DIET AND FORAGING BEHAVIOR OF SEA OTTERS IN SOUTHEAST ALASKA

Abstract: Direct observations of feeding sea otters ( Enhydra lutris ) at 11 sites in southeast Alaska showed infaunal clams to be the primary prey utilized by otters throughout the region. Foraging dive times associated with clam and sea urchin prey were significantly longer than those for more easily captured prey (crabs and mussels). Dive times and surface intervals were also generally correlated with water depth or apparent difficulty in obtaining buried prey. Male otters, which fed more extensively on clams than females, made significantly longer foraging dives than females. Foraging success remained high, even at sites where prey numbers were found to be very low during a related study. The very deeply burrowing geoduck clam ( Panope abrupta ), while common at several otter feeding sites, was rarely captured by otters. These results, combined with those of a companion study on prey numbers, indicate that butter clams ( Saxidomus giganteus ) account for the majority of the sea otter diet in southeast Alaska, and that sea urchins may represent relatively short-term prey in comparison to infaunal bivalves in regions where both prey types co-exist. Furthermore, the importance of butter clams in the sea otter diet and the tendency for this bivalve to retain chronically high levels of paralytic shellfish poisoning toxins in southeast Alaska increases the probability that toxic phytoplankton blooms influence sea otter distribution in this region.     

Prey selection in coastal Eurasian otters Lutra Iutra

Prey preferences and dietary differences between sex and age categories of Eurasian otters were studied in coastal Norwegian habitats Relative to their trapping frequency potential prey species with hard, spiny exoskeletons (crabs and sea urchins) or otherwise tough, spiny integuments (Labridae) were much less frequently found in spraints than fish species with soft integuments Spines did not protect fish with otherwise soft integuments from otter predation The number of non-fish taxa per otter stomach did not vary significantly between otter age categories despite presumed differences in hunting abilities (small cubs large cubs and subadults, adults) Relative frequency of occurrence of crabs and sea urchins was < 5% in the stomachs in each of these otter categories Anadromous, katadromous and freshwater fish species were infrequently eaten The coastal otter population during the study period probably had access to an adequate, and preferred, supply of marine fish prey
At the otter population level no prey size selection was conclusively demonstrated within the range of fish sizes sampled However, fish sizes eaten differed significantly between otter sex and age categories The fish sizes per stomach were on average larger in males than in females, regardless of age Adult males tended to eat the largest fishes Among the self provisioning age categories (subadult and adult otters) fish lengths differed significantly between otter males and females, but not between the otter age categories, and did not covary significantly with otter body length Fish eaten by females with old placental scars (potential mothers of fisheating cubs) were significantly smaller than those eaten by small cubs, provisioned by their mothers     

Reductions in the dietary niche of southern sea otters (Enhydra lutris nereis) from the Holocene to the Anthropocene

The sea otter (Enhydra lutris) is a marine mammal hunted to near extinction during the 1800s. Despite their well‐known modern importance as a keystone species, we know little about historical sea otter ecology. Here, we characterize the ecological niche of ancient southern sea otters (E. lutris nereis) using δ¹³C analysis and δ¹⁵N analysis of bones recovered from archaeological sites spanning ~7,000 to 350 years before present (N = 112 individuals) at five regions along the coast of California. These data are compared with previously published data on modern animals (N = 165) and potential modern prey items. In addition, we analyze the δ¹⁵N of individual amino acids for 23 individuals to test for differences in sea otter trophic ecology through time. After correcting for tissue‐specific and temporal isotopic effects, we employ nonparametric statistics and Bayesian niche models to quantify differences among ancient and modern animals. We find ancient otters occupied a larger isotopic niche than nearly all modern localities; likely reflecting broader habitat and prey use in prefur trade populations. In addition, ancient sea otters at the most southerly sites occupied an isotopic niche that was more than twice as large as ancient otters from northerly regions. This likely reflects greater invertebrate prey diversity in southern California relative to northern California. Thus, we suggest the potential dietary niche of sea otters in southern California could be larger than in central and northern California. At two sites, Año Nuevo and Monterey Bay, ancient otters had significantly higher δ¹⁵N values than modern populations. Amino acid δ¹⁵N data indicated this resulted from shifting baseline isotope values, rather than a change in sea otter trophic ecology. Our results help in better understanding the contemporary ecological role of sea otters and exemplify the strength of combing zooarchaeological and biological information to provide baseline data for conservation efforts.     

