Merav Ben-David

Consumption of salmon by Alaskan brown bears: a trade-off between nutritional requirements and the risk of infanticide?

The risk of infanticide may alter foraging decisions made by females, which otherwise would have been based on nutritional requirements and forage quality and availability. In systems where meat resources are spatially aggregated in late summer and fall, female brown bears ( Ursus arctos) would be faced with a trade-off situation. The need of reproductive females to accumulate adequate fat stores would likely result in a decision to frequent salmon streams and consume the protein- and lipid-rich spawning salmon. In contrast, aggregations of bears along salmon streams would create conditions of high risk of infanticide. We investigated consumption of salmon by brown bears on Admiralty and Chichagof Islands in Southeast Alaska from 1982 to 2000 using stable isotope analysis and radiotelemetry. While nearly all males (22 of 23) consumed relatively large amounts of salmon (i.e., >10% relative contribution to seasonal diet), not all females ( n =56) did so. Five of 26 females for which we had reproductive data, occupied home ranges that had no access to salmon and thus did not consume salmon when they were mated or accompanied by young. Of females that had access to salmon streams ( n =21), all mated individuals ( n =16) had δ15N values indicative of salmon consumption. In contrast, 4 out of 16 females with cubs avoided consuming salmon altogether, and of the other 12, 3 consumed less salmon than they did when they were mated. For 11 of 21 females with access to salmon streams we had data encompassing both reproductive states. Five of those altered foraging strategies and exhibited significantly lower values of δ15N when accompanied by young than when mated, while 6 did not. Radiotelemetry data indicated that females with spring cubs were found, on average, further away from streams during the spawning season compared with females with no young, but both did not differ from males and females with yearlings and 2-year-olds. Females with young that avoided salmon streams were significantly lighter indicating that female choice to avoid consumption of salmon carries a cost that may translate to lower female or cub survivorship. The role of the social hierarchy of males and females, mating history, and paternity in affecting the risk of infanticide and foraging decisions of female brown bears merit further investigation.

Stable isotopes in mammalian research: a beginner's guide

Abstract We open this Special Feature on stable isotopes in mammalian research with a beginners guide, an introduction to the novice and a refresher to the well-versed. In this guide we provide the background needed to understand the more advanced papers that follow. We describe the basic principles of isotopic fractionation and discrimination, briefly explain the processes that govern isotopic incorporation into animal tissues, list some innovative studies, and provide cautionary notes and caveats. In addition to discussing the uses of natural abundance we present the concepts and applications of enriched isotopes and the potential combination of these 2 methodologies. We end with descriptions of analytical and conceptual developments that we believe will be cardinal to the future of isotopic analyses in mammalian research.

Niche partitioning among mule deer, elk, and cattle: Do stable isotopes reflect dietary niche?

Abstract We examined dietary niches of mule deer (Odocoileus hemionus), North American elk (Cervus elaphus), and free-ranging cattle (Bos taurus) that frequently co-occur in western North America. We tested the hypothesis that those three species would exhibit little overlap in diet and that mule deer, the smallest in body size of the three species, would forage more selectively than either elk or cattle. We determined diet composition from microhistological analysis and used principal components analysis to assess dietary niches. In addition to those conventional methods, we also assessed whether dietary overlap among those three ruminants would be reflected in stable isotope ratios (δ13C and δ15N) from fecal pellets. Principal component 1 represented a foraging axis based on plant classes, whereas principal component 2 represented a continuum from grazing to browsing, which revealed complete separation among those three large herbivores. Similarly, δ13C and δ15N differed significantly among species and indicated differences in moisture regimes within habitats and types of forages used by those three ruminants. Mule deer had the greatest variability in diet and foraged on more xeric forages than did either elk or cattle. Stable isotopes elucidated differences in dietary niche among the three ruminants that were not evident from dietary analysis alone.

