Habitat requirements of weasels <Emphasis Type="Italic">Mustela nivalis</Emphasis> constrain their impact on prey populations in complex ecosystems of the temperate zone

Differences in habitat use by prey and predator may lead to a shift of occupied niches and affect dynamics of their populations. The weasel Mustela nivalis specializes in hunting rodents, therefore habitat preferences of this predator may have important consequences for the population dynamics of its prey. We investigated habitat selection by weasels in the Bia?owie?a Forest in different seasons at the landscape and local scales, and evaluated possible consequences for the population dynamics of their prey. At the landscape scale, weasels preferred open habitats (both dry and wet) and avoided forest. In open areas they selected habitats with higher prey abundance, except during the low-density phase of the vole cycle, when the distribution of these predators was more uniform. Also in winter, the distribution of weasels at the landscape scale was proportional to available resources. In summer, within open dry and wet habitats, weasels preferred areas characterised by dense vegetation, but avoided poor plant cover. In winter, weasels used wet open areas proportionally to availability of habitats when hunting, but in contrast to summer, they rested only in habitats characterized by a lower water level, which offered better thermal conditions. At the local scale, the abundance of voles was a less important factor affecting the distribution of these predators. Although we were not able to provide direct evidence for the existence of refuges for voles, our results show that they may be located within habitat patches, where availability of dense plant cover and physiological constraints limit the activity of weasels. Our results indicate that in complex ecosystems of the temperate zone, characterized by a mosaic pattern of vegetation types and habitat specific dynamics of rodents, impact of weasels on prey populations might be limited.     

Predation rate, prey preference and predator switching: experiments on voles and weasels

We studied the predation rate and prey selection of the least weasel ( Mustela nivalis nivalis ) on its two most common prey species in boreal environments, the bank vole ( Clethrionomys glareolus ) and the field vole ( Microtus agrestis ), in large outdoor enclosures. We also studied the response of weasels to odours of the two species in the laboratory. The enclosure experiment was conducted using constant vole densities (16 voles/ha) but with varying relative abundance of the two species. Weasels showed higher predation rates on bank voles, and males had higher predation rate than females. Females killed disproportionately more of the more abundant prey species, but they preferred bank voles to field voles when both were equally available. Overall, the predation rate also increased with increasing abundance of bank voles. Therefore our results are in agreement with earlier laboratory results showing preference for bank voles, even if no intrinsic preference for odours of either species was observed in our laboratory study. We suggest that the least weasel hunts according to prey availability, prey aggregation and suitability of hunting habitat, and that this causes the observed dependence of least weasels on field voles and emphasises the role of the field vole in the vole-weasel interaction in cyclic vole populations. Furthermore, our results suggest that predation by weasels may facilitate the coexistence of the two vole species via predator switching, and that it may cause the observed synchrony in dynamics between vole species.     

Food habits of stoatsMustela erminea and weaselsMustela nivalis in Denmark

StoatsMustela erminea Linnaeus, 1758 and weaselsMustela nivalis Linnaeus, 1766 exploit the same array of prey species at different frequencies according to body size. The rabbitOryctolagus cuniculus, which typically is the dominant prey for stoats in temperate parts of Europe, is absent in Denmark. The present study based on gastrointestinal tract contents examines the food habits of sympatric stoats (n=112) and weasels (n=132) from Denmark. Rodents were the most important prey group for both stoats and weasels, constituting 77% and 84% of their diet respectively, expressed as frequency of occurrence. No differences were detected in the proportions of major prey groups between sexes within species, but between species the diets differed. Stoats ate birds and birds’ eggs more often than weasels did, while weasels ate more insectivores. Stoats ate more oftenMicrotus voles and water voleArvicola terrestris than weasels did, while weasels ate more bank voleClethrionomys glareolus and moleTalpa europaea. There was a larger dietary overlap between sympatric stoats and weasels in Denmark than in other dietary studies in areas where rabbits were available.     