Influence of occupation history and habitat on Washington sea otter diet

Habitat characteristics are primary determinants of nearshore marine communities. However, biological drivers like predation can also be important for community composition. Sea otters (Enhydra lutris ssp.) are a salient example of a keystone species exerting top‐down control on ecosystem community structure. The translocation and subsequent population growth and range expansion of the northern sea otter (Enhydra lutris kenyoni) in Washington State over the last five decades has created a spatio‐temporal gradient in sea otter occupation time and density, and acts as a natural experiment to quantify how sea otter population status and habitat type influence sea otter diet. We collected focal observations of sea otters foraging at sites across the gradient in varying habitat types between 2010 and 2017. We quantified sea otter diet composition and diversity, and long‐term rates of energy gain across the gradient. We found that sea otter diet diversity was positively correlated with cumulative sea otter density, while rate of energy gain was negatively correlated with cumulative density. Additionally, we found that habitat type explained 1.77 times more variance in sea otter diet composition than sea otter cumulative density. Long‐term diet studies can provide a broader picture of sea otter population status in Washington State.     

Indirect food web interactions: sea otters and kelp forest fishes in the Aleutian archipelago

Although trophic cascades—the effect of apex predators on progressively lower trophic level species through top-down forcing—have been demonstrated in diverse ecosystems, the broader potential influences of trophic cascades on other species and ecosystem processes are not well studied. We used the overexploitation, recovery and subsequent collapse of sea otter (Enhydra lutris) populations in the Aleutian archipelago to explore if and how the abundance and diet of kelp forest fishes are influenced by a trophic cascade linking sea otters with sea urchins and fleshy macroalgae. We measured the abundance of sea urchins (biomass density), kelp (numerical density) and fish (Catch per unit effort) at four islands in the mid-1980s (when otters were abundant at two of the islands and rare at the two others) and in 2000 (after otters had become rare at all four islands). Our fish studies focused on rock greenling (Hexagrammos lagocephalus), the numerically dominant species in this region. In the mid-1980s, the two islands with high-density otter populations supported dense kelp forests, relatively few urchins, and abundant rock greenling whereas the opposite pattern (abundant urchins, sparse kelp forests, and relatively few rock greenling) occurred at islands where otters were rare. In the 2000, the abundances of urchins, kelp and greenling were grossly unchanged at islands where otters were initially rare but had shifted to the characteristic pattern of otter-free systems at islands where otters were initially abundant. Significant changes in greenling diet occurred between the mid-1980s and the 2000 although the reasons for these changes were difficult to assess because of strong island-specific effects. Whereas urchin-dominated communities supported more diverse fish assemblages than kelp-dominated communities, this was not a simple effect of the otter-induced trophic cascade because all islands supported more diverse fish assemblages in 2000 than in the mid-1980s.     

Diet composition of the Eurasian otter Lutra lutra in different freshwater habitats of temperate Europe: a review and meta‐analysis

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Commensal association of the common kingfisher with foraging Eurasian otters

Positive interactions between birds and mammals are a fascinating aspect of animal behaviour. Feeding associations may consist of local enhancement or facilitation, and in the latter case, of commensalism or mutualism depending on the benefits received by the facilitator. We report here on a previously undescribed feeding association between piscivorous birds and Eurasian otters Lutra lutra. In Spain, common kingfisher Alcedo atthis and grey heron Ardea cinerea were observed closely following foraging otters and benefited from feeding opportunities provided by these. Behavioural observations of otters in central Spain (28.4 hr; 19 days) revealed that an association with kingfishers occurred in 33% of otter foraging events (n = 92). Simultaneous observations in northern Spain (14.2 hr; 16 days) showed an association between otters and kingfishers or grey herons in 41.6% and 11.7% of otter foraging events (n = 77), respectively. The association probability between kingfishers and otters increased significantly when otters foraged closer to the shore and on small fish rather than other prey (crayfish or large fish). Birds fed on prey remain left by the feeding otters, on small fish captured by otters when these were satiated and playing, or on prey displaced by otters. Our observations are consistent with facilitation and commensalism: piscivorous birds gained feeding opportunities provided by the otters, with no apparent costs or benefits to the latter. Similar feeding associations have been described between other species of otters and piscivorous birds (kingfishers, herons, egrets, storks) in Asia, South America and Southern Africa, but had not yet been described in Europe. The occurrence of piscivorous bird–otter associations in different species and regions suggests that this commensalism may be often overlooked but widespread. We have shown that the association can be frequent and is context‐dependent, with benefits for associating birds depending on otters´ behaviour and targeted prey.     

