurrence of fish was much higher than in previous studies; the pattern of fish consumption showed a consistent seasonal pattern in 3 of the 4 years studied. Of the total number of the myctophid Protomyctophum choriodon, the most numerous fish taxon, 98% were taken between early February and the middle of March. Champsocephalus gunnari and Lepidonotothen larseni agg., which both feed on krill, dominated the fish component of the diet outside this period and together constituted 94% of the total estimated biomass of fish consumed. The intra- and inter-annual variability in the diet of Antarctic fur seals emphasise the need for diet studies to be conducted during the entire pup-rearing periods of several years. Received: 15 March 1995/Accepted: 13 May 1995     

Harp seal foraging behaviour during summer around Svalbard in the northern Barents Sea: diet composition and the selection of prey

The harp seal Pagophilus groenlandicus is a major high trophic level predator in the Barents Sea, and to better understand their function in the Barents Sea ecosystem, we need to understand their foraging behaviour during their most intensive feeding period. We analysed the diet composition and prey preference of 184 harp seals and 94 faeces samples, sampled in the northern Barents Sea (around Svalbard) during the period May–August in 1996, 1997, and 2004–2006. Concurrent with the sampling of seals, prey availability was assessed in one area in 1996 and 1997 and in two areas in 2006 using standard acoustic methods. Our study showed that harp seal diet composition varied significantly both in time (year) and space, and that their diets appeared to be size dependent. Both subadult (<150 cm) and adult seals were associated with pelagic crustaceans (particularly krill), whereas primarily adult seals were associated with fish (capelin, gadoids and flatfish). Krill was the most important prey group (63 %) followed by polar cod (16 %) and other fish species (10 %). The prey preference of harp seals varied in time and space; polar cod was often preferred by the seals whereas krill was commonly consumed in lower proportion than observed in the survey area. Gadoids and capelin had either been exploited in the same or less proportion as observed in the survey sea. This study emphasises the ecological significance of krill as prime food for harp seals during their intensive feeding period in summer.     

Pre-partum diet of adult female bearded seals in years of contrasting ice conditions

Changing patterns of sea-ice distribution and extent have measurable effects on polar marine systems. Beyond the obvious impacts of key-habitat loss, it is unclear how such changes will influence ice-associated marine mammals in part because of the logistical difficulties of studying foraging behaviour or other aspects of the ecology of large, mobile animals at sea during the polar winter. This study investigated the diet of pregnant bearded seals (Erignathus barbatus) during three spring breeding periods (2005, 2006 and 2007) with markedly contrasting ice conditions in Svalbard using stable isotopes (δ(13)C and δ(15)N) measured in whiskers collected from their newborn pups. The δ(15)N values in the whiskers of individual seals ranged from 11.95 to 17.45 ‰, spanning almost 2 full trophic levels. Some seals were clearly dietary specialists, despite the species being characterised overall as a generalist predator. This may buffer bearded seal populations from the changes in prey distributions lower in the marine food web which seems to accompany continued changes in temperature and ice cover. Comparisons with isotopic signatures of known prey, suggested that benthic gastropods and decapods were the most common prey. Bayesian isotopic mixing models indicated that diet varied considerably among years. In the year with most fast-ice (2005), the seals had the greatest proportion of pelagic fish and lowest benthic invertebrate content, and during the year with the least ice (2006), the seals ate more benthic invertebrates and less pelagic fish. This suggests that the seals fed further offshore in years with greater ice cover, but moved in to the fjords when ice-cover was minimal, giving them access to different types of prey. Long-term trends of sea ice decline, earlier ice melt, and increased water temperatures in the Arctic are likely to have ecosystem-wide effects, including impacts on the forage bases of pagophilic seals.     

Rates of water turnover and energy expenditure of free-living male common seals (Phoca vitulina)

The water and energy metabolism of free-living male common seals ( Phoca vitulina ) during the mating season was investigated using labelled water methods. All three seals, which were captured on two occasions, were in negative energy balance during the study. The daily energy expenditure of one animal, estimated using doubly-labelled water was 52·5 MJ. This is equivalent to six times the basal metabolic rate predicted from Kleiber's (1975) allometric equation. Rates of water turnover were slightly lower than predicted from the allometric equation of Richmond, Langham & Trujillo (1962). The observed rates of water turnover and energy expenditure are considerably higher than those of seals which fast during the mating season, and are consistent with the observed differences in behaviour between males of the common seal and other pinniped males during mating.     

