Differential habitat use promotes sustainable coexistence between the specialist otter and the generalist mink

Optimal foraging and habitat selection theories predict that heterogeneous environments should favour the coexistence of competitors, especially when the dominant competitor is a specialist and the sub-ordinate is a generalist. In this paper, we analysed differential habitat use as a potential mechanism for the coexistence of two competing riparian mammals, the specialist and dominant Eurasian otter ( Lutra lutra ) and the generalist and sub-ordinate American mink ( Mustela vison ). We tested three hypotheses: H1: mink coexist with otters for longer in areas with abundance of habitats hosting terrestrial prey because, by not relying on aquatic prey, mink can segregate from its competitor. H2: the characteristics of the habitat closer to the riverbank will affect the length of time the two species coexist, because mink are still tied to the water even in the presence of otters. H3: denser vegetative cover along the bank increases the duration of coexistence of mink and otters because it reduces the frequency of their encounters. The first hypothesis was supported by the data and we found that in areas where terrestrial prey was abundant mink coexisted for longer with otters. The second hypothesis was also supported by the data and the characteristics of the habitat closer to the riverbank were the most important in determining coexistence time. Finally, we did not find supporting evidence for the third hypothesis. This study provides strong evidence that habitat heterogeneity plays an important role in determining the likelihood of coexistence of American mink with Eurasian otters. This result is particularly important from a conservation standpoint. Mink are invasive and a threat to endangered species in parts of their range. The knowledge that mink have a higher chance to persist in the presence of otters when terrestrial prey is abundant should be used to target areas for preferential mink management.     

Evidence for food competition between mink (Mustela vison) and otter (Lutra lutra) on Scottish islands

Food competition between American mink and otters was measured by comparing the diets of sympatric mink and otter populations with those of allopatric populations. Niche breadth was narrower for otters than mink. Niche breadth was wider for both mink and otters on islands where they co-existed in comparison to that of the allopatric populations. Niche overlap was lower in sympatric populations on islands with mammalian prey, however, niche overlap was not reduced on small islands without mammalian prey. Obtained data suggest that mink and otter compete for food resources and, when alternative prey sources are available, mink become more generalist predators to avoid competition with otters. However, when alternative prey sources are not available, both species become more generalist.     

Food niche variation of European and American mink during the American mink invasion in north-eastern Belarus

Understanding processes allowing the co-existence of ecologically similar species is important but difficult to study in community ecology. Introductions of alien species are unplanned experiments allowing investigation of co-adaptation of both native and invasive species over a short period. We analysed food niche differentiation between native European mink and alien American mink after invasion of the latter species in Belarus. European mink feed mainly on crayfish, frogs and fish whereas American mink prefer small mammals, fish and frogs. The diet of both species varied between seasons and during the period of alien mink invasion. Concurrent with the progress of American mink invasion, the European mink food niche has narrowed to feeding mainly on frogs, with the proportion of aquatic prey (fish and crayfish) in their diet drastically reduced. In contrast, the American mink food niche became wider during invasion. The breadth was stable but included a varied proportion of different prey categories: namely an increased proportion of aquatic prey and a decreased proportion of water vole and waterfowl. The increase in abundance of American mink saw a decrease in the proportion of larger prey in their diet. When American mink preyed more often on frogs, food niche overlap of both predators increased. This result suggests that arrival of an alien competitor reduced food abundance (exploitative competition) and caused a change in native mink diet.     

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.     

Impact of released Eurasian otters on a population of American mink: a test using an experimental approach

Competition is considered to be a major evolutionary driving force within assemblages of related and morphologically similar species, but it is notoriously difficult to test and quantify especially amongst species that cannot be easily manipulated in a laboratory or in the field. By exploiting a re-introduction of Eurasian otters ( Lutra lutra ) in the Upper Thames catchment (UK) in 1999 we performed an experiment to test whether there was evidence of competition between otters and American mink ( Mustela vison ). Mink and otters are semi-aquatic mustelids belonging to the same guild. Otters are expected to be the dominant competitor because they are larger and better adapted at exploiting aquatic resources. Our hypothesis was that mink declines when the density of otters increases. We measured the effect of competition at the population level, by observing whether mink distribution and densities changed in an area of 1353 km² in association with the arrival of its putative competitor. We estimated distribution and densities by means of sign surveys and trapping. The results showed that otters were associated with a significant and rapid reduction in the densities of mink, while mink occupancy remained approximately the same in an area of 2464 km² that was used as a control. We observed that the spatial distribution of the mink population throughout its decline was influenced by the yearly cycle of mink activities, with areas being temporarily re-colonized during the dispersal season. Mink is an invasive species in the UK threatening the survival of some native species. Our findings suggest that the re-establishment of otter populations is likely to lead to a decline of mink that may, in turn, be beneficial to native species threatened by this invasive.     