SEA OTTER DIETS IN A DECLINING POPULATION IN ALASKA

Diet of sea otters ( Enhydra lutris ) using a haulout site on the north side of the Alaska Peninsula was determined from 50 scats. Dominant prey species were mussels ( Mytilus edulis ), followed by three species of clams ( Siliqua spp., Spisula polynyma , and Tellina lutea ), sand dollars ( Echinarachnius parma ), and helmet crabs ( Telmessus cheiragonus ). Our results support preliminary findings by Cim-berg et al. (1984) that this sea otter population preys heavily on mussels and that the presence of low caloric value sand dollars in their diet is significant. Coupled with population estimates, our results also provide evidence that this population of sea otters may be declining due, in part, to overdepletion of food resources.     

Eurasian otter (<Emphasis Type="Italic">Lutra lutra</Emphasis>) diet and prey selection in Mediterranean streams invaded by centrarchid fishes

The diet of the Iberian otter (Lutra lutra) was determined by analysing 547 spraints collected at 28 sites within a wide area invaded by centrarchid fishes (pumpkinseed sunfish, Lepomis gibbosus and largemouth bass, Micropterus salmoides): the middle Guadiana basin (South-west Iberian Peninsula). Fish was the otters’ main prey, representing more than 60% of total individuals and more than 80% of total biomass. Otters preyed on most of the fish species captured in the field; however, the consumption of centrarchids was low compared to their abundance in the streams, and Jacobs’ index of preference showed a clear rejection of both species by the otter. Consumption of native fish genera (Squalius, Barbus and Chondrostoma) by otters increased in relation to their increase in the environment. In contrast, increasing numbers of L. gibbosus in the field was not reflected in otter consumption. The general decline of native freshwater fishes in Iberian rivers, the preferred prey of otters, together with the spread of exotic fish species (centrarchids and others) could put otter populations at risk.     

Non‐trophic impacts from white sharks complicate population recovery for sea otters

Complex interactions between protected populations may challenge the recovery of whole ecosystems. In California, white sharks (Carcharodon carcharias) mistargeting southern sea otters (Enhydra lutris nereis) are an emergent impact to sea otter recovery, inhibiting the broader ecosystem restoration sea otters might provide. Here, we integrate and analyze tracking and stranding data to compare the phenology of interactions between white sharks and their targeted prey (elephant seals, Mirounga angustirostris) with those of mistargeted prey (sea otters, humans). Pronounced seasonal peaks in shark bites to otters and humans overlap in the late boreal summer, immediately before the annual adult white shark migration to elephant seal rookeries. From 1997 to 2017, the seasonal period when sharks bite otters expanded from 2 to 8 months of the year and occurred primarily in regions where kelp cover declined. Immature and male otters, demographics most associated with range expansion, were disproportionately impacted. While sea otters are understood to play a keystone role in kelp forests, recent ecosystem shifts are revealing unprecedented bottom‐up and top‐down interactions. Such shifts challenge ecosystem management programs that rely on static models of species interactions.     