Inter-island variation in the diet of female northern fur seals (Callorhinus ursinus) in the Bering Sea

The diet of adult female northern fur seals ( Callorhinus ursinus ) is examined through the analysis of faecal material collected during the summer breeding season at three breeding locations in the Bering Sea: St. Paul Island (1988, 1990) and St. George Island (1988, 1990) of the Pribilof Islands Group (USA), and Medny Island (1990) of the Commander Islands Group (Russia). Prey consumption varies annually and accordingly with the physical and biological environment surrounding each island. Juvenile walleye pollock ( Theragra chalcogramma ) is the most common prey of northern fur seals from St. Paul Island; the island is surrounded by a broad neritic environment with widely separated frontal zones and is the greatest distance from the continental shelf-edge. Gonatid squid ( Gonatopsis borealis/Berryteuthis magister and Gonatus madokail Gonatus middendorffi ) were the most common prey of northern fur seals from Medny Island; the island is surrounded by a compressed neritic environment and is adjacent to the continental shelf-edge and the oceanic marine environment. A combination of walleye pollock and gonatid squid is consumed by northern fur seals from St. George Island; the island has a surrounding oceanographic environment intermediate between the other two islands.
Variability in predation on walleye pollock is consistent with fishery information concerning the relative abundance and availability of walleye pollock around St. George and St. Paul Islands. The abundance and availability of these prey resources during the summer breeding season are key factors which influence the health and growth of the northern fur seal populations in the Bering Sea.     

Food habits of harbor seals (Phoca vitulina richardii) as an indicator of invasive species in San Francisco Bay,California

The diet of harbor seals (Phoca vitulina richardii) in San Francisco Bay (SFB), California, was determined from July 2007 to July 2008 using prey hard parts recovered from 442 scats collected at five haul‐out sites. Twenty‐two species of fish and one species of crustacean were identified, but harbor seals primarily ate a nonnative invasive species, yellowfin goby (Acanthogobius flavimanus), which increased in dietary importance since the diet was last studied in 1991/1992. Additionally, another nonnative invasive fish species, chameleon goby (Tridentiger trigonocephalus), was found for the first time in the diet of harbor seals in SFB. Harbor seal diet was statistically different between years (1991/1992 and 2007/2008), between the pupping and nonpupping seasons, and between North SFB and South SFB haul‐out locations. The diet of harbor seals was significantly correlated with fish species caught in trawl surveys conducted by the California Department of Fish and Wildlife (CDFW) during the same time periods as this study (2007/2008). Harbor seals currently are influencing the health of the SFB ecosystem in a positive manner by consuming large quantities of nonnative invasive fish species.     

Diet composition and biomass consumption of harbour seals in Telemark and Aust-Agder,Norwegian Skagerrak

ABSTRACT

To explore ecosystem dynamics and functions it is vital to obtain knowledge on predator–prey relations. Harbour seals are piscivorous predators that can come into conflict with fisheries. Recently, as the Skagerrak and Kattegat population of harbour seals has increased, claims have emerged that seals are depleting coastal cod populations. The diet of harbour seals in Norwegian Skagerrak was investigated based on otolith identification from scats. The overall seal diet included 20 different fish species/groups. The most important prey (combined index Q _i ) were haddock/pollack/saithe (32.7%), genus Trisopterus (Norway pout/poor cod/bib, 12.5%), plaice (12.4%) and herring (10.0%). Plaice also had the largest biomass (24.1%). Gadoids and pleuronectids comprised 88.6% of the diet (combined index Q _i ) and 87.1% in terms of biomass. Cod constituted 0.7% (combined index Q _i ) of the overall diet and 2.3% in terms of biomass. Fish length estimates showed that seals generally prefer small fish below minimum allowed landing size. Estimated total amount of fish consumed was 315 tons per year and was dominated by non-commercial species. Annual cod consumption was an estimated 7.1 tons, representing 5% of annual cod landings, suggesting that competition between local fisheries and harbour seals is limited.