Two complementary approaches were used to assess year-round variation in the diet of sea otters Enhydra lutris around Prince of Wales Island (POW) in southern Southeast Alaska, a region characterized by mixed-bottom habitat. We observed sea otters foraging to determine diet composition during the spring and summer. Then, we obtained sea otter vibrissae, which record temporal foraging patterns as they grow, from subsistence hunters to identify year-round changes in sea otter diets via stable isotope analysis of carbon (δ 13 C) and nitrogen (δ 15 N). We compared the stable isotopes from sea otter vibrissae and sea otter prey items that were collected during spring, summer, and winter. Overall, year-round sea otter diet estimates from stable isotope signatures and visual observations from spring and summer were dominated by clams in terms of biomass, with butter clams Saxidomus gigantea the most common clam species seen during visual observations. Our results indicate that these sea otters, when considered together at a regional level around POW, do not exhibit shifts in the main prey source by season or location. However, sea otter diets identified by stable isotopes had a strong individual-level variation. Behavioral variation among individual sea otters may be a primary driving factor in diet composition. This study provides quantitative diet composition data for modeling predictions of invertebrate population estimates that may aid in the future management of shellfisheries and subsistence hunting and the development of co-management strategies for this protected species.
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Sea otter effects on trophic structure of seagrass communities in southeast Alaska
Previous research in southeast Alaska on the effects of sea otters Enhydra lutris in seagrass Zostera marina communities identified many but not all of the trophic relationships that were predicted by a sea otter-mediated trophic cascade. To further resolve these trophic connections, we compared biomass, carbon (δ 13 C) and nitrogen (δ 15 N) stable isotope (SI), and fatty acid (FA) data from 16 taxa at 3 sites with high and 3 sites with low sea otter density (8.2 and 0.1 sea otters km -2 , respectively). We found lower crab and clam biomass in the high sea otter region but did not detect a difference in biomass of other seagrass community taxa or the overall community isotopic niche space between sea otter regions. Only staghorn sculpin differed in δ 13 C between regions, and Fucus , sugar kelp, butter clams, dock shrimp, and shiner perch differed in δ 15 N. FA analysis indicated multivariate dissimilarity in 11 of the 15 conspecifics between sea otter regions. FA analysis found essential FAs, which consumers must obtain from their diet, including 20:5ω3 (EPA) and 22:6ω3 (DHA), were common in discriminating conspecifics between sea otter regions, suggesting differences in consumer diets. Further FA analysis indicated that many consumers rely on diverse diets, regardless of sea otter region, potentially buffering these consumers from sea otter-mediated changes to diet availability. While sea otters are major consumers in this system, further studies are needed to understand the mechanisms responsible for the differences in biomarkers between regions with and without sea otters.
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- Award ID(s):
- 1600230
- NSF-PAR ID:
- 10324673
- Date Published:
- Journal Name:
- Marine Ecology Progress Series
- Volume:
- 674
- ISSN:
- 0171-8630
- Page Range / eLocation ID:
- 37 to 58
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Abstract The sea otter (
Enhydra lutris ) population of Southeast Alaska has been growing at a higher rate than other regions along the Pacific coast. While good for the recovery of this endangered species, rapid population growth of this apex predator can create a human‐wildlife conflict, negatively impacting commercial and subsistence fishing. Previous foraging studies throughout the sea otter range have shown they will reduce invertebrate prey biomass when recolonizing an area. The goal of this study was to examine and quantify the energy content of sea otter diets through direct foraging observations and prey collection. Our study area, Prince of Wales Island in southern Southeast Alaska, exhibits a gradient of sea otter recolonization, thus providing a natural experiment to test diet change in regions with different recolonization histories. Sea otter prey items were collected in three seasons (spring, summer, and winter) to measure caloric value and lipid and protein content. We observed 3523 sea otter dives during the spring and summer. A majority of the sea otter diet consisted of clams. Sea otters in newly recolonized areas had lower diet diversity, higher energetic intake rates (EIR, kcal/min), and prey had higher energy content (kcal/g). Females with pups had the highest diet diversity and the lowest EIR. Sea otter EIR were higher in the fall and winter vs. spring and summer. Sea cucumber energy and lipid content appeared to correspond with times when sea otters consumed the highest proportion of sea cucumbers. These caloric variations are an important component of understanding ecosystem‐level effects sea otters have in the nearshore environment. -
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Abstract Intraspecific variation, including individual diet variation, can structure populations and communities, but the causes and consequences of individual foraging strategies are often unclear.