Food partitioning and spatial subsidy in shelter‐limited fishes inhabiting patchy reefs of Patagonia

The diets of the most conspicuous reef‐fish species from northern Patagonia, the carnivorous species Pseudopercis semifasciata, Acanthistius patachonicus, Pinguipes brasilianus and Sebastes oculatus were studied. Pinguipes brasilianus had the narrowest diet and most specialized feeding strategy, preying mostly on reef‐dwelling organisms such as sea urchins, limpets, bivalves, crabs and polychaetes. The diet of A. patachonicus was characterized by the presence of reef and soft‐bottom benthic organisms, mainly polychaetes, crabs and fishes. Pseudopercis semifasciata showed the broadest spectrum of prey items, preying upon reef, soft‐bottom and transient organism (mainly fishes, cephalopods and crabs). All S. oculatus guts were empty, but stable‐isotope analyses suggested that this species consumed small fishes and crabs. In general, P. brasilianus depended on local prey populations and ate different reef‐dwelling prey than the other species. Pseudopercis semifasciata, A. patachonicus and probably S. oculatus, however, had overlapping trophic niches and consumed resources from adjacent environments. The latter probably reduces the importance of food as a limiting resource for these reef‐fish populations, facilitating their coexistence in spite of their high trophic overlap.     

Eurasian otters modify their trophic niche after the introduction of non-native prey in Mediterranean fresh waters

Biological invasions are a major driver behind disturbances in freshwater community structure. We investigated how the foraging strategy of the Eurasian otter Lutra lutra (L.) has responded to this change in a Mediterranean catchment of the Iberian Peninsula, over a 30-year time span (1980 vs. 2010). We found that, despite continuing to feed on a considerable amount of native fish, this carnivore has adapted its trophic niche to accommodate non-native species. Prior to the massive introductions of these new food items (1980), otters diversified their diet during the limiting season (i.e. summer drought). However, otters had largely incorporated non-native prey into their diet in 2010, with red-swamp crayfish being their main food resource, especially in the summer. This increased crayfish consumption reflects a narrowing of trophic niche during the summer drought in 2010. We discuss how this heavy specialisation may compromise the conservation of otters and their endemic fish prey.     

The influence of wave exposure on the foraging activity of marine otter, <Emphasis Type="Italic">Lontra felina</Emphasis> (Molina, 1782) (Carnivora: Mustelidae) in northern Chile

The marine otter Lontra felina has been said to prefer wave-exposed habitats over more protected sites in response to a greater prey abundance in exposed habitat. We examined how the foraging activity of L. felina is affected by the regime of wave exposure and prey availability at Isla Choros, northern Chile. Through focal sampling we recorded time spent by otters in foraging, the duration of dives, and the hunting success on a wave-exposed and a wave-protected site on the island. In addition, we quantified the abundance of prey in both habitats. Marine otters spent more time foraging in the wave-protected site compared with the wave-exposed habitat. Successful dives reached 26.9% in the wave-exposed habitats, and 38.2% in the wave-protected habitat. Foraging dives were 18% shorter in wave-exposed as compared with wave-protected habitat. Numerically, available prey did not differ significantly with habitat. Our results are more consistent with the hypothesis that wave-exposed habitats represent a sub-optimal habitat to foraging marine otters. Marine otters’ use of wave-exposed patches through northern and central Chile coastal areas probably reflects a low availability of suitable protected areas and greater human disturbance of more protected habitat.     

Growth-related changes in diet and foraging behavior of the yellow wrasseThalassoma lutescens at Kuchierabu Island

Ontogenetic changes in diet and foraging behavior ofThalassoma lutescens were examined in shallow reef habitats around Kuchierabu Island, southern Japan. This species mainly took small benthic invertebrates, including gammarids, polychaetes, sipunculids, chitons, crabs, gastropods, pelecypods and urchins from algal mats. Larger fish consumed correspondingly larger prey, although most of the latter were armored with hard exoskeletons, shells or body plates (e.g., crabs, gastropods, pelecypods and urchins). Such hard parts were crushed with the molar-like, pharyngeal teeth which develop with fish growth, allowing exploitation of such larger, hard-bodied prey. Because the densities of larger prey species were relatively low in the initial habitats foraged, larger fish shifted their foraging attention to rock and coral crevices, where the prey species dwelt in greater numbers, as well as foraging over larger areas. Such behavioral changes maintained high foraging efficiency in larger fish.     

