European water voles in a reconnected lowland river floodplain: habitat preferences and distribution patterns following the restoration of flooding

Water voles have suffered large population declines in the United Kingdom due to habitat degradation and predation by invasive American mink. Habitat restoration of floodplain wetlands could help to reverse this decline, but the detailed habitat preferences of water voles in these environments have not been well studied, and the impacts of restoration practices on water vole populations are not known. This study investigated the habitat preferences of water voles in a reconnected lowland river floodplain. The results show that water voles preferred wider water bodies, and taller and more diverse vegetation. The impact of flooding on water voles was also investigated by comparing their occurrence between two survey periods which were separated by large flood events, and by comparing distribution patterns before and after restoration. Contrary to previous reports, there was no observed negative impact of flood events on water vole distribution, which has slightly expanded since the floodplain was reconnected to the river in 2009. Overall this study demonstrates that restored wetlands can provide suitable habitat for water voles, and provides guidance on some of the factors which should be considered when designing floodplains for water vole conservation.     

Dynamic impacts of feral mink predation on vole metapopulations in the outer archipelago of the Baltic Sea

Predation impacts by introduced predators are predicted to be most intense in island ecosystems, and also variable depending on environmental conditions, but large-scale experimental field testing is rare. In this study we examine the factors that determine the distribution and abundance of vole metapopulations preyed upon by feral American mink Mustela vison in the outer Finnish archipelago of the Baltic Sea. Specifically, we follow the dynamics of field voles Microtus agrestis and bank voles Clethrionomys glareolus on 40 small islands under variable rainfall as part of a large-scale mink removal experiment. For both vole species occupancy rates were negatively influenced by island isolation, as were extinction events for field voles. High summer rainfall in 1998 corresponded to large vole populations where mink were absent, populations that then crashed in 1999 and 2000 when below average rains fell during the summer breeding season. Where mink were present however, vole abundance remained more constant between years with no boom-bust apparent. We conclude that weather and predation may drive vole abundance whereas habitat patchiness and metapopulation processes more strongly drive vole distributions. There may also be potential for interaction between these factors: because feral mink prevent rapid vole population growth after good summer rains, and vole dispersal is influenced by population size, feral mink may be changing vole dispersal patterns to disrupt the natural metapopulation dynamic. Hence this indirect impact of mink could lead to gradual erosion of vole populations in the outer archipelago by reducing recolonisation processes.     

The influence of American mink odour on the spatial distribution and behaviour of water voles

The perception and assessment of predation risk often cause changes in the activities of animals and induce behavioural responses that may in turn affect their movements and distribution. To simulate high predation risk in a midfield pond riparian habitat, we used fresh faeces from ranch American mink Neovison vison and recorded behavioural responses of water voles Arvicola amphibius. In areas where mink odour was deployed, the numbers of captured vole individuals and their trappability were significantly lower than in control areas. Several voles migrated from the zones with deployed mink faeces to the areas without faeces, thus proving that increased predation risk affects the distribution of individuals in a population. The response to mink odour was much more pronounced in females than in males; in areas with deployed mink faeces, not a single female was trapped. We conclude that although American mink is a non‐native, invasive predator, water voles respond to mink odour by reducing their activity and/or by avoiding places with higher predation risk.     

Movements of radio-tracked American mink (<Emphasis Type="Italic">Neovison vison</Emphasis>) in extensive wetland in the UK,and the implications for threatened prey species such as the water vole (<Emphasis Type="Italic">Arvicola amphibius</Emphasis>)

In a number of countries around the world, introduced American mink are acknowledged to have had a negative effect on a number of native species, many of which are of particular conservation concern. In the UK, there has been an observed correlation between the spread of mink and the decline of the, once common and widespread, water vole. Large wetlands, such as extensive reed bed, appear to mitigate the impact of mink predation on water voles and some bird species. The present study was carried out to test the hypothesis that the observed refuge effect of reed beds arises from the way mink forage in this type of wetland. The results suggest that the interior of reed bed offers a spatial refuge for water voles, and other species, from predation by mink, because more than 60% of mink foraging activity occurred within 10 m of a main (>10 m wide) channel. Where mink ventured within the reed bed itself, they associated closely with scrub. The implication is that easily navigable channels and areas of scrub probably compromise the refuge effect of reed beds. This can be used to inform management recommendations at these sites.     