Population biology of small mammals in Pureora forest park .1. Carnivores (

Populations of four species of carnivores were sampled over the five years 1983-87 at Pureora Forest Park, by regular three- monthly Fenn trap index lines supplemented with occasional control campaigns by shooting and additional traps. Stoats were the most frequently collected (63 captures), followed by weasels (18), cats (15) and ferrets (13). Stoats ranged throughout the mosaic of forest types but especially the older exotic blocks, hunting rabbits, rats, possums and birds. The mean age of 55 stoats trapped was 15 months, and their maximum life span about 5 years. The age-specific mortality rate of first year stoats was about 0.76, and the proportion of older stoats (>1 year) declined from 52% of 21 killed in summer/autumn of 1983 to 27% of 22 killed in the same seasons of 1984-87. Weasels were collected mainly from habitats favouring mice, such as a young plantation and the road verges, and 40% of 15 non-empty weasel guts contained mice. Cats and ferrets hunted the native forest blocks where their main prey, rats and possums, were abundant. The body sizes and reproductive patterns of mustelids at Pureora were similar to those recorded in podocarp-broadleaf forests elsewhere in New Zealand.     

Food habits of British stoats (Mustela ermined) and weasels (Mustela nivalis)

Information on the food habits of stoats and weasels was obtained chiefly from gut analyses of carcasses sent from various parts of Britain. Lagomorphs, small rodents and birds form the bulk of the food taken by these predators. The three food classes appear to be equally important to stoats, whereas weasels obtain nearly half of their food from small rodents. Microtus was the most important small rodent in the diet of both stoats and weasels. Game birds were taken more by stoats than weasels; the chief bird prey of the latter was passerines.     

Movement patterns of a specialist predator,the weasel<Emphasis Type="Italic">Mustela nivalis</Emphasis> exploiting asynchronous cyclic field vole<Emphasis Type="Italic">Microtus agrestis</Emphasis> populations

We investigated habitat selection and movement characteristics of male weaselsMustela nivalis Linnaeus, 1766 during the breeding season through radio-telemetry in Kielder Forest (KF) in order to assess how weasel movement is influenced by prey dynamics, mate searching and predation risk, and whether the scale of weasel movement corresponds to the spatial scale of the asynchronous, multi-annual vole population cycles observed in KF. Weasels used habitats with a high proportion of grass cover to a much larger extend than habitats with less grass cover and moved through the latter habitats faster and / or straighter. Habitats with high amounts of grass cover also had the highest field vole abundance, although total rodent abundance did not differ between habitats. The selection of this habitat by weasels might reflect weasels preferring field voles as prey or avoiding habitats with little grass cover and high intraguild predation risk. Five out of 8 male weasels radio-tracked had low day-to-day site fidelity and moved between different clear cuts. Three other males were resident in a single clear cut. This variation may reflect mate searching by male weasels. The observation that most weasels (5 out of 8) roamed over large areas and the scale of their dispersal potential suggests, that if they regulated vole populations, they should have a greater synchronising effect on the spatial scale of vole population dynamics than what is observed in vole populations in KF.     

Body size of the weasel Mustela nivalis and the stoat M. erminea in Sweden

In this study we examined temporal and geographical variations in a sample of 124 skulls of the weasel Mustela nivalis and 146 skulls of the stoat M. erminea, collected in Sweden between 1959-1992 and 1913-1990, respectively.We used Principal Component Analysis (PCA) to combine the effects of latitude, longitude, year of collection, mean ambient temperature and Net Primary Productivity (NPP). The first principal component (PC1) contained latitude, ambient temperature and NPP and was significantly and positively related to male (but not female) skull size of both stoats and weasels. None of the other factors or their interactions were significantly related to skull size.We conclude that ambient temperature, either directly through energy savings, or indirectly through improved food availability (increased NPP), had a significant effect on determining body size of male stoats and weasels in Sweden. Our results support the hypothesis that male and female of these species are affected by different selection pressures and thus react differently to changing environmental conditions.     