Feeding habits of the otter and the American mink in Bialowieza Primeval Forest (Poland) compared to other Eurasian populations

Diets of the otter Lutra lutra and the American mink Mustela vison were studied by scat analysis on five woodland rivers and streams in eastern Poland. Fish constituted 51% of food biomass consumed by otters in spring‐summer and 40% in autumn‐winter, with common fish (perch Perca fluviatilis, pike Esox lucius, and roach Rutilus rutilus) being captured most frequently by the otters. Amphibians (mainly Rana temporaria, which also dominated in the living community) made up 34% of otters’ food biomass in spring‐summer and 58% in autumn‐winter. American mink relied on three prey groups: fish (40% in spring‐summer, and 10% in autumn‐winter), frogs (32% and 51%, respectively), and small mammals (21% and 36%). Out of available Micromammalia, mink strongly selected the root vole Microtus oeconomus. The cold season diet of both otter and mink depended on river size. On small rivers with forested valleys, otters and mink fed nearly exclusively on amphibians (72–90% of food biomass). With size of a river increasing and riverside habitat becoming more open (sedge and reed marshes instead of forests), otters shifted to catching predominantly fish (up to 76% in diet) and mink to preying on small mammals (up to 65% in diet).
Review of literature on otter and mink in Eurasia showed that their diets did not change with latitude (as indicators of climate severity and duration of water freezing) but they depended on habitats. In otter diet, the mean share of fish declined from 94% (SE 1.7) on sea shores, to 71% (SE 2.9) on lakes and fish ponds, to 64% (SE 2.8) on rivers and streams. The roles of amphibians and crustaceans increased in the same gradient (from 0 to 15%, and from 3 to 7%, respectively). On inland waters, the abundance of crayfish was the essential factor differentiating otters’ diet composition. In Eurasia, the staple food types of American mink on rivers and streams were fish (on average, 27% in diet, SE 3.9), mammals (30%, SE 5.0), and amphibians (17%, SE 4.8), whereas on lakes and ponds mink fed predominantly on birds (on average, 33% in diet, SE 10.1) and fish (28%, SE 9.5). In the Palaearctic region, over a wide gradient of habitats, otters appeared strongly specialised on prey taken from water, whereas American mink was a typical generalist capable of utilising several prey groups originating from both water and land.     

Dietary complexity and hidden costs of prey switching in a generalist top predator

1. Variation in predator diet is a critical aspect of food web stability, health, and population dynamics of predator/ prey communities. Quantifying diet, particularly among cryptic species, is extremely challenging, however, and differentiation between demographic subsets of populations is often overlooked.
2. We used prey remains and data taken postmortem from otter Lutra lutra to determine the extent to which dietary variation in a top predator was associated with biotic, spatial, and temporal factors.
3. Biotic data (e.g., sex, weight, and length) and stomach contents were taken from 610 otters found dead across England and Wales between 1994 and 2010. Prey remains were identified to species where possible, using published keys and reference materials. Multi‐model inference followed by model prediction was applied to test for and visualize the nature of associations.
4. Evidence for widespread decline in the consumption of eels (Anguilla anguilla) reflected known eel population declines. An association between eel consumption and otter body condition suggested negative consequences for otter nutrition. Consumption of Cottus gobio and stickleback spp. increased, but was unlikely to compensate (there was no association with body condition). More otters with empty stomachs were found over time. Otter sex, body length, and age‐class were important biotic predictors of the prey species found, and season, region, and distance from the coast were important abiotic predictors.
5. Our study is unique in its multivariate nature, broad spatial scale, and long‐term dataset. Inclusion of biotic data allowed us to reveal important differences in costs and benefits of different prey types, and differences between demographic subsets of the population, overlaid on spatial and temporal variation. Such complexities in otter diet are likely to be paralleled in other predators, and detailed characterization of diet should not be overlooked in efforts to conserve wild populations.

     

Inter-decadal patterns of population and dietary change in sea otters at Amchitka Island, Alaska

