Trade-offs between activity and thermoregulation in a small carnivore,the least weasel Mustela nivalis

We studied factors influencing daily energy expenditures (DEE) of male least weasels (Mustela nivalis) using the doubly labelled water technique. The relationship between ambient temperature and DEE formed a triangular pattern, characterized by invariance of the maximum DEE and an inverse relationship between minimum DEE and temperature. A simple energetic model relating the DEE of male weasels to activity time (AT) and ambient temperature predicted that, across seasons, less than 10 per cent of measurements approach the upper bound of observed DEE. Male weasels were able to maintain a relatively constant maximum energy output across varying temperatures by adjusting their AT to changes in temperature. They achieved maximum energy expenditures in winter due to high thermoregulatory costs, and in spring and summer due to high levels of physical activity. This pattern exemplifies a ‘metabolic niche’ of a small mammal having extremely high energy expenditures primarily driven by ambient temperature.     

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.     

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.     

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.     

No evidence of past bottlenecks in two Danish mustelids: results of craniometric and genetic studies in time and space

A craniometric and molecular genetic investigation was conducted in Danish stoat (Mustela erminea) and weasel (Mustela nivalis) populations. Specimens used were collected over a wide time span (stoat: 1864–2002; weasel: 1863–1990) and from several geographical regions (Jutland peninsula and the two islands of Funen and Zealand). The study was made with a temporal and a spatial perspective, allowing the estimation of differences in genetic diversity and craniometrical trait means between geographical regions and through time with the use of ancient DNA techniques. Univariate statistics of 11 trait lengths did not reveal geographical differentiation in size and shape among the different regions for the stoat, but a geographical differentiation in shape was found for the weasel. There was evidence for reductions in skull size with the year of collection in male stoats, but not in females, which suggests that some selective pressures or environmental factors have affected male stoats to a greater extent than female stoats and the weasel. Relatively high values of heterozygosity were found in both the stoat and weasel, using microsatellite markers. The level of genetic variability of the stoat collected recently was compared with the level of genetic variability in the historical samples, demonstrating that the stoat has not suffered severe loss of genetic variability through the investigated period. A comparison of recent and historical genetic variability of the weasel was not possible because the ancient DNA extracted from the weasels was too degraded. Pairwise F_ST values and assignment tests showed small but significant genetic differentiation between the different geographical regions for both the stoat and weasel. No genetic differentiation between the recent and historical samples of the stoat was found. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 88 , 541–553.     

A population study of the metastrongylid nematode Skrjabingylus nasicola in the weasel Mustela nivalis

Analyses of adult worms of Skrjabingylus nasicola occurring in the nasal sinuses of the weasel Mustela nivalis from three localities in the British Isles show that the intensity of infestation can be correlated with increasing severity of skull damage, which in turn increases as the weasel ages. It is suggested that, apart from hormonal differences, the smaller sized female skull is likely to be linked with lower levels of infestation in female compared with male weasels. Also a reduction in mean worm length in female skulls and in heavy infestations of S. nasicola is probably the result of a "crowding effect", previously described for other helminth infections in the definitive host. Crowding appears to have no effect on the sex ratio of worm populations, although there is a tendency for the ratio of female: male worms to increase in female weasels. Reasons for this are discussed.     

Variation in body size, sexual dimorphism and age-specific survival in stoats, Mustela erminea (Mammalia: Garnivora), with fluctuating food supplies