Feeding habits of three sympatric mammals in NE Spain: the American mink, the spotted genet, and the Eurasian otter

Diet composition, niche measures, and prey consumption of three sympatric species of carnivores, one non-native and introduced, the American minkNeovison vison Schreber, 1777, and two native, the spotted genetGenetta genetta Linnaeus, 1758 and the Eurasian otterLutra lutra Linnaeus, 1758, were studied in a Spanish Mediterranean area. The study was based on the analysis of prey remains in the faeces of the predators. Faeces of mink (n = 444), genet (n = 310), and otter (n = 108) were collected all year round for four years along the Llobregat (21 km) and Gavarresa rivers (12 km). Simultaneously, and in the same area, prey species density and weight were estimated by means of trapping. The diet of mink and genet was dominated by the American crayfishProcambarus clarkii, while the otter fed almost exclusively on fish. Compared to the mink, the genet seemed to specialise more on small mammals while the mink specialised more on aquatic birds, especially in autumn-winter. Niche overlap was higher between mink and genet than between either of them and the otter.     

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.     

Do alien North American mink compete for resources with native South American river otter in Argentinean Patagonia?

American mink Mustela vison , originally bred in fur farms, have become established in areas occupied by native endangered Southern river otter Lontra provocax , in Patagonia. In accordance with European experience, this biological invasion in South America raises questions about the interaction between invasive mink and native otter, from the viewpoints of both community assembly and conservation. We set out (1) to find which aspects of habitat structure were related to the distribution of signs of both this invasive species and Southern river otter Lontra provocax , in Argentinean Patagonia and their most common prey and (2) to test general predictions of niche partitioning between these two species. Based on surveys of 447 of 600 m transects for otter and mink scats/footprints along the waterside of lakes and rivers in the Andean Patagonian region, we compared diet composition (from scat analysis) and micro-habitat preferences (from field signs) of the two species. Otters were more specialist than mink in habitat use and diet. Mink used different habitats in other river basins where otters were absent. Where they occurred together in the basin of the Limay River, the distributions of their signs were similar, and mink diet was more similar to that of otters. There was no detectable difference in otter diet before and after mink arrival in the Limay basin. Contrary to the prediction of niche partitioning, and to the findings of European studies, resource use by mink was more similar to that of otters where the species occurred sympatrically than where they were allopatric.     

A Comparison of the diet and distribution of southern river otter (Lutra provocax) and mink (Mustela vison) in Southern Chile

In Chile, between latitudes 39S and 4330'S, southern river otter ( Lutra provocax ) and introduced mink ( Mustela vison ) scats were collected. Mink sign was recorded in 29% of the aquatic habitat where otter sign was found. Sixty-eight per cent of mink scats were collected at otter rest sites. A significant difference between the diets and low percentage of habitat use overlap of the two species (5-22%) suggest that, in Chile, river otter and mink coexist with little competition for space or food. There are no data supporting a relationship between the introduction of mink and the decline of southern river otters.     

The food of feral mink (Mustela vison) in Scotland

Feral American mink ( Mustela vison ) thrive in Scotland. The potential impact of the mink on the native fauna was investigated through its food. Fifty-five stomachs and 33 scats were analysed by identifying the indigestible remains. Fish formed the main prey (49% occurrence) and birds and mammals 28% and 23% respectively. Most of the fish eaten were Salmonidae (brown trout and salmon). No remains of eels were found.
Mink may compete with otters for food, but the smaller prey taken by the mink and its greater use of land separates it from the otter. Mink seem to be occupying a vacant feeding niche in Scotland.     

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.     