Dynamics of voles and small mustelids in the taiga landscape of northern Fennoscandia in relation to habitat quality

In the forests of northern Fennoscandia during the I980's, the dynamics of microtine rodents changed from multiannual high amplitude fluctuations (cycles) to, depending on species, fluctuations with a strong seasonal component or fluctuations with smaller amplitude and lower frequency. Microtine and predator data from the Pallasjarvi area, Finnish Lapland, suggest that this transition took place at different rates in different parts of the taiga landscape. Generally, densities in forest habitats have been primarily seasonal since 198S-86. In mesic spruce taiga and in drier forest habitats microtines had a prolonged peak in 1981-83 and a crash in 1984-83. At the timberline, however, microtine populations dropped from peak to low densities already in 1982-83 but the final crash did not occur until spring 1985. The synchronous decrease in microtines densities in all habitat types in 1984-85 coincided with increase in weasel activity. Activity of other carnivores was consistently high in mesic lowland habitats. The data support following three conjectures. 1) Periodic abundance of least weasels is crucial for sustained vole cycles. 2) Predominance of stoats and other generalist predators lead to less regular fluctuations with a strong seasonal component where density declines occur in autumn and early winter. 3) In barren tundra areas, the vegetation cannot sustain high densities of microtines and. consequently, predation is not a necessary condition for population crashes.     

Role of fruits in the diet of small mustelids (<Emphasis Type="Italic">Mustela</Emphasis> sp.) from the western Italian Alps

The food habits of both weasels and stoats (Mustela nivalis and M. erminea) and the occurrence of fruit in their diet were studied by scat analysis in the Gran Paradiso National Park (western Italian Alps). Being impossible to distinguish between stoats and weasels scats, they were grouped together as the genus Mustela. Despite the high consumption of small rodents, which represented the main food item in the diet of small mustelids (41.5% in mean volume), wild fruits (mainly Rosaceae and Ericaceae) were also relatively common (23.3% in mean volume). Fruits were exploited throughout the year, with an intensive use in seasons of high availability. Their consumption did not significantly differ between habitat types, despite a significant decrease of fruit availability from deciduous mixed forest to alpine prairie. This evidence suggests that this trophic behaviour could represent a consolidated feeding strategy of both weasels and stoats, which seem to adopt an opportunistic behaviour as shown by larger size mustelids. We argue that fruits, owing to their good availability and high profitability, could play an important role in the diet of small mustelids in alpine habitats, slackening their well-documented dependence on rodents.     

Distribution and abundance of small mammals in relation to habitat in Pureora Forest Park

Populations of ship rats (Rattus rattus), Norway rats (R. norvegicus), feral house mice (Mus musculus), stoats (Mustela erminea), weasels (M. nivalis), and ferrets (M. furo) were sampled with killtraps every three months from November 1982 to November 1987 in logged and unlogged native forest and in exotic plantations of various ages at Pureora Forest Park, central North Island. Mice (n=522 collected) were fewest in unlogged native forest, more abundant in road edge cutover forest, and most abundant in a young (5-10 year old) plantation. Traps catching most mice were set in dense ground cover under a low, sparse canopy. Ship rats (n=1793) were absent from the young plantation, present but not abundant in older exotic forest, and abundant in all native forest regardless of logging history. Traps set on warmer, steeper sites caught most ship rats, and those set in early successional habitats caught fewest. There was a marked reciprocal relationship between the distributions of ship rats and of mice: the proportion of mice in the total catch of rodents decreased significantly at the least, disturbed forest sites (P <0.001). Most (81%) Norway rats (n=43) were caught in a single trap in unlogged native forest on the bank of a stream. Stoats (n=57) were most abundant in the older exotic plantations; weasels (n=16) in the young plantation and along road edges in native forest; and ferrets (n=11) in unlogged native forest. Hedgehogs (n=290) were common in unlogged native forest far from any roads and also in older exotic forest. Our data suggest that selective logging and conversion to exotics have different effects on each of the six species we monitored. We hypothesise that (1) selective logging is likely to stimulate temporary increases in the numbers of mice and weasels, but not rats or stoats, and (2) after conversion to exotic forest, mice and occasionally weasels will be abundant at first but will gradually be replaced by ship rats and stoats as the forest matures.     

