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.     

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.     

Life‐history variability of non‐native centrarchids in regulated river systems of the lower River Guadiana drainage (south‐west Iberian Peninsula)

Life‐history variability of two non‐native centrarchids, pumpkinseed Lepomis gibbosus and largemouth bass Micropterus salmoides, was evaluated in three stream stretches of the lower River Guadiana drainage (south‐west Iberian Peninsula) with different degrees of regulated flows. Abundance, condition and population structure differed among populations for both species, but invasion success was lower in the least regulated river. Lepomis gibbosus were abundant and had multiple age classes in the three river sites, whereas M. salmoides were less abundant and mainly represented by young‐of‐the‐year fish. Juvenile growth in L. gibbosus was similar in all three populations, though longevity was slightly greater in the population from the River Guadiana mainstream. Lepomis gibbosus exhibited a long reproductive season, but the duration of season, size at maturity and reproductive effort varied among populations. The life‐history differences found demonstrate the importance of species adaptation to local conditions which might favour their invasion success. Lepomis gibbosus were more adaptable and resilient to local conditions, whereas M. salmoides seemed dependent on reservoirs and large rivers for maintenance of riverine populations.     

Morphological variation between non‐native lake‐ and stream‐dwelling pumpkinseed Lepomis gibbosusin the Iberian Peninsula

The objective of this study was to test if morphological differences in pumpkinseed Lepomis gibbosus found in their native range (eastern North America) that are linked to feeding regime, competition with other species, hydrodynamic forces and habitat were also found among stream‐ and lake‐ or reservoir‐dwelling fish in Iberian systems. The species has been introduced into these systems, expanding its range, and is presumably well adapted to freshwater Iberian Peninsula ecosystems. The results show a consistent pattern for size of lateral fins, with L. gibbosus that inhabit streams in the Iberian Peninsula having longer lateral fins than those inhabiting reservoirs or lakes. Differences in fin placement, body depth and caudal peduncle dimensions do not differentiate populations of L. gibbosus from lentic and lotic water bodies and, therefore, are not consistent with functional expectations. Lepomis gibbosus from lotic and lentic habitats also do not show a consistent pattern of internal morphological differentiation, probably due to the lack of lotic–lentic differences in prey type. Overall, the univariate and multivariate analyses show that most of the external and internal morphological characters that vary among populations do not differentiate lotic from lentic Iberian populations. The lack of expected differences may be a consequence of the high seasonal flow variation in Mediterranean streams, and the resultant low‐ or no‐flow conditions during periods of summer drought.     

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     

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.     

Conflict between Fishermen and Giant Otters Pteronura brasiliensis in Western Brazilian Amazon

The recovery of giant otter populations after the hunting prohibition and restriction of the pelt trade resulted in more frequent conflicts with fishermen. In this study, fisherman–giant otter conflicts were analyzed in the Uacari Sustainable Development Reserve, where giant otters are accused of interfering with fisheries by eating the fish (predation), frightening the fish away (local interference), and damaging fishing equipment (direct interference). Interference by predation was analyzed by evaluating overlap in fish species consumption between humans (measured by subsistence and commercial catches) and giant otters. The giant otter diet was assessed from fecal samples, and the human diet through questionnaires. Local and direct interferences were analyzed through fish samples using gillnets and comparing capture efficiency with and without giant otters’ presence. The overlap between human and giant otter diets was low (0.37), varied seasonally, and was smaller in the low water (0.24) than in the high water period (0.60), when both species tend to be more generalists. Overlap between fish species consumed by giant otters and those exploited by commercial fisheries was small (0.34). Giant otter presence during the experimental fishing was low (9.5%), restricted to the high water period, and did not significantly reduce the captures (U = 13, P = 0.61). The low overlap in diet may be a result of differences in preferences and fishing strategies. The conflict between giant otters and fishermen is greater in the high water period, when the income of the fisheries decreases; however, the conflict seems to be mainly motivated by the resident's prejudice against giant otters.     

