DIFFERENCES IN FORAGING LOCATION OF MEXICAN AND CALIFORNIA ELEPHANT SEALS: EVIDENCE FROM STABLE ISOTOPES IN PUPS

Female northern elephant seals, Mirounga angustirostris, from Año Nuevo (AN) in central California feed offshore in mid‐latitude waters (40°–55°N). Migratory patterns and foraging locations of seals from Mexico are unknown. Rookeries on San Benitos (SB) islands in Baja California Sur, Mexico, are ～1,170 km south of AN. Although the colonies are similar in size, seals from SB begin breeding earlier and have an earlier breeding birthing peak than seals from AN. To determine if the foraging location of seals from Mexico was similar to that of seals from California, we measured δ¹³C and δ¹⁵N values in the hair of 48 suckling pups at SB and 37 from AN, assuming that their isotopic signatures reflected those of mothers' milk, their exclusive diet. The mean δ¹³C and δ¹⁵N values for SB pups (?16.1‰± 0.9‰ and 17.7‰± 0.9‰, respectively) were significantly higher than those for AN pups (?17.6‰± 0.4‰ and 15.6‰± 1.0‰, respectively). From data on environmental isotope gradients and known behavior of SB and AN populations, we hypothesize that the isotope differences are due to females in the SB colony foraging ～8° south of seals from AN. This hypothesis can be tested by deployment of satellite tags on adult females from the SB colony.     

Stable isotope composition and parasitic infections of harbor seal young‐of‐the‐year used as prey‐based diet indicators

The stable isotope composition (δ¹³C and δ¹⁵N values) of harbor seals (Phoca vitulina) is influenced by their diet. Young‐of‐the‐year during lactation and postweaning fast are expected be enriched in ¹⁵N compared to foraging seals. We studied the temporal variation of stable isotope composition of young‐of‐the‐year and adults to determine from which point in time the young‐of‐the‐year tissues (i.e., muscles and vibrissae) are influenced by independent foraging only. These results were compared with the development of trophically transmitted parasitic infections. The δ¹⁵N values in young‐of‐the‐year muscles decreased from June (20.3‰ ± 0.5‰) to October (18.5‰ ± 0.4‰), while those of foraging seals were all year long below 19.2‰. This decrease coincides with the increase of parasitic infections in young‐of‐the‐year, reflecting a shift to fish diet. Together these results suggest that the muscles of the young‐of‐the‐year older than 5–6 mo reflect independent foraging and that they can therefore be used in community diet studies. The nursing signal in vibrissae was unclear, as the δ¹⁵N values of young‐of‐the‐year were stable over time, whereas those of adults varied seasonally. However, δ¹⁵N values of nursing pups were significantly higher than those of adults in May and June, maybe due to their reliance on milk.     

Fueling phocids: Divergent exploitation of primary energy sources and parallel ontogenetic diet switches among three species of subarctic seals

Determining how marine predators partition resources is hindered by the difficulty in obtaining information on diet and distribution. Stable isotopes (SI) of carbon (¹³C/¹²C, δ¹³C) and nitrogen (¹⁵N/¹⁴N, δ¹⁵N) provide a two‐dimensional estimate of the dietary space of consumers; an animal's isotopic composition is directly influenced by what they consume and where they feed. Harp (Pagophilus groenlandicus) and hooded (Cystophora cristata) seals are abundant phocid species found in the North Atlantic. We measured and contrasted SI values between seals sampled at nearshore and offshore sites to test for effects of sampling location, sex, age‐class, and body size to gain insight into how these species partition space and prey resources. In addition we contrasted previously published results for gray seals (Halichoerus grypus). Isotope values differed significantly by age class and location in harp and hooded seals. We found significant differences in SI values (mean δ¹³C and δ¹⁵N ± SE) between all species. Hooded seals, a continental shelf‐edge, deep‐diving species, exhibited low SI values (juveniles: ?20.9‰ ± 0.03‰, 13.36‰ ± 0.05‰; adults: ?20.41‰ ± 0.03‰, 14.81‰ ± 0.04‰) characteristic of feeding on meso‐ to bathypelagic prey. Harp seals, which dive to moderate depths primarily on the shelf had intermediate SI values (juveniles: ?20.53‰ ± 0.01‰, 13.91‰ ± 0.01‰; adults: ?20.13‰ ± 0.01‰, 14.96‰ ± 0.01‰) characteristic of feeding on epipelagic prey, whereas gray seals, which feed on or near the sea floor in shallow shelf waters, had high SI values (juveniles: ?19.74‰ ± 0.04‰, 17.51‰ ± 0.05‰; adults: ?18.86‰ ± 0.01‰, 17.23‰ ± 0.02‰) characteristic of feeding on demersal prey. In all species, δ¹³C values increased with body size and age in the same manner, indicating that seals exploit or forage in deeper habitats as they get larger and older. We hypothesize that the consistent ontogenetic shift in foraging niche, despite large differences between species in their diving behavior, geographic range and habitat use, not only reflects increased access to different prey due to increased diving capacity, but a progressive adjustment to balance energy budgets by reducing foraging costs.     