Scats and den contents as indicators of the diet of stoats (Mustela erminea) in the Tasman Valley,South Canterbury,New Zealand

Stoats are significant predators of native fauna in New Zealand. They occur in many habitat types and consume a wide range of prey. The diet of stoats in the Tasman River, South Canterbury, was studied by analysis of scats and den contents. Analysis of 206 scats showed that stoats ate mainly lagomorphs, birds and invertebrates. Minor components included mice, lizards, fish and hedgehogs. Stoats ate more birds in spring than in autumn, and female stoats ate more invertebrates than did males. The contents of 219 dens collected in the same area at the same time provided further information. Birds and lagomorphs occurred at high frequency in dens, and other components were minor. Remains in dens were larger than in scats and allowed identification of many more prey items to species level. Den contents revealed a potentially substantial impact of stoats on threatened shorebirds locally; this impact was not detected by analysis of scats.     

Species‐specific limitation of vole population growth by least weasel predation: facilitation of coexistence?

Interspecific competition is usually understood as different species competing directly with each other for limited resources. However, predators can alter such competitive interactions substantially. Predation can promote the coexistence of species in a situation where it would otherwise be impossible, for example if a tradeoff between the competitive abilities and predation resistance of the prey species exists. The field vole Microtus agrestis and the sibling vole M. rossiaemeridionalis are sympatric grassland species, which compete for the same resources. At the population level sibling voles are suggested to be superior competitors to field voles, yet more vulnerable to predation. We tested the effects of predation on the two species in 0.5 ha outdoor enclosures by exposing vole populations to radio-collared freely-hunting least weasels Mustela nivalis nivalis for three weeks. Lethal and non-lethal impacts of predation limited population densities of both species during and after the experimental period, but the effect was more pronounced in sibling voles in which population densities decreased markedly during the treatment period and even after that. Field vole population densities remained stable under weasel predation, while densities increased in controls. Survival in both species was lower in treatment populations compared to controls, but the effect tended to be more pronounced in sibling voles and in females of both species. The average mass of adults in both species declined in the treatment populations. These results suggest that predation by least weasels can limit vole populations locally, even during favourable summer conditions, and have extended negative effects on the dynamics of vole populations. In addition, predation alleviated interspecific competition between the vole species and is, therefore, a potential factor enabling the coexistence of them.     

Multiple predators induce risk reduction in coexisting vole species

Large predators may affect the hunting efficiency of smaller ones directly by decreasing their numbers, or indirectly by altering their behaviour. Either way this may have positive effects on the density of shared prey. Using large outdoor enclosures, we experimentally studied whether the presence of the Tengmalm's owl Aegolius funereus affects the hunting efficiency of the smallest member of the vole-eating predator guild, the least weasel Mustela nivalis, as measured by population responses of coexisting prey species, the field vole Microtus agrestis and the sibling vole M. levis . We compared the density and survival probability of vole populations exposed to no predation, weasel predation or combined predation by a weasel and an owl. The combined predation of both owl and weasel did not result in obvious changes in the density of sibling and field vole populations compared to the control populations without predators, while predation by least weasel alone decreased the densities of sibling voles and induced a similar trend in field vole densities. Survival of field voles was not affected by predator treatment while sibling vole survival was lower in predator treated populations than in control populations. Our results suggest that weasels are intimidated by avian predators, but without changing the effects of predators on competitive situations between the two vole species. Non-lethal effects of intraguild predation therefore will not necessarily change competitive interactions between shared prey species.     

Responses of stoats and least weasels to fluctuating food abundances: is the low phase of the vole cycle due to mustelid predation?