Voles on small islands: effects of food limitation and alien predation

Ecosystems of three trophic levels may be bottom-up (by food-plant availability) and/or top-down (by predators) limited. Top-down control might be of greater consequence when the predation impact comes from an alien predator. We conducted a replicated two-factor experiment with field voles (Microtus agrestis) during 2004-2005 on small islands of the outer archipelago of the Baltic Sea, south-west Finland, manipulating both predation impact by introduced American mink (Mustela vison) and winter food supply. In autumn 2004, we live-trapped voles on five islands from which mink had been consistently removed, and on four islands where mink were present, and provided half of these islands with 1.8 kg oats per vole. Body mass of female voles increased as a response to supplementary food, whereas both food supplementation and mink removal increased the body mass of male voles in subsequent spring. During winter, there was a positive effect of supplementary food, but in the subsequent summer, possible positive long-term impacts of food supplementation on field voles were not detected. Mink removal appeared not to affect density estimates of field voles during the winter and summer immediately after food addition. Trapping data from 2004 to 2005 and 2007 suggested, however, that in two out of three summers densities of voles were significantly higher in the absence than in the presence of mink. We conclude that vole populations on small islands in the archipelago of the Baltic Sea are mainly bottom-up limited during winter (outside the growing season of food plants), when food availability is low, and limited by mink predation during summer which slows population growth during the reproductive season of voles.     

Metapopulation processes and persistence in remnant water vole populations

We examined the spatial distribution of water vole populations in four consecutive years and investigated whether the regional population processes of extinction, recolonisation and migration influence distribution and persistence. We examined how such regional processes are influenced by spatial variation in habitat quality. In addition, we assessed the relevance of metapopulation concepts for understanding the dynamics of species that deviate from classical metapopulation assumptions and developing conservation measures for them. Populations were patchy and discrete, and the patchy distribution was not static between years. Population turnover occurred even in the absence of predatory mink, which only influenced the network of populations at the end of the study. Most populations were clustered close together in the upper tributaries. Local population persistence was predominantly influenced by population size: large populations were more persistent. Recolonisation rates were influenced by isolation and habitat quality. The isolation estimates which best explained the distribution of water vole populations incorporated straight‐line distances, suggesting water voles disperse overland. The distribution of recolonised sites indicated that dispersing voles actively selected habitat on the basis of its quality. Water voles depart from some of the assumptions made by frequently used metapopulation models. In particular there is no clear binary distinction between suitable and non‐suitable habitat. Accounting for variation in habitat quality before investigating temporal changes in population distribution allowed us to demonstrate that the key metapopulation processes were important. The significance of regional population processes relative to local population processes may have increased in declining, fragmented populations compared to pristine regional populations. We hypothesise that although mink predation is likely to eventually cause regional extinction in many areas, metapopulation processes have delayed this decline. Consequently, conservation measures should take into account mink predation rates and regional population processes, before considering aspects of habitat quality.     

The stoat Mustela erminea population decline in northern Belarus and its consequences for weasels Mustela nivalis

Abstract

In northern Belarus, we have documented a decline in the local stoat Mustela erminea population following the naturalisation of the American mink Mustela vison. The most likely cause is the reduction in the density and distribution of the main prey of stoats, the riparian voles (the water vole Arvicola terrestris and the root vole Microtus oeconomus), due to excessive predation by mink. Since the stoat population has declined, the number of weasels Mustela nivalis in marshlands has increased and their mean body mass has increased, correlated with the higher number and mean weights of rodents available for weasels in marshland compared with forest habitats.     

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.     

Feeding habits of invasive American mink (Neovison vison) in northern Germany—potential implications for fishery and waterfowl

In order to collect ecological data of invasive American mink (Neovison vison) at a fishpond area in northeastern Germany, we conducted a telemetry study in which 14 mink were radio-tracked. During this project, 2,502 scats from radio-tracked individuals were collected in the period from October 2003 to October 2005. Investigated mink principally prey on fish, small mammals and birds (eggs inclusive), whereas amphibians, reptiles and invertebrates were caught infrequently. Analysing mink scats of different seasons, we found significant seasonal variations of diet composition. In spring, fish, mammals and birds were hunted in similar amounts. During summer, birds made up the main part of the diet followed by mammals. In autumn, the proportion of birds in the mink diet decreased, whereas fish gained in importance. This trend continued during the winter period, when mink preyed almost exclusively on fish. Amphibians, crustaceans, insects, molluscs and reptiles were found only occasionally in scat samples. Among birds, the mink preyed mainly on the Eurasian Coot (Fulica atra) followed by the Mallard (Anas platyrhynchos). Mammalian prey was clearly dominated by the water vole (Arvicola terrestris) and among fish, mink hunted especially perch (Perca fluviatilis), roach (Rutilus rutilus) and carp (Cyprinus carpio). Results clearly demonstrate that mink is an opportunistic predator, which hunts its prey according to availability and vulnerability, respectively. Despite the high portions of fish in their autumn and winter diet, the economic damage caused by mink seems to be negligible. However, high predation rates on birds during the breeding season indicate a potential negative impact of mink on waterfowl.     