Sex-specific foraging strategies and resource partitioning in the southern elephant seal (Mirounga leonina)

The evolution of resource specializations is poorly understood, especially in marine systems. The southern elephant seal (Mirounga leonina) is the largest of the phocid seals, sexually dimorphic, and thought to prey predominantly on fish and squid. We collected vibrissae from male and female southern elephant seals, and assessed stable C and N isotope ratios along the length of the vibrissae. Given that whiskers grow slowly, this sampling strategy reflects any variation in feeding behaviour over a period of time. We found that isotopic variation among females was relatively small, and that the apparent prey choice and trophic level of females was different from that for males. Further, males showed a very broad range of trophic/prey choice positions, grouped into several clusters, and this included isotopic values too low to match a broad range of potential fish and cephalopod prey tested. One of these clusters overlapped with data for South American sea lions (Otaria flavescens), which were measured for comparison. Both male southern elephant seals and southern sea lions forage over the continental shelf, providing the potential for competition. We discuss the possibility that individual southern elephant seals are pursuing specialist foraging strategies to avoid competition, both with one another, and with the South American sea lions that breed nearby.     

Whisker isotopic signature depicts migration patterns and multi-year intra- and inter-individual foraging strategies in fur seals

The movement and dietary history of individuals can be studied using stable isotope records in archival keratinous tissues. Here, we present a chronology of temporally fine-scale data on the trophic niche of otariid seals by measuring the isotopic signature of serially sampled whiskers. Whiskers of male Antarctic fur seals breeding at the Crozet Islands showed synchronous and regular oscillations in both their δ¹³C and δ¹⁵N values that are likely to represent their annual migrations over the long term (mean 4.8 years). At the population level, male Antarctic fur seals showed substantial variation in both δ¹³C and δ¹⁵N values, occupying nearly all the ‘isotopic space’ created by the diversity of potential oceanic habitats (from high Antarctica to the subtropics) and prey (from Antarctic krill to subantarctic and subtropical mesopelagic fishes). At the individual level, whisker isotopic signatures depict a large diversity of foraging strategies. Some seals remained in either subantarctic or Antarctic waters, while the migratory cycle of most animals encompassed a wide latitudinal gradient where they fed on different prey. The isotopic signature of whiskers, therefore, revealed new multi-year foraging strategies of male Antarctic fur seals and is a powerful tool for investigating the ecological niche during cryptic stages of mammals'' life.     

Stable isotopes document seasonal changes in trophic niches and winter foraging individual specialization in diving predators from the Southern Ocean