Interactions between competition and resources are thought to dictate foraging strategies (e.g. specialization vs. generalization), but classical paradigms such as optimal foraging and niche theory offer contrasting predictions for individual consumers. Furthermore, both paradigms assume that individual foraging strategies maximize fitness, yet this prediction is rarely tested.
We used repeated stable isotope measurements (δ13C, δ15N;
N = 3,509) and 6 years of capture–mark–recapture data to quantify the relationship between environmental variation, individual foraging and consumer fitness among four species of desert rodents. We tested the relative effects of intraspecific competition, interspecific competition, resource abundance and resource diversity on the foraging strategies of 349 individual animals, and then quantified apparent survival as function of individual foraging strategies.Consistent with niche theory, individuals contracted their trophic niches and increased foraging specialization in response to both intraspecific and interspecific competition, but this effect was offset by resource availability and individuals generalized when plant biomass was high. Nevertheless, individual specialists obtained no apparent fitness benefit from trophic niche contractions as the most specialized individuals exhibited a 10% reduction in monthly survival compared to the most generalized individuals. Ultimately, this resulted in annual survival probabilities nearly 4× higher for generalists compared to specialists.
These results indicate that competition is the proximate driver of individual foraging strategies, and that diet‐mediated fitness variation regulates population and community dynamics in stochastic resource environments. Furthermore, our findings show dietary generalism is a fitness maximizing strategy, suggesting that plastic foraging strategies may play a key role in species' ability to cope with environmental change.
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Humans were considered external drivers in much foundational ecological research. A recognition that humans are embedded in the complex interaction networks we study can provide new insight into our ecological paradigms. Here, we use time-series data spanning three decades to explore the effects of human harvesting on otter–urchin–kelp trophic cascades in southeast Alaska. These effects were inferred from variation in sea urchin and kelp abundance following the post fur trade repatriation of otters and a subsequent localized reduction of otters by human harvest in one location. In an example of a classic trophic cascade, otter repatriation was followed by a 99% reduction in urchin biomass density and a greater than 99% increase in kelp density region wide. Recent spatially concentrated harvesting of otters was associated with a localized 70% decline in otter abundance in one location, with urchins increasing and kelps declining in accordance with the spatial pattern of otter occupancy within that region. While the otter–urchin–kelp trophic cascade has been associated with alternative community states at the regional scale, this research highlights how small-scale variability in otter occupancy, ostensibly due to spatial variability in harvesting or the risk landscape for otters, can result in within-region patchiness in these community states.more » « less
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Abstract Animals often consume resources from multiple energy channels, thereby connecting food webs and driving trophic structure. Such ‘multichannel feeding’ can dictate ecosystem function and stability, but tools to quantify this process are lacking. Stable isotope ‘fingerprints’ are unique patterns in essential amino acid (EAA) δ13C values that vary consistently between energy channels like primary production and detritus, and they have emerged as a tool to trace energy flow in wild systems. Because animals cannot synthesize EAAs de novo and must acquire them from dietary proteins, ecologists often assume δ13C fingerprints travel through food webs unaltered. Numerous studies have used this approach to quantify energy flow and multichannel feeding in animals, but δ13C fingerprinting has never been experimentally tested in a vertebrate consumer.
We tested the efficacy of δ13C fingerprinting using captive deer mice
Peromyscus maniculatus raised on diets containing bacterial, fungal and plant protein, as well as a combination of all three sources. We measured the transfer of δ13C fingerprints from diet to consumer liver, muscle and bone collagen, and we used linear discriminant analysis (LDA) and isotopic mixing models to estimate dietary proportions compared to known contributions. Lastly, we tested the use of published δ13C source fingerprints previously used to estimate energy flow and multichannel feeding by consumers.We found that EAA δ13C values exhibit significant isotopic (i.e. trophic) fractionation between consumer tissues and diets. Nevertheless, LDA revealed that δ13C fingerprints are consistently routed and assimilated into consumer tissues, regardless of isotopic incorporation rate. Isotopic mixing models accurately estimated the proportional diets of consumers, but all models overestimated plant‐based protein contributions, likely due to the digestive efficiencies of protein sources. Lastly, we found that δ13C source fingerprints from published literature can lead to erroneous diet reconstruction.
We show that δ13C fingerprints accurately measure energy flow to vertebrate consumers across tissues with different isotopic incorporation rates, thereby enabling the estimation of multichannel feeding at various temporal scales. Our findings illustrate the power of δ13C fingerprinting for quantifying food web dynamics, but also reveal that careful selection of dietary source data is critical to the accuracy of this emerging technique.
- Free Publicly Accessible Full Text
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Accepted Manuscript1.0
- Journal Article:
- https://doi.org/10.3354/meps13819