After having been hunted to near-extinction in the Pacific maritime fur trade, the sea otter population at Amchitka Island, Alaska increased from very low numbers in the early 1900s to near equilibrium density by the 1940s. The population persisted at or near equilibrium through the 1980s, but declined sharply in the 1990s in apparent response to increased killer whale predation. Sea otter diet and foraging behavior were studied at Amchitka from August 1992 to March 1994 and the data compared with similar information obtained during several earlier periods. In contrast with dietary patterns in the 1960s and 1970s, when the sea otter population was at or near equilibrium density and kelp-forest fishes were the dietary mainstay, these fishes were rarely eaten in the 1990s. Benthic invertebrates, particularly sea urchins, dominated the otter’s diet from early summer to mid-winter, then decreased in importance during late winter and spring when numerous Pacific smooth lumpsuckers (a large and easily captured oceanic fish) were eaten. The occurrence of spawning lumpsuckers in coastal waters apparently is episodic on a scale of years to decades. The otters’ recent dietary shift away from kelp-forest fishes is probably a response to the increased availability of lumpsuckers and sea urchins (both high-preference prey). Additionally, increased urchin densities have reduced kelp beds, thus further reducing the availability of kelp-forest fishes. Our findings suggest that dietary patterns reflect changes in population status and show how an ecosystem normally under top-down control and limited by coastal zone processes can be significantly perturbed by exogenous events. Received: 23 March 1999 / Accepted: 19 January 2000     

Testing the generality of the trophic cascade paradigm for sea otters: a case study with kelp forests in northern Washington,USA

Trophic cascade hypotheses for biological communities, linking predation by upper trophic levels to major features of ecological structure and dynamics at lower trophic levels, are widely subscribed and may influence conservation policy. Few such hypotheses have been evaluated for temporal or spatial generality. Previous studies of sea otter (Enhydra lutris) predation along the outer coast of North America suggest a pattern, often elevated to the status of paradigm, in which sea otter presence leads to reduced sea urchin (Strongylocentrotus spp.) biomass and rapid increases in abundance and diversity of annual algal species, followed by a decline in diversity as one or a few perennial algal species become dominant. Both sea otter predation and commercial sea urchin harvest are ecologically and economically important sources of urchin mortality in nearshore benthic systems in northern Washington marine waters. We recorded changes in density of macroalgae in San Juan Channel, a marine reserve in the physically protected inland waters of northern Washington, resulting from three levels of experimental urchin harvest: (1) simulated sea otter predation (monthly complete harvest of sea urchins), (2) simulated commercial urchin harvest (annual size-selective harvest of sea urchins), and (3) no harvest (control). The two experimental urchin removal treatments did not significantly increase the density of perennial (Agarum and Laminaria) or annual (Desmarestia, Costaria, Alaria and Nereocystis) species of macroalgae after 2 years, despite significant and persistent decreases in urchin densities. Our results suggest that other factors such as grazing by other invertebrates, the presence of dense Agarum stands, and recruitment frequency of macroalgae and macroinvertebrates may play a large role in influencing community structure in San Juan Channel and other physically protected marine waters within the range of sea otters. Handling editor: J. Trexler     

Do non‐native fish as prey favour the conservation of the threatened indigenous Eurasian otter?

1. Biological invasions are considered a major threat to biodiversity. Most research has focused on the distribution, biology and impacts of non‐native species on native fauna and flora. However, few studies have explored their role as prey for native predators of conservation concern. 2. To assess the incidence and intensity of predation by the Eurasian otter Lutra lutra on established non‐native fish species, data were collated from the published literature. To be selected, studies had to cover at least 1 year, analyse more than 100 spraints and report the study period and percentage relative frequency (%RF) of all prey fish species. 3. To permit reliable, time‐related comparisons with %RF of non‐native fishes in otter diet, we also reviewed available information about both the distribution of non‐native fishes and history of their introductions to European countries, revealing a decrease with longitude in the number of naturalised non‐native fishes taken (ranging between 5 and 34) and their percentage in each fish assemblage. 4. Our selective criteria were met by 30 dietary studies from 44 study areas in 15 European countries during 1970–2010. The extent to which otters rely on non‐native fishes was almost negligible (mean %RF = 4.8), with the number of non‐native fishes preyed upon by otters decreasing with both latitude and longitude. 5. The %RF of non‐native fish in the diet increased slightly with time, with otters preying significantly more on non‐native fish in study areas where alterations of the fish assemblage had been highlighted in the reference papers. No relationship was found between otter diet breadth and the occurrence of non‐native fishes in their diet. 6. The current role of non‐native species in otter diet suggests that effective otter conservation management plans should focus on the maintenance and/or enhancement of native fish assemblages.     

Amphibians in Eurasian otter Lutra lutra diet: osteological identification unveils hidden prey richness and male‐biased predation on anurans

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