1. Climatic variation outside the breeding season affects fluctuations in population numbers of seabirds and marine mammals. A challenge in identifying the underlying biological mechanisms is the lack of information on their foraging strategies during winter, when individuals migrate far from their breeding grounds. 2. We investigated the temporal variability in resource partitioning within the guild of five sympatric Subantarctic penguins and fur seals from Crozet Islands. The stable isotopic ratios of carbon (delta(13)C) and nitrogen (delta(15)N) for whole blood were measured for penguins and fur seals, as were the isotopic ratios for penguin nails and food. Animals were sampled at two periods, during breeding in summer and at their arrival in the colonies in spring (hereafter winter, since the temporal integration of blood amounting to several months). 3. In summer, delta(13)C and delta(15)N for blood samples defined three foraging areas and two trophic levels, respectively, characterizing four nonoverlapping trophic niches. King penguins and female Antarctic and Subantarctic fur seals are myctophid eaters foraging in distinct water masses, while both macaroni and rockhopper penguins had identical isotopic signatures indicating feeding on crustaceans near the archipelago. 4. Isotopic ratios were almost identical in summer and winter suggesting no major changes in the species niches, and hence, in the trophic structure of the guild during the nonbreeding period. A seasonal difference, however, was the larger variances in delta(13)C (and also to a lesser extent in delta(15)N) values in winter, thus verifying our hypothesis that trophic niches widen when individuals are no longer central place foragers. 5. Winter isotopic ratios of macaroni penguins and male Antarctic fur seals had large variances, indicating individual foraging specializations. The range of delta(13)C and delta(15)N values of male fur seals showed, respectively, that they dispersed over a wide latitudinal gradient (from Antarctica to north of the archipelago) and fed on different prey (crustaceans and fish). 6. By comparing summer and winter isotopic ratios and examining the summer diet, we highlight the feeding habits of marine predators that were not previously addressed. The findings have a number of implications for understanding the functioning of the pelagic ecosystem and on the demography of these species.     

Detecting alternate foraging ecotypes in Australian sea lion (Neophoca cinerea) colonies using stable isotope analysis

Stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes are used frequently to describe the trophic ecology of top marine mammal predators. Australian sea lions (Neophoca cinerea) are one of the world's rarest otariid seals and exhibit the highest levels of natal site philopatry of any seal. We report the development of a screening technique to identify different foraging ecotypes and assess their relative frequencies in Australian sea lion breeding colonies using stable isotope ratios in pups. Geospatial and dive data from 15 adult females at three breeding colonies revealed alternate foraging strategies (inshore and offshore foraging) that were reflected in significant changes in δ¹³C and δ¹⁵N. Isotope fractionation from mother to pup was validated using paired whisker and blood serum samples with no significant difference between δ¹³C and δ¹⁵N enrichment of +1.27‰ (whiskers) and +1.92‰ (blood serum) from mothers to pups. Isotope ratios from whisker samples representing over 50% of pups born at three colonies revealed significant intercolony differences in maternal foraging ecotype frequencies. These results are unique in that ecological partitioning over such a small spatial scale has not been described in any other otariid species.     

Is individual consistency in body mass and reproductive decisions linked to individual specialization in foraging behavior in a long‐lived seabird?

Individual specialization in diet or foraging behavior within apparently generalist populations has been described for many species, especially in polar and temperate marine environments, where resource distribution is relatively predictable. It is unclear, however, whether and how increased environmental variability – and thus reduced predictability of resources – due to global climate change will affect individual specialization. We determined the within‐ and among‐individual components of the trophic niche and the within‐individual repeatability of δ¹³C and δ¹⁵N in feathers and red blood cells of individual female southern rockhopper penguins (Eudyptes chrysocome) across 7 years. We also investigated the effect of environmental variables (Southern Annular Mode, Southern Oscillation Index, and local sea surface temperature anomaly) on the isotopic values, as well as the link between stable isotopes and female body mass, clutch initiation dates, and total clutch mass. We observed consistent red blood cell δ¹³C and δ¹⁵N values within individuals among years, suggesting a moderate degree of within‐individual specialization in C and N during the prebreeding period. However, the total niche width was reduced and individual specialization not present during the premolt period. Despite significant interannual differences in isotope values of C and N and environmental conditions, none of the environmental variables were linked to stable isotope values and thus able to explain phenotypic plasticity. Furthermore, neither the within‐individual nor among‐individual effects of stable isotopes were found to be related to female body mass, clutch initiation date, or total clutch mass. In conclusion, our results emphasize that the degree of specialization within generalist populations can vary over the course of 1 year, even when being consistent within the same season across years. We were unable to confirm that environmental variability counteracts individual specialization in foraging behavior, as phenotypic plasticity in δ¹³C and δ¹⁵N was not linked to any of the environmental variables studied.     