Understanding the inter-specific dynamics of two co-existing predators in the Tierra del Fuego Archipelago: the native southern river otter and the exotic American mink

Knowledge about interactions between endangered native southern river otters (Lontra provocax) and introduced American mink (Neovison vison) is essential for effective management of both species. We evaluated competition for spatial and trophic niches between otter and mink in overlapping and non-overlapping areas, comparing distribution, habitat preference, diet and mink marking behavior. We surveyed otter and mink signs along 250 km of Beagle Channel coastline. Habitat suitability models were constructed based on species presence/absence and habitat characteristics, using generalized linear models. Feces were collected for diet analyses. Otters used forested coasts with 12°–32° shoreline slope and without human influence, and our evidence suggests they were not affected by mink presence. Mink preferred forested and shrubland coasts with 10°–28° shoreline slope. Neither human influence nor otter presence affected mink habitat occupation, but in the presence of otters, mink left fewer signs. Otters consumed more aquatic prey than mink, and mink modified their diet in the presence of otters, consuming more exotic small terrestrial mammals and less fish as well as shifting to smaller and shallower fish species that are less consumed by otters. Mink showed more plastic, generalist behavior than otters, being more tolerant of human presence, using more habitat types and having greater diet breadth. At the same time, otters apparently affect mink adversely and could help limit their invasion in sympatric areas. Conservation and recovery of otters, therefore, may produce a secondary benefit of simultaneously reducing the effect of mink, thereby providing an additional way to control this exotic predator’s population.     

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.     

Foraging strategies and prey switching in the California sea otter

Summary Southern sea otters (Enhydra lutris nereis), in recovering from near extinction, are gradually extending their range to include areas from which they have been absent for more than one hundred years. This study took advantage of the otters' relatively sudden arrival in the area near Santa Cruz, California, to monitor their prey selection in the first two years of residence there. Foraging observations revealed that sea urchins (Strongly-locentrotus franciscanus) were heavily preyed upon initially, but virtually disappeared from the diet after one year of sea otter residence. The disappearance of sea urchins was accompanied by an increased use of kelp crabs (Pugettia producta) and the appearance of clams (Gari californica) in the otters' diet. Abalones (Haliotis rufescens) and cancer crabs (Cancer spp.) remained fairly stable as dietary items throughout the two year period. An electivity index was used to quantify sea otter preferences, which corresponded closely with a ranking scheme based on energy intake/unit foraging time calculated for each major prey species. As predicted by optimal foraging theory, sea otters prefer food species of high rank and replace depleted dietary items with those of next highest rank. The process of dietary switching was analyzed with respect to foraging success rates, and it appears that poor success rates, associated with predation on an increasingly rarer prey species (sea urchins), drive sea otters to hunt for different prey. Both patch selection and search image formation appear to function in this process. The potential effects on community structure and stability of predators exhibiting a preference for the most profitable prey are discussed.     

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.     

Piscivorous fishes diet dominated by the Asian cyprinid invader,topmouth gudgeon (<Emphasis Type="Italic">Pseudorasbora parva</Emphasis>)

Predation on the non-native cyprinid, Pseudorasbora parva, was reported during an annual study in a small connecting canal about 40 km south of Brno, Czech Republic. Two cyprinid fish species (Pseudorasbora parva, Rutilus rutilus) were identified in the diet of 23 piscivores belonging to three native fishes (Perca fluviatilis, Sander lucioperca, Esox lucius). The relative abundance of prey fishes changed seasonally and was dominated by the topmouth gudgeon in spring. Thereafter roach incidence continuously increased. In contrast, topmouth gudgeon was found to be a more sized-available prey and was the most dominant prey species in the diet of all piscivorous fishes during the whole season. Strong piscivory towards this alien might be beneficial for its eradication or control to support ecological integrity in natural and modified seminatural habitats.     

Applying social and ecological approaches to evaluate factors influencing river otter (<Emphasis Type="Italic">Lontra canadensis</Emphasis>) visitation to fish-rearing facilities in Pennsylvania

In Pennsylvania the production of game and other fish species is a $1.6 billion industry, and loss from depredation can take a serious toll on fish stocks and associated revenues. Following a reintroduction project and natural expansion of native populations, river otters (Lontra canadensis) are now widely distributed throughout Pennsylvania. To determine occurrence and extent of depredation by river otters, we conducted telephone or mail surveys with 156 (80%) of Pennsylvania’s 196 fish-rearing facilities. River otters were reported in the vicinity of 50 (32%) facilities, of which 23 (46%) reported river otter visitation. Of the facilities that reported river otter visitation, most (87%) reported losing <5% of fish stock to otters and 91% reported an annual economic loss of <$500 from river otter depredation. Binary logistic regression determined that greater amounts of tree cover (%), facilities that reared >100,000 fish annually, and private facilities were more likely to experience visitation by river otters. Our results demonstrate that the existence of river otter populations in Pennsylvania does not appear to be a significant economic threat to fish-rearing facilities and physical and landscape characteristics of the facilities are important in determining the likelihood of river otter visitation.     