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.     

The combined effect of environmental factors and neighbouring populations on the distribution and abundance of Arvicola terrestris. An approach using rule-based models

We investigated how the interplay between environmental factors and presence of neighbouring populations determines the distribution and abundance of a small, endangered rodent, the water vole ( Arvicola terrestris ). We studied thriving and non-fragmented populations of water voles in the absence of their main predator, the introduced American mink ( Mustela vison ). A low degree of population fragmentation, such as the one characterizing the studied populations, was probably typical of water voles before their decline started. We found that under these conditions water voles' distribution is mainly determined by three environmental factors: presence of freshwater, adequate food, and cover. Variance in other factors is well tolerated by water voles. We obtained this result by the use of rule-based models in two separate areas. The two models correctly classified 81% and 83% of the observed cases, respectively. When optimised on one area and cross-validated on the other area the performance of the models was still high (73% and 79%) indicating that the models were robust and generalizable. We also found that the density of animals was lower in sub-optimal than in optimal habitat. We then tested the hypothesis that the number of neighbouring colonies determines the probability of finding voles in a given section. We found that the presence of nearby colonies was an important factor in determining the presence of water voles in sub-optimal habitat, while isolated patches of suitable habitat were less likely to host water voles. These observations suggest the possible presence of a source-sink dynamic, where an optimal habitat acts as a source for populating sub-optimal habitats that may be considered a sink habitat. These findings are discussed in the context of water vole conservation.     

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.     

Habitat utilization by sympatric European mink<Emphasis Type="Italic">Mustela lutreola</Emphasis> and polecats<Emphasis Type="Italic">Mustela putorius</Emphasis> in south-western France

The European minkMustela lutreola Linnaeus, 1761 and the European polecatMustela putorius Linnaeus, 1758 are related species sympatric in southwestern France. The European mink is rapidly disappearing whereas the polecat maintains good populations. Seasonal habitat use of both species was compared in the Landes de Gascogne region to identify if some vulnerability factors of the European mink were associated with habitats occupied by this mustelid. Potential habitats were mapped using a satellite picture and 12 main types of habitats were defined. Animal locations were recorded by radiotracking 9 European mink and 14 polecats from March 1996 to August 1999. Resting animals were located by triangulation, and, when possible, resting places were described. Animals in activity were tracked by continuous monitoring. Data collected revealed a strong preference of European mink for flooded habitats, particularly open marshes, flooded woodlands and moorlands. They seldom left the corridor of the riparian forest and their resting places were mainly in flooded environments, above ground (under herbs or bushes) or in cavities between tree roots. European polecats were less tightly linked to wetlands. Most of their locations were in the pine forests outside the valleys and their resting places were mainly in burrows. The strong specialisation of European mink in aquatic habitats is probably one of the main reasons for its decline because wetlands suffer drastic damages throughout all of its range. Maintaining adequate water levels is crucial for satisfying habitat requirements of mink.     

Variation in the diet composition of a generalist predator, the red fox, in relation to season and density of main prey

Diet composition of a generalist predator, the red fox (Vulpes vulpes) in relation to season (winter or summer) and abundance of multi-annually cyclic voles was studied in western Finland from 1983 to 1995. The proportion of scats (PS; a total of 58 scats) including each food category was calculated for each prey group. Microtus voles (the field vole M. agrestis and the sibling vole M. rossiaemeridionalis) were the main prey group of foxes (PS = 0.55) and they frequently occurred in the scats both in the winter and summer (PSs 0.50 and 0.62, respectively). There was a positive correlation between the PSs of Microtus voles in the winter diet of foxes and the density indices of these voles in the previous autumn. Other microtine rodents (the bank vole Clethrionomys glareolus, the water vole Arvicola terrestris and the muskrat Ondatra zibethicus) were consumed more in winter than in summer. The unusually high small mustelid predation by red foxes (PS = approx. 0.10) in our study area gives qualitative support for the hypothesis on the limiting impact of mammalian predators on least weasel and stoat populations. None of the important prey groups was preyed upon more at low than at high densities of main prey (Microtus voles). This is consistent with the notion that red foxes are generalist predators that tend to opportunistically subsist on many prey groups. Among these prey groups, particularly hares and birds (including grouse), were frequently used as food by foxes.     