Foraging segregation by sex and age class in the Guadalupe fur seal from Guadalupe Island,Mexico

A proper assessment of the foraging habits of the Guadalupe fur seal (GFS; Arctocephalus townsendi) is a priority to better understand its recovery, in which the potential for intraspecific competition for prey and space resources is expected to lead to segregation. This study aimed to determine the foraging habits of different sex and age classes. A total of 146 GFS fur samples was collected at Guadalupe Island, Mexico (2014–2020) for stable isotopes (δ¹³C and δ¹⁵N) analysis. Isotopic areas were created (SIBER package in R). Significant isotopic differences were observed between classes. Male (3.6‰²) and female (3.0‰²) juveniles had the largest isotopic areas due to a greater foraging dispersion. Adult females showed the lowest mean δ¹⁵N value (16.1‰ ± 0.5‰) due to foraging trips that are mostly performed towards high latitudes. Except for pups, adult and subadult males presented the highest mean δ¹⁵N (17.4‰ ± 0.4‰) and δ¹³C values (−17.0‰ ± 0.8‰) due to a possible higher trophic level and coastal foraging habits, whereas pups presented the highest mean δ¹⁵N value (17.6‰ ± 0.3‰) because of lactation, which reflects their mothers δ¹⁵N signal plus their own enrichment. Our findings suggest a segregation explained by differences in life history, energy requirements, and a possible strategy to avoid competition.     

INVESTIGATING TROPHIC RELATIONSHIPS OF PINNIPEDS IN ALASKA AND WASHINGTON USING STABLE ISOTOPE RATIOS OF NITROGEN AND CARBON

We measured stable-nitrogen (δ¹⁵N) and stable-carbon (δ¹³C) isotope ratios in muscle and hair from 7 northern fur seals (Callorhinus ursinus) from the Pribilof Islands, Alaska, and 27 Steller sea lions (Eumetopias jubatus), and 14 harbor seals (Phoca vitulina) from the Gulf of Alaska and coast of Washington State, in order to contrast dietary information derived from isotopic vs. available conventional dietary studies. Stable-nitrogen-isotope analysis of muscle revealed that harbor seals were enriched over sea lions (mean δ¹⁵N = 18.6‰vs. 17.5‰) which were in turn enriched over northern fur seals (mean δ¹⁵N = 16.6‰). Trophic segregation among these species likely results primarily from differential reliance on herring (Clupea harengus), Atka mackerel (Pleurogrammus monopterygius), and large vs. small walleye pollock (Theregra chalcogramma). According to their δ¹⁵N values, adult male Steller sea lions showed a higher trophic position than adult females (mean δ¹⁵N: 18.0‰vs. 17.2‰), whereas adult female northern fur seals were trophically higher than juvenile male fur seals (mean δ¹⁵N: 16.5‰vs. 15.0‰). Each of these observed differences likely resulted from differential reliance on squid or differences in the size range of pollock consumed. Three northern fur seal pups showed higher δ¹⁵N enrichment over adults (mean 17.7‰vs. 15.8‰) due to their reliance on their mother's milk. Stable-carbon isotope measurements of hair revealed a cline toward more negative values with latitude. Segregation in hair δ¹³C between Steller sea lions and harbor seals off the coast of Washington (mean δ¹³C: ?13.6‰vs.?15.0‰) reflected the greater association of harbor seals with freshwater input from the Columbia River. Our study demonstrates the utility of the stable isotope approach to augment conventional dietary analyses of pinnipeds and other marine mammals.     