Review and quantitative meta-analysis of diet suggests the Eurasian otter (Lutra lutra) is likely to be a poor bioindicator

The Eurasian otter (Lutra lutra L.) is a top predator in aquatic systems and plays an important role in ecosystem functioning. However, it has undergone dramatic declines throughout Europe as a result of environmental degradation. We examine the putative role of the otter as a bioindicator in Ireland which remains a stronghold for the species and affords a unique opportunity to examine variation in its ecological niche. We describe diet, using spraint contents, along rivers during 2010 and conduct a review and quantitative meta-analysis of the results of a further 21 studies. We aimed to assess variation in otter diet in relation to river productivity, a proxy for natural nutrification and anthropogenic eutrophication, and availability of salmonid prey (Salmo trutta and Salmo salar), to test the hypothesis that otter diet is related to environmental quality. Otter diet did not vary with levels of productivity or availability of salmonids whilst Compositional Analysis suggested there was no selection of salmonid over non-salmonid fish. There was a distinct niche separation between riverine and lacustrine systems, the latter being dominated by Atlantic eel (Anguilla anguilla). Otters are opportunistic and may take insects, freshwater mussels, birds, mammals and even fruit. Otters living along coasts have a greatest niche breath than those in freshwater systems which encompasses a wide variety of intertidal prey though pelagic fish are rarely taken. It is concluded that the ability of the otter to feed on a wide diversity of prey taxa and the strong influence of habitat type, renders it a poor bioindicator of environmental water quality. It seems likely that the plasticity of the habitat and dietary niche of otters, and the extent of suitable habitat, may have sustained populations in Ireland despite intensification of agriculture during the 20th century.     

Decline of invasive alien mink (Mustela vison) is concurrent with recovery of native otters (Lutra lutra)

Invasive species often cause the decline of native prey or competitors. We highlight a contrasting example of the large-scale recovery of a native species and the concurrent decline and likely displacement of an established invasive competitor. Invasive American mink Mustela vison became widespread in the British Isles at the same time as native Eurasian otters Lutra lutra were declining as a result of water pollution. In common with other invasive predators, mink cause conservation problems for a range of native prey species, most notably water voles Arvicola terrestris . Recent trends in the distribution of native otters and invasive mink in north-east England were examined using a novel regression modelling approach to analyse presence/absence data from field surveys, corroborated by contemporary predator culling records. Between 1991 and 2002, the percentage of sites where mink signs were found decreased from 80% to 20%, while otter signs increased from 18% to 80%. Annual indices of mink captures on shooting estates increased between 1980 and 1996, but were followed by a decline thereafter. Indices of the incidence of native otters were significantly related to those indicating the decline or displacement of invasive mink. This large-scale field study is supportive of localized experimental evidence for the return of dominant, native otters being concurrent with the decline of the invasive alien mink. The recovery of a dominant native species may represent a reversal of the mesopredator release that allowed invasive mink to establish and may eventually serve to mitigate their impact on native prey species.     

Stream community richness predicts apex predator occupancy dynamics in riparian systems

Streams and adjacent riparian habitats represent linked terrestrial and aquatic ecosystems that exchange materials and energy. Recognized relationships among apex predators and ecosystem biodiversity led us to hypothesize that these predators in riparian‐stream systems were more likely to be found in sites with high stream quality, defined as increased ecosystem function and integrity. In our freshwater study system, river otter Lontra canadensis and mink Neovison vison play critical roles as apex predators. We used multi‐season occupancy modelling across three sampling years (2012–2014) to compare aspects of the stream communities that explain occupancy dynamics of river otter and mink, including their interactions with other semi‐aquatic mammals. We surveyed for semi‐aquatic mammals at 77 sites in 12 major watersheds in southern Illinois, USA (44 526 km²). Naïve occupancy differed among years but generally increased for river otter, and remained high (≥93.5%) for mink. Increasing substrate availability increased detectability of river otter, whereas mink detection varied by survey period. Occupancy of river otter during the initial survey period was higher in sites closer to reintroduction points. Probability of colonization of river otter was positively associated with macroinvertebrate index of biotic integrity, fish species richness, and beaver presence. Sites with high species richness of fish families preferred by river otter also had increased river otter persistence. Mink occupied sites with increased fish richness, muskrat presence and mussel community index. Taken together, our results show occupancy of both mink and river otter were predicted by aspects of prey diversity and presence, indicating the importance of community composition in occupancy dynamics of riparian predators. Ultimately, these relationships suggest that habitat heterogeneity and system stability are important to apex predator site use. However, the relative role of bottom–up and top–down forcing in stream systems remains to be resolved.     