The feeding ecology of mink (Mustela vison) in coastal habitat

The diet of a coast-living population of mink was investigated from the scats collected over a three-year period, and compared with information on the availability of principal prey species. Lagomorphs were the single most important prey, and predation upon them matched the abundance of rabbits as determined by monthly counts. Aquatic foraging was particularly important, with rockpool-inhabiting fish accounting for 29–1% occurrence of food items. Fish predation was more pronounced during winter months when lagomorph prey was less available. Crustacean prey, particularly the shore crab, Carcinus maenas , occurred frequently in the diet. Seabirds figured regularly in the diet; these were either taken as carrion from the strand-line or through predation on breeding colonies during the summer months.     

Fox predation on cyclic field vole populations in Britain

The diet of the red fox Vulpes vulpes L. was studied during three winter periods in spruce pklantations in Britain, during which time the cyclic field vole Microtus agrestis L. populations varied in abundance. Field voles and roe deer Capreolus capreolus L. were the two main prey species in the diet of the red fox. The contribution of lagomorphs to fox diet never exceeded 35% and species of small mammal other than field voles were of minor importance. The contribution of field voles was dependent on vole density. The non-linear density dependent relationship with a rather abrupt increase of field voles in fox did when vole density exceeded ca 100 voles ha⁻¹ was consistent with a prey-switching response. The contribution of field voles to fox diet during the low phase of population cycles was lower in Kielder Forest than in other ecosystems with cyclic vole populations. The number of foxes killed annually by forestry rangers was consistent with the evidence from other studies that foxes preying on cyclic small rodents might show a delayed numerical response to changes in vole abundance. Estimates of the maximum predation rate of the fox alone (200–290 voles ha⁻¹ of vole habitat year⁻¹) was well above a previously predicted value for the whole generalist predator community in Kielder Forest. Our data on the functional response of red foxes and estimates of their predation rates suggest that foxes should have a strong stabilising impact on vole populations, yet voles show characteristic 3-4 yr cycles.     

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.     

The impact of the American mink (Neovison vison) on native vertebrates in mountainous streams in Central Spain

The American mink (Neovison vison) is responsible for the widespread decline of its prey species in the regions where it is an invasive species. The current expansion of the mink in the Iberian Peninsula has aroused concern among conservationists about its negative impact on the rich native fauna. However, evidence for this is still scarce, although there are several studies establishing a direct causal relationship between declining native species and the presence of the American mink. Thus, it is important to further investigate the responses of native species to the American mink in several habitats and locations to enhance our knowledge about the patterns of the effect of the mink in Spain, as well as to inform conservation actions. A field study of the impact of the American mink on a mountainous vertebrate community in central Spain is presented. We studied six species: two fish, one amphibian, one bird, and two mammals. The general results showed a species-specific sensitivity to mink presence, with the Mediterranean water shrew (Neomys anomalus) and the southern water vole (Arvicola sapidus) being the most affected because their ranges were significantly decreased after the introduction of the mink. Regarding the other species, neither their abundance nor range was apparently affected by the American mink. The predatory behavior of the mink and interactions with other carnivores could account for these results. These data aid in shedding light about the current impact of the mink on invaded areas of the Iberian Peninsula and highlight the variability of its effects, as well as the urgent need to establish a general program of control of the mink to avoid negative effects upon native prey communities. Furthermore, given the different responses of native species, we propose that measures to protect native species should be based on species-specific goals and attributes.     

The diet of the feral mink (Mustela vison) in southwest Britain

Mink diet in three study areas in southwest Britain was studied by faeces (scat) analysis. In these areas mink ate a wide range of prey, mainly fish, birds and mammals. Proportions of prey in the diet were determined largely by availability, but were also affected by prey behaviour, the physical nature of the habitat and overall prey abundance. Seasonal variation in predation was recorded on moorhens and fish. Changes in rod and net fishing success on the river Teign could not be correlated with the arrival of the mink. Changes in fish populations in Slapton Ley appeared to have causes other than mink. It is concluded that, in the areas studied, domestic stock make up a very small proportion of the mink's diet, and that the wild prey species are not suffering any obvious population depletion.