Stable isotope series from elephant ivory reveal lifetime histories of a true dietary generalist

Longitudinal studies have revealed how variation in resource use within consumer populations can impact their dynamics and functional significance in communities. Here, we investigate multi-decadal diet variations within individuals of a keystone megaherbivore species, the African elephant (Loxodonta africana), using serial stable isotope analysis of tusks from the Kruger National Park, South Africa. These records, representing the longest continuous diet histories documented for any extant species, reveal extensive seasonal and annual variations in isotopic--and hence dietary--niches of individuals, but little variation between them. Lack of niche distinction across individuals contrasts several recent studies, which found relatively high levels of individual niche specialization in various taxa. Our result is consistent with theory that individual mammal herbivores are nutritionally constrained to maintain broad diet niches. Individual diet specialization would also be a costly strategy for large-bodied taxa foraging over wide areas in spatio-temporally heterogeneous environments. High levels of within-individual diet variability occurred within and across seasons, and persisted despite an overall increase in inferred C(4) grass consumption through the twentieth century. We suggest that switching between C(3) browsing and C(4) grazing over extended time scales facilitates elephant survival through environmental change, and could even allow recovery of overused resources.     

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.     

Trophic niche overlap between sympatric harbour seals (Phoca vitulina) and grey seals (Halichoerus grypus) at the southern limit of their European range (Eastern English Channel)

Sympatric harbour (Phoca vitulina) and grey seals (Halichoerus grypus) are increasingly considered potential competitors, especially since recent local declines in harbour seal numbers while grey seal numbers remained stable or increased at their European core distributions. A better understanding of the interactions between these species is critical for conservation efforts. This study aimed to identify the trophic niche overlap between harbour and grey seals at the southern limit of their European range, in the Baie de Somme (BDS, Eastern English Channel, France), where numbers of resident harbour seals and visiting grey seals are increasing exponentially. Dietary overlap was identified from scat contents using hierarchical clustering. Isotopic niche overlap was quantified using δ¹³C and δ¹⁵N isotopic values from whiskers of 18 individuals, by estimating isotopic standard ellipses with a novel hierarchical model developed in a Bayesian framework to consider both intraindividual variability and interindividual variability. Foraging areas of these individuals were identified from telemetry data. The three independent approaches provided converging results, revealing a high trophic niche overlap due to consumption of benthic flatfish. Two diet clusters were dominated by either small or large benthic flatfish; these comprised 85.5% [CI95%: 80.3%–90.2%] of harbour seal scats and 46.8% [35.1%–58.4%] of grey seal scats. The narrower isotopic niche of harbour seals was nested within that of grey seals (58.2% [22.7%–100%] overlap). Grey seals with isotopic values similar to harbour seals foraged in coastal waters close to the BDS alike harbour seals did, suggesting the niche overlap may be due to individual grey seal strategies. Our findings therefore provide the basis for potential competition between both species (foraging on benthic flatfish close to the BDS). We suggest that a continued increase in seal numbers and/or a decrease in flatfish supply in this area could cause/amplify competitive interactions and have deleterious effects on harbour seal colonies.     

Long-term species, sexual and individual variations in foraging strategies of fur seals revealed by stable isotopes in whiskers

Background

Individual variations in the use of the species niche are an important component of diversity in trophic interactions. A challenge in testing consistency of individual foraging strategy is the repeated collection of information on the same individuals.

Methodology/Principal Findings

The foraging strategies of sympatric fur seals (Arctocephalus gazella and A. tropicalis) were examined using the stable isotope signature of serially sampled whiskers. Most whiskers exhibited synchronous δ¹³C and δ¹⁵N oscillations that correspond to the seal annual movements over the long term (up to 8 years). δ¹³C and δ¹⁵N values were spread over large ranges, with differences between species, sexes and individuals. The main segregating mechanism operates at the spatial scale. Most seals favored foraging in subantarctic waters (where the Crozet Islands are located) where they fed on myctophids. However, A. gazella dispersed in the Antarctic Zone and A. tropicalis more in the subtropics. Gender differences in annual time budget shape the seal movements. Males that do not perform any parental care exhibited large isotopic oscillations reflecting broad annual migrations, while isotopic values of females confined to a limited foraging range during lactation exhibited smaller changes. Limited inter-individual isotopic variations occurred in female seals and in male A. tropicalis. In contrast, male A. gazella showed large inter-individual variations, with some males migrating repeatedly to high-Antarctic waters where they fed on krill, thus meaning that individual specialization occurred over years.