Diet of coastal foraging Eurasian otters (<Emphasis Type="Italic">Lutra lutra</Emphasis> L.) in Pembrokeshire south-west Wales

The importance of the marine environment to Eurasian otters is currently poorly understood. Wales is one of the few countries where coastal activity has been recorded and an increase in marine otter sightings could indicate remarkable developments within Welsh populations. The trophic niche of coastal otter populations around Pembrokeshire was investigated over a 12-month period. Marine activity was more widespread than previously thought and marine prey formed the largest component of otter diet, although, otters also consumed freshwater and terrestrial prey throughout the year. Otter diet was very diverse compared to other European coastal populations and a spring contraction in trophic niche width coincided with the estimated timing of breeding activity. Seasonal variation in prey composition was predominantly due to differences in the consumption of alternate prey types. In areas where wetlands are fragmented and populations of freshwater fish are declining, the marine environment may become an increasingly important habitat for otters. It is necessary to define the historical importance of coastal populations to otter conservation. Coastal areas are often subject to pressure from human activities, so the impact of disturbance needs to be assessed. Importantly, there is no verified otter survey method for coastal areas, so the use of marine habitat is likely to be underestimated.     

Genetic diversity and individual identification of reintroduced otters (<Emphasis Type="Italic">Lutra lutra</Emphasis>) in north-eastern Spain by DNA genotyping of spraints

During the last century, otter populations in the Mediterranean area of the Iberian Peninsula were dramatically reduced and disappeared in many localities. A reintroduction programme was established in north-eastern Spain (Muga and Fluvià basins and the “Aiguamolls de l’Empordà” wetlands), by releasing 42 otters from four different Iberian populations, between 1995 and 2002. In order to evaluate the success of the reintroduction programme, we investigated microsatellite variation in the native populations of released otters as well as in the population present in the release area in 2004. We used non-invasively collected samples as the DNA source to avoid disturbing the animals in the wild. Laboratory procedures included the screening of samples and a multiple-tubes approach to detect and correct genotyping errors. Our results show that founders have been replaced by descendants in the release area and the population is in Hardy-Weinberg equilibrium, with a 1:1 sex ratio. Western Iberian otters—representing two-thirds of the founder group and released earlier—have greatly contributed to the genetic composition of the current population. The genetic patterns of otters found in a basin north of the studied area suggest a common origin with the released population. We also detected in a few otters the presence of alleles not identified in the original founder group, and we shall discuss the possible origin of these alleles. This paper is dedicated to the memory of Dr. Xavier Domingo-Roura.     

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.     

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

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Resource seasonality and fish diets in an Illinois stream

Synopsis The purpose of this study was to evaluate the intensity of competition for food among 9 species of stream fishes that primarily eat aquatic invertebrates. The taxonomic and size composition, and numerical abundance of aquatic invertebrates were monitored for one year using drift and benthic samples. Diet data were obtained from stomachs of fishes captured at the same time and place that invertebrate sampling was done. Diet characteristics examined included taxonomic and size composition, number of prey per fish, and diet breadth. Drifting invertebrates were more abundant early in the year (March–June) than later (July–January). The summer-early fall scarcity of invertebrates was especially notable among those>3.6 mm long, which comprised the bulk of prey found in fish stomachs. Average prey size eaten by a fish species was positively correlated with fish mouth size, but interspecific overlap in prey size was extensive. Cyprinids as a group (5 species) ate proportionally fewer small (< 3.6 mm long) prey from July to January than did the centrarchids and stonecat. Taxonomic compositions of available invertebrates and fish diets varied markedly among sampling dates, but the use of prey taxa by fishes was not correlated with the availability of those taxa. Use of aquatic prey taxa was generally similar among fish species, but cyprinids as a group ate proportionally more terrestrial prey from July to January than did the centrarchids and stonecat. Diet breadths for all species increased as food levels declined, indicating that these fishes experienced resource depression. Food scarcity was evidently more severe for cyprinids since their stomachs contained few prey through the summer and fall relative to the centrarchids and stonecat. Though the fish species studied probably compete for food in the summer and fall, this competition did not account for the community structure observed.     

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.