Summary We studied responses of stoats and least weasels to fluctuating vole abundances during seven winters in western Finland. Density indices of mustelids were derived from snow-tracking, diet composition from scat samples, and vole abundances from snap-trapping. Predation rate was estimated by the ratio of voles to mustelids and by the vole kill rate by predators (density of predator x percentage of voles in the diet). We tested the following four predictions of the hypothesis that small mustelids cause the low phase of the microtine cycle. (1) The densities of predators should lag well behind the prey abundances, as time lags tend to have destabilizing effects. The densities of stoats fluctuated in accordance with the vole abundances, whereas the spring densities of least weasels tracked the vole abundances with a half-year lag and the autumn densities with a 1-year lag. (2) Predators should not shift to alternative prey with declining vole densities. The yearly proportion of Microtus voles (the staple prey) in the diet of stoats varied widely (range 16–82%) and was positively correlated with the winter abundance of these voles. In contrast, the same proportion in the food of least weasels was independent of the vole abundance. (3) The ratio of voles to small mustelids should be smallest in poor vole years and largest in good ones. This was also observed. (4) Vole densities from autumn to spring should decrease more in those winters when vole kill rates are high than when they are low. The data on least weasels agreed with this prediction. Our results from least weasels were consistent with the predictions of the hypothesis, but stoats behaved like semi-generalist predators. Accordingly, declines and lows in the microtine cycle may be due to least weasel predation, but other extrinsic factors may also contribute to crashes.     

Predation on two vole species by a shared predator: antipredatory response and prey preference

In prey communities with shared predators, variation in prey vulnerability is a key factor in shaping community dynamics. Conversely, the hunting efficiency of a predator depends on the prey community structure, preferences of the predator and antipredatory behavioural traits of the prey. We studied experimentally, under seminatural field conditions, the preferences of a predator and the antipredatory responses of prey in a system consisting of two Myodes species of voles, the grey-sided vole (M. rufocanus Sund.) and the bank vole (M. glareolus Schreb.), and their specialist predator, the least weasel (Mustela nivalis nivalis L.). To quantify the preference of the weasels, we developed a new modelling framework that can be used for unbalanced data. The two vole species were hypothesised to have different habitat-dependent vulnerabilities. We created two habitats, open and forest, to provide different escape possibilities for the voles. We found a weak general preference of the weasels for the grey-sided voles over the bank voles, and a somewhat stronger preference specifically in open habitats. The weasels clearly preferred male grey-sided voles over females, whereas in bank voles, there was no difference. The activity of voles changed over time, so that voles increased their movements immediately after weasel introduction, but later adjusted their movements to times of lowered predation risk. Females that were more active had an elevated mortality risk, whereas in the case of males, the result was the opposite. We conclude that, in vulnerability to predation, the species- or habitat-specific characteristics of these prey species are playing a minor role compared to sex-specific characteristics.     

Endozoochorous seed dispersal by sympatric mustelids, Martes melampus and Mustela itatsi, in western Tokyo, central Japan

We investigated seed dispersal by two sympatric mustelid species, the Japanese marten (Martes melampus) and Japanese weasel (Mustela itatsi), along an intercity forest path in western Tokyo, central Japan, from Jul 2007 to Jul 2008. We aimed to investigate the effect of food/habitat preference of these mustelids (martens are semi-arboreal frugivores while weasels are terrestrial carnivores) on their seed dispersal characteristics, which determine their efficacy as seed dispersers. In total, we analyzed 478 fecal samples collected from the two mustelids (N_marten = 381, N_weasel = 97). The proportions of feces containing seeds for martens and weasels were 81.4% and 55.7%, respectively. The number of plant species whose seeds were found within the feces were 28 and 17, respectively. Almost all seeds within feces of both mustelids were intact. The number of plant species whose seeds were found within a single fecal sample ranged from one to four, but no significant difference was detected between the two mustelids. However, marten feces contained a significantly greater number of seeds of most plant species as well as total number of seeds than did weasel feces. The numbers of plant species and seeds represented in marten feces varied seasonally, but those represented in weasel feces did not. Our findings suggest the possibility that both mustelids act in some ways as seed dispersers, although martens seem to disperse a greater diversity and total amount of seeds.     