Conclusions/Significance

Whisker isotopic signature yields unique long-term information on individual behaviour that integrates the spatial, trophic and temporal dimensions of the ecological niche. The method allows depicting the entire realized niche of the species, including some of its less well-known components such as age-, sex-, individual- and migration-related changes. It highlights intrapopulation heterogeneity in foraging strategies that could have important implications for likely demographic responses to environmental variability.     

Stable isotope records of sei whale baleens from Chilean Patagonia as archives for feeding and migration behavior

Carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope variations in baleen plates of sei whales (Balaenoptera borealis) stranded after a mass mortality event in Chilean Patagonia were investigated to assess potential dietary and migratory patterns. Carbon and nitrogen isotope ratios of seven baleens from six individuals were analyzed. The δ¹³C values ranged from ? 19.1 to ? 15.9‰ and the δ¹⁵N values from 8.7 to 15.4‰. Variations of up to 2.9‰ for δ¹³C and 5.3‰ for δ¹⁵N were observed within one baleen. Carbon and nitrogen isotope records of each baleen were significantly correlated and showed recurring oscillations confirmed by wavelet analyses. Oscillations slightly differed in periodicity indicating variable baleen growth rates between 10.0 and 16.5 cm/year. Food sources of the whales are discussed in terms of available isotope data for potential prey taxa and potential migratory behavior on the basis of latitudinal isotope gradients of particulate organic matter. Cyclicity could be explained by regular migrations of the sei whales from subtropical calving areas to high‐latitude foraging grounds. δ¹⁵N records of baleens differed between individuals eventually pointing to diverse feeding and migratory preferences among sei whale individuals.     

Specialized foraging habits of adult female California sea lions Zalophus californianus

Individual foraging specialization occurs when organisms use a small subset of the resources available to a population. This plays an important role in population dynamics since individuals may have different ecological functions within an ecosystem related to habitat use and prey preferences. The foraging habitat fidelity and degree of specialization of California sea lions (Zalophus californianus) were evaluated by analyzing the stable isotopes values of carbon and nitrogen in vibrissae collected from 16 adult females from the reproductive colony on Santa Margarita Island, Magdalena Bay, Mexico, in 2012 and 2013. Based on the degree of individual specialization in δ¹⁵N, 62.5% of the females assessed can be considered specialist consumers focusing on the same prey or different prey from the same trophic level. The degree of individual specialization in δ¹³C indicated that 100% of the individuals showed fidelity to their foraging habitat as some fed in the lagoon, others foraged along the coast, and a third group preferred prey from the pelagic environment during both the breeding and nonbreeding seasons, suggesting diversification of foraging areas. Foraging area fidelity persisted despite the 2°C increase in the sea surface temperature over the course of the study period.     

Temporal variability in stable isotope ratios of C and N in the vibrissa of captive and wild adult South American sea lions Otaria byronia: More than just diet shifts

We analyzed the δ¹³C and δ¹⁵N values in the vibrissae of captive adult breeding South American sea lions (Otaria byronia) fed at a constant diet and then used this information to analyze the change in stable isotope values along the vibrissae from wild individuals. The overall diet‐to‐vibrissa discrimination factor of the captive animals was 3.0‰ ± 0.1‰ for δ¹³C and 3.6‰ ± 0.1‰ for δ¹⁵N, but the stable isotope ratios fluctuated periodically despite constant diet. The δ¹³C and δ¹⁵N values of the captive male declined at the end of the breeding season, whereas the δ¹³C values of the female increased during the central part of pregnancy and the δ¹⁵N values peaked during lactation. The δ¹³C and δ¹⁵N values of adult wild specimens also fluctuated periodically and vibrissae growth rate (0.15 mm/d in both sexes) was slightly lower than in captivity (0.17 mm/d), assuming an annual periodicity for oscillations. Similarities in the amplitude of the cycles of captive and wild males suggested that fasting was probably the main source of periodic variability in the δ¹⁵N of wild males, whereas pregnancy and lactation were probably the main source of periodic variability for the δ¹³C of wild females.     