Characteristics of sex-biased dispersal and gene flow in coastal river otters: implications for natural recolonization of extirpated populations

River otters (Lontra canadensis) were extirpated from much of their historic distribution because of exposure to pollution and urbanization, resulting in expansive reintroduction programmes that continue today for this and other species of otters worldwide. Bioaccumulation of toxins negatively affects fecundity among mustelids, but high vagility and different dispersal distances between genders may permit otter populations to recover from extirpation caused by localized environmental pollution. Without understanding the influence of factors such as social structure and sex‐biased dispersal on genetic variation and gene flow among populations, effects of local extirpation and the potential for natural recolonization (i.e. the need for translocations) cannot be assessed. We studied gene flow among seven study areas for river otters (n = 110 otters) inhabiting marine environments in Prince William Sound, Alaska, USA. Using nine DNA microsatellite markers and assignment tests, we calculated immigration rates and dispersal distances and tested for isolation by distance. In addition, we radiotracked 55 individuals in three areas to determine characteristics of dispersal. Gender differences in sociality and spatial relationships resulted in different dispersal distances. Male river otters had greater gene flow among close populations (within 16–30 km) mostly via breeding dispersal, but both genders exhibited an equal, low probability of natal dispersal; and some females dispersed 60–90 km. These data, obtained in a coastal environment without anthropogenic barriers to dispersal (e.g. habitat fragmentation or urbanization), may serve as baseline data for predicting dispersal under optimal conditions. Our data may indicate that natural recolonization of coastal river otters following local extirpation could be a slow process because of low dispersal among females, and recolonization may be substantially delayed unless viable populations occurred nearby. Because of significant isolation by distance for male otters and low gene flow for females, translocations should be undertaken with caution to help preserve genetic diversity in this species.

Analysis of stable isotope data : A K nearest-neighbors randomization test

The use of stable isotope analysis in ecological and wildlife studies is rapidly increasing. Studies include evaluating flow of nutrients in ecosystems and studying dietary composition of individual animals. Several mixing models have been developed to evaluate the relative contribution of different foods to the diet of consumers. All these mixing models require that all prey types will be significantly different in bivariate space. This requirement usually poses a problem in analyzing data of stable isotope ratios because sample sizes in most studies are small and seldom normally distributed. We propose a randomization test that we based on the K nearest-neighbor approach. Results from our simulations of power revealed that the K nearest-neighbor test appears to have high power even with small sample sizes and comparatively low displacement. The K nearest-neighbor test described here provides the preliminary statistical analysis necessary for the use of the mixing models, and therefore is a new, powerful tool for analyzing stable isotope data. In evaluating the test performance on data collected from American martens (Martes americana) and their prey on Chichagof Island, Southeast Alaska, we were able to reject our null hypothesis that all samples of prey were drawn from identical populations (P = 0.05). A program written in Pascal or S-Plus is available from the authors to evaluate the K nearest-neighbor statistic for several groups.

Why do river otters scent-mark? An experimental test of several hypotheses

We tested several alternative hypotheses about the function of scent marking by the North American river otter, Lontra canadensis. Otters may mark at latrine sites with spraints ( faeces) to (1) signal species identity, (2) advertise their reproductive status, (3) establish and maintain territories, and (4) communicate social status and identity to group members. Olfactory preference tests were conducted at the Alaska Sealife Center in Seward, Alaska, on a group of 15 wild-caught male otters in February 1999. We found that male otters investigated otter scent more than sealion faeces. The male otters also showed a preference for male scent over the scent of anoestrous females. No preference for the scent of unfamiliar males, compared with the scent of familiar males, was observed, and no preference for the scent of close kin was detected. However, an investigation of dominant relationships of the captive otters showed that dominant males spent more time investigating male scent than did subordinate males. Thus, spraints deposited at latrine sites may function to communicate social status of males.

Communication in river otters: Creation of variable resource sheds for terrestrial communities

Movements and behavior of animals can result in transfer of nutrients between discrete spatial patches, leading to spatial and temporal variability in resource sheds, modification of nutrient cycling, changes in productivity and in community structure and function, and increases in landscape heterogeneity. In this study, we explored the function of scent-marking at latrines by coastal river otters (Lontra canadensis), through investigating spatial distributions of otters with respect to gender, sociality, and the distribution of their food resources. We then calculated the amounts of nitrogen (N) and phosphorus (P) transported to latrine sites based on otter foraging behavior and the function of scent-marking at latrines. Locations of 55 radio-tagged otters in Prince William Sound, Alaska, USA, were obtained through aerial telemetry over a period of four years. Data on fish densities and marine habitat features were concurrently obtained from scuba transects and aerial surveys. A plastic social organizati...