Habitat use of small mustelids in north Fennoscandian tundra: a test of the hypothesis of patchy exploitation ecosystems

The habitat use of small mustelids in a tundra area in Norwegian Lapland was studied chiefly by means of snow-tracking 1986-89 Stoats showed strong peference to a habitat complex immediately beneath the thrust line of the Scandes, with exceptional abundance of luxuriant habitats, whereas weasel activity was more evenly spread over the lowland tundra Mustelid activity on the high tundra above the thrust line was consistently low Within each subarea. stoat activity was concentrated to the most luxuriant habitats Similar preferences were shown by weasels on the lowland but not in the vicinity of the thurst cliff Daily movements of both species varied from local (c 200 m) to extensive (up to 4 km), no consistent interspecific differences in travel distances could be observed
The results largely conform to the hypothesis of patchy exploitation ecosystems (T Oksanen 1990a), according to which predator activity tends to "spill over' from luxuriant habitats, capable of supporting predator populations, to adjacent barren ones, due to despotic behavior within and between species and due to opportunistic predation by transient predators However, predator activity in barren habitats during the crash winter could not be explained by these mechanisms alone A third mechanism - breakdown of habitat preferences of predators during crash phases of a cyclic prey population - was thus introduced     

Coexistence of Habitat Specialists and Generalists in Metapopulation Models of Multiple-Habitat Landscapes

In coarse-grained environments specialists are generally predicted to dominate. Empirically, however, coexistence with generalists is often observed. We present a simple, but previously unrecognized, mechanism for coexistence of a habitat generalist and a number of habitat specialist species. In our model all species have a metapopulation structure in a landscape consisting of patches of different habitat types, governed by local extinction and colonization. Each specialist is limited to its specific type of habitat. The generalist can use more types of habitat, has a lower local competitive ability but can exploit patches left open by the specialists. Our modeling shows that coexistence is easily possible. The mechanism amounts to a colonization/competition trade-off at the landscape level, where the colonization advantage of the inferior competitor does not arise from a higher colonization rate but from its ability to use more types of habitat. Habitat availability has to be intermediate: when there are few patches of each habitat, only the generalist is able to maintain itself and when there are many patches, high propagule pressure of the specialists excludes the generalist. Habitat selection or temporal variations in relative habitat quality are not necessary for coexistence. Increased niche-width, colonization rate or local competitive ability of the generalist enhances its performance compared to the specialists. Various types of habitat degradation favour generalism. When able to use a broad range of habitats, generalists can generate so much propagule pressure that only a low level of local competitive ability is needed to globally exclude the specialists. Hence, in a reversal of the original problem, the question is why there are so many specialist metapopulations?     

The fleas of a population of weasels in Wytham Woods, Oxford

Weasels were regularly livetrapped, anaesthetized and searched for fleas over two years, in Wytham Woods, near Oxford. Eleven species of fleas were identified among 262 collected. Seven normally occur on small rodents, one on rats, two on moles and one on birds. Fleas were collected on 103 of 338 occasions when an unconscious weasel was examined; on average males carried 0·6 fleas, females 1·5 fleas, but there was no significant difference between the proportion of males and females infested. The species distribution of the fleas on weasels in Wytham suggests that they acquire their fleas mainly from the nests and runways of their prey, which they have hunted through or taken over. Important prey, e.g. birds (22% of prey items identified) with uninviting nests contributed few fleas (2%); non-prey, e.g. moles (0–4% of items) with highly desirable nests, contributed disproportionately many, including 35 fleas (15%) of two species otherwise monoxenous on moles. A similar pattern is shown by the fleas carried by stoats in New Zealand.     

The use of poison eggs for the control of stoats

Abstract

Stoats (Mustela erminea) are an important predator of many forest bird species in New Zealand, and more effective methods for their control are being sought. Stoat control using Fenn traps has been shown to prevent predation on mohua (Mohoua ochrocephala), but this technique is labour‐intensive and costly to use for protection of large areas of habitat. We evaluated 1080 delivered in eggs as a poison for control of stoats. The lethal dose has been determined by captive and field trials, but attempts to implement a large‐scale control operation have given inconclusive results. To clarify the effectiveness of 1080 eggs as a control technique, we carried out further field trials with radio‐tagged stoats in the Makarora Valley. Twenty animals were monitored by radio tracking, and data loggers and video cameras recorded their visits to bait stations. The precise time an individual stoat ate a poison egg could be determined from data logger and video information, and its fate was followed. Sixteen of twenty stoats were killed by 1080 eggs, three died of other causes and one remained alive at the end of the trials.