Stable carbon and nitrogen isotopes in multiple tissues of wild and captive harbor seals (Phoca vitulina) off the California coast

Stable carbon and nitrogen isotope ratios (δ¹³C and δ¹⁵N) of serum, red blood cells (RBC), muscle, and blubber were measured in captive and wild northeast Pacific harbor seals (Phoca vitulina richardii) at three coastal California sites (San Francisco Bay, Tomales Bay, and Channel Islands). Trophic discrimination factors (Δ_{Tissue‐Diet}) were calculated for captive seals and then applied in wild counterparts in each habitat to estimate trophic position and feeding behavior. Trophic discrimination factors for δ¹⁵N of serum (+3.8‰), lipid‐extracted muscle (+1.6‰), and lipid‐blubber (+6.5‰) are proposed to determine trophic position. An offset between RBC and serum of +0.3‰ for δ¹³C and ?0.6‰ for δ¹⁵N was observed, which is consistent with previous research. Specifically, weaner seals (<1 yr) had large offsets, suggesting strong trophic position shifts during this life stage. Isotopic values indicated an average trophic position of 3.6 at both San Francisco Bay and Tomales Bay and 4.2 at Channel Islands. Isotopic means were strongly dependent on age class and also suggested that mean diet composition varies considerably between all locations. Together, these data indicate that isotopic composition of blood fractions can be an effective approach to estimate trophic position and dietary behavior in wild pinnipeds.     

中国古代儿童断奶模式与喂养方式初探——以安徽薄阳城遗址人骨的C、N稳定同位素分析为例

了解古代儿童的断奶模式和喂养方式,可望为揭示古代儿童的营养状况、生长发育以及其死亡原因提供有价值的重要信息。近些年来,通过对古代儿童骨骼的C、N稳定同位素分析,重建古代儿童的断奶模式和喂养方式,成为国际生物考古界的研究前沿,但在我国尚无任何相关报道。为此,本文以安徽滁州薄阳城遗址人骨(包含不同年龄阶段的儿童和成年人)样品为研究材料,通过对同一个体肋骨和肢骨的C、N稳定同位素分析和比较,重点探讨了我国西周时期儿童的断奶模式及喂养方式。研究结果显示:儿童在2岁已有辅食的摄入,大多数儿童在3-4岁已经完成断奶;断奶后的儿童,其食物中包含更多的植物作为辅食。本研究,印证了我国历史文献中有关儿童喂养模式的最早记载(唐代孙思邈的医书《千金方》),也为系统探索我国古代儿童的断奶模式和喂养方式提供了很好的范例。     

Individual dietary specialization in a generalist predator: A stable isotope analysis of urban and rural red foxes

Some carnivores are known to survive well in urban habitats, yet the underlying behavioral tactics are poorly understood. One likely explanation for the success in urban habitats might be that carnivores are generalist consumers. However, urban populations of carnivores could as well consist of specialist feeders. Here, we compared the isotopic specialization of red foxes in urban and rural environments, using both a population and an individual level perspective. We measured stable isotope ratios in increments of red fox whiskers and potential food sources. Our results reveal that red foxes have a broad isotopic dietary niche and a large variation in resource use. Despite this large variation, we found significant differences between the variance of the urban and rural population for δ¹³C as well as δ¹⁵N values, suggesting a habitat‐specific foraging behavior. Although urban regions are more heterogeneous regarding land cover (based on the Shannon index) than rural regions, the dietary range of urban foxes was smaller compared with that of rural conspecifics. Moreover, the higher δ¹³C values and lower δ¹⁵N values of urban foxes suggest a relatively high input of anthropogenic food sources. The diet of most individuals remained largely constant over a longer period. The low intraindividual variability of urban and rural red foxes suggests a relatively constant proportion of food items consumed by individuals. Urban and rural foxes utilized a small proportion of the potentially available isotopic dietary niche as indicated by the low within‐individual variation compared to the between‐individual variation. We conclude that generalist fox populations consist of individual food specialists in urban and rural populations at least over those periods covered by our study.