A comparison of the feeding biology of Mink Mustela vison and otter Lutra lutra

The feeding habits and prey selectivity of Mink Mustela vison and otters Lutra lutra were compared in two localities in Devon: a eutrophic lake and a moorland river, in which both species occurred and had access to the same prey populations. The effects of prey availability on the predators' diets were assessed by comparing prey consumed, as revealed by scat analysis, with estimates of prey abundance and size range. Otters specialized in fish at all times of year but showed seasonal variation in species taken. Selection for slow-moving fish and seasonal changes in behaviour of some fish species were the probable causes of this variation. Otters diversified more into non-fish food in summer, when fish availability was reduced. The main alternative prey in the lacustrine habitat was waterfowl, but in the riverine habitat, rabbits. Mink were more generalized carnivores, taking a variety of fish, waterside and terrestrial prey in all seasons. These three prey categories were taken to an almost equal extent in the lake but terrestrial prey dominated in the riverine habitat. Fish were taken most frequently in winter and birds and mammals in summer. Neither predator showed selection in respect of prey size. In each area, about one third of the otter and Mink diets was common to both species. Fish was the principal group of the shared component, and dietary overlap in respect of them was greatest in autumn and winter. In view of the dietary preferences of each predator, the existence of alternative prey items and limited degree of dietary overlap, it is considered unlikely that the two species competed for food to any extent. Other factors must therefore be responsible for the spread of feral Mink and the decline in otter populations in many parts of Britain.     

Interactions of exotic and native carnivores in an ecotone, the coast of the Beagle Channel, Argentina

In coasts bordering highly productive seas, there can be a flux of resources to the terrestrial ecosystem, and terrestrial carnivores can use marine prey extensively. Two native, endangered species (otter Lontra provocax and culpeo fox Pseudalopex culpaeus lycoides) and two exotic species (mink Neovison vison and grey fox Pseudalopex griseus) inhabit the Beagle coast. The objectives of this paper are: (1) to describe the diet, habitat use and distribution of otters, mink and foxes on the coast of the Beagle channel and (2) to discuss the role of marine resources in the ecological interactions among these species. Diet was determined from the analysis of 245 faeces, and distribution was established from sign surveys. Marine prey occurred in the scats of 98.3, 70.4, 35.5 and 18.2% of otters, mink, culpeo and grey foxes, respectively. Other terrestrial species also use marine resources in Southern Patagonia. All this evidence suggests that the Fueguian coastal channels provide an illuminating example of allochthonous food subsidies from the sea. In the community of four sympatric predators, two native (and endangered) and two exotic, coexistence appears to be facilitated by a renewing marine food subsidy.     

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.     

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.     

Seasonal exploitation of amphibians by otters (Lutra lutra) in north-east Scotland

The dietary importance of amphibians to otters was studied in north-east Scotland in 1987. The proportion of amphibians identified in otter droppings varies seasonally and according to the habitat. The consumption of amphibians increases in late winter and spring while they are either hibernating or spawning. Estimations of amphibian population density at spawning sites have been carried out. There is a significant correlation between the availability of amphibians and their consumption by otters. The main amphibian species involved in the diet is the common frog, Rana temporaria (60.8%), while the common toad, Bufo bufo , represents only 1.7% of the amphibian prey.     

The Smell of New Competitors: The Response of American Mink, Mustela vison, to the Odours of Otter, Lutra lutra and Polecat, M. putorius

We tested the response of wild American mink (an established alien species in the UK), to the odours of unfamiliar mink, European polecat and Eurasian otter. Polecats are similar in size and habits to mink, otters are larger than mink and a dominant competitor; both are native to the UK and both were absent during the original colonization by mink but are now undergoing natural population recoveries. The response of mink to experimental odours was assessed by counting the numbers of tracks (footprints) on rafts treated with anal gland secretions, and compared with response to a control raft, on two rivers in the Upper Thames valley, UK. Remote video showed that the number of tracks was positively correlated with the time that mink spent investigating an odour. We found that mink were attracted to the odours of both unfamiliar mink and polecats. There was little evidence that mink avoided the odour of otters. We suggest that, during an encounter with a polecat, mink may behave much as they would to a conspecific. We infer from the response of mink to the odour of otters, that, if mink do avoid otters, the mechanism of avoidance is likely to be complex, situation-dependent and perhaps affected by prior experience.