Assigning birds to wintering and breeding grounds using stable isotopes: lessons from two feather generations among three intercontinental migrants

AbstractGeographic origins of populations and migration patterns of several vertebrate and invertebrate species have been inferred from geographically distinct isotopes in their tissues. To test the hypothesis that feathers grown on different continents would reflect continental differences of δD in precipitation and have significantly different stable isotope ratios, we analyzed stable isotopes in two generations of feathers from three bird species (American and Pacific golden-plovers, Pluvialis dominica and P. fulva, and northern wheatears Oenantheoenanthe) that breed in North America and winter in South America, the South Pacific and Asia, and Africa. We found significant differences in stable isotope signatures between summer- and winter-grown feathers in the plovers, and our use of two generations of feathers provided similar variation to that reported in studies using larger sample sizes. In contrast to plovers, no differences were detected in isotope values between summer- and winter-grown feathers in wheatears. Discriminant analyses separated 80% of summer- and winter-grown feathers for each plover species. Nonetheless, an “assignment with exclusion” method adapted from population genetics to impart a measure of confidence in assigning individuals to groups of origin resulted in an overall accuracy among plovers of only 41%, compared with a 63% assignment accuracy when the exclusion criterion was removed. Thus, we were unable to accurately assign feathers to origin of growth on the continental scale. Moreover, using δD expectations for North America, we were unable to assign summer-grown plover feathers to within better than several thousand kilometers of their true origins. We urge researchers to carefully consider the ecology and physiology of their study organisms, statistical methodology, and the interpretation of results when using stable isotopes to infer the geographic origins of feather growth.

Maternal condition determines birth date and growth of newborn bear cubs

Abstract The number, size, and survival of bear cubs emerging from winter dens depend on maternal condition prior to entering the den. We hypothesized that delayed implantation provides flexibility in timing of birth such that pregnant females are able to track environmental or body conditions long after conception to optimize reproductive output in a changing environment. We tested the hypotheses that causative links between maternal condition and size of newly emerging brown bear (Ursus arctos) cubs were females in superior condition give birth earlier and, thereby, lactate longer in the den than females in poorer condition; and females in superior condition produce more milk or higher quality milk, which accelerates cub growth relative to females in poorer condition. No brown bear with a body fat content ≤ 20% produced cubs even though breeding occurred. Brown bears that were fat gave birth earlier than those that were lean. Cubs nursing from fat mothers grew faster than those nursing from lean mothers. The combination of an earlier birth date and faster growth by cubs produced from fat mothers increased mass of brown bear and polar bear (U. maritimus) twins at den emergence by 330–360 g for each unit increase in percent maternal body fat content when entering hibernation.

Controlling for anthropogenically induced atmospheric variation in stable carbon isotope studies

Increased use of stable isotope analysis to examine food-web dynamics, migration, transfer of nutrients, and behavior will likely result in expansion of stable isotope studies investigating human-induced global changes. Recent elevation of atmospheric CO2 concentration, related primarily to fossil fuel combustion, has reduced atmospheric CO2 δ13C (13C/12C), and this change in isotopic baseline has, in turn, reduced plant and animal tissue δ13C of terrestrial and aquatic organisms. Such depletion in CO2 δ13C and its effects on tissue δ13C may introduce bias into δ13C investigations, and if this variation is not controlled, may confound interpretation of results obtained from tissue samples collected over a temporal span. To control for this source of variation, we used a high-precision record of atmospheric CO2 δ13C from ice cores and direct atmospheric measurements to model modern change in CO2 δ13C. From this model, we estimated a correction factor that controls for atmospheric change; this correction reduces bias associated with changes in atmospheric isotopic baseline and facilitates comparison of tissue δ13C collected over multiple years. To exemplify the importance of accounting for atmospheric CO2 δ13C depletion, we applied the correction to a dataset of collagen δ13C obtained from mountain lion (Puma concolor) bone samples collected in California between 1893 and 1995. Before correction, in three of four ecoregions collagen δ13C decreased significantly concurrent with depletion of atmospheric CO2 δ13C (n ≥ 32, P ≤ 0.01). Application of the correction to collagen δ13C data removed trends from regions demonstrating significant declines, and measurement error associated with the correction did not add substantial variation to adjusted estimates. Controlling for long-term atmospheric variation and correcting tissue samples for changes in isotopic baseline facilitate analysis of samples that span a large temporal range.