Most hypotheses attempting to explain the evolution of pronounced sexual dimorphism in body size in the three species of weasels (Mustela erminea, M. frenata, M. nivalis) assume that sexual dimorphism is a long-term adaptation, associated with the different reproductive strategies of the two sexes. We here examine an auxiliary hypothesis which predicts that the degree of sexual dimorphism may also vary over the short-term, because when food is temporarily abundant, sexual selection should favour a greater growth rate of males than of females. This hypothesis concerns a phenotypic response which could introduce temporarily increased variation into an existing genotypic trait. We document the present size and sexual dimorphism of stoats introduced last century to New Zealand from Britain in relation to between-year variation in food supply in a single habitat (forests of southern beech, Nothofagus sp.). Southern beech trees produce heavy crops of flowers and seed at 3–5 year intervals, which are associated with very variable supplies of important prey of stoats, including several species of seed-eating birds, litter-feeding insects, and feral house mice (Mus musculus). Alternative prey are scarce. Regressions of condylobasal length and head-body length on mouse population indices were significant in both sexes. Mean condylobasal length was larger in both male and female stoats born after a heavy seedfall compared with those born in non-seedfall years. However, the largest males born in years of heavy seedfall were removed by selective mortality before the age of 3 years, so the condylobasal lengths for old (≥ 3.0 yr) males converged on a common mean regardless of food supply in their birth year. Sexual dimorphism did not vary with food supplies (as reflected in seedfall records or mouse population indices) at any age. First-year survivorship, at least from the age of independence, was significantly negatively correlated with density of stoats in the summer of their birth year.     

Habitat selection of coexisting competitors: a study of small mustelids in northern Norway

The coexistence of two very similar species, stoat and least weasel, has puzzled many researchers. From their ecology it is expected that they do not coexist, not locally at least, and still they seem to do. We reviewed the specific hypotheses proposed to explain their coexistence and related these to general theories of competitive coexistence. To test these conjectures, we studied the habitat selection of least weasels and stoats on landscape and on local scale. The study was performed during the winters, the most critical season, in years 1986–2001 in northern Norway. Stoats were usually more numerous than least weasels. Stoats showed preference for productive areas both at the landscape and at the habitat scale and appeared stereotypic in their habitat selection. Least weasels were more generalized and flexible in their habitat selection. Contrary to results reported in many studies, least weasel did not react to the presence of stoats and were not excluded from the areas with stoats. We suggest that in the conditions of northernmost Fennoscandia, the two species exhibit a variant of classical competitive coexistence. Both species have a shared preference for rodents, but the access to exclusive alternative prey in stoats allows their coexistence with least weasels, which are more efficient predators on rodents. We suggest that more attention should be paid on survival resources, exploited during times of low resource density, when studying the coexistence between close competitors.     

Pandora’s box down-under: origins and numbers of mustelids transported to New Zealand for biological control of rabbits

This paper describes one of the world’s first large-scale experiments in biological control of a major vertebrate pest of agriculture, which was tried in New Zealand during the second half of the nineteenth century. Starting from the late 1860s, pasture damage in Southland and Otago by European rabbits was causing serious reductions in productivity of sheep (wool clip and lambing percentages) associated with malnutrition of the breeding ewes, and a consequent decline in the value of pastoral land. In response, and despite repeated local and international warnings, ferrets, stoats and weasels (Mustela furo, M. erminea and M. nivalis) were liberated on the worst of the rabbit-infested pastures. They were perceived as the ‘natural enemies of the rabbit’ but (unlike foxes) too small to threaten lambs. Over the 50 years after 1870, upwards of 75,000 ferrets, most imported from Australia or locally bred, were released in the South Island. Over the decade 1883–1892, at least 7838 stoats and weasels arrived from Britain. At least 25 shipments are known, with an average of only 10% mortality per shipment. Of the 3585 animals listed by species, 73% were weasels. The total cost of the ferret programme cannot now be estimated; that of stoats and weasels alone was at least £5441, probably twice that, or >$NZ 1–2 million in today’s money. Mustelids (and cats) killed many young rabbits, which was helpful because rates of change in rabbit populations are sensitive to variations in juvenile mortality, but in the most rabbit-prone semi-arid lands, mustelids could not remove enough rabbits to prevent the continuing damage to sheep pastures. The era of deliberate introductions of mustelids to control rabbits in New Zealand was short, expensive, and unsuccessful.     

Population biology of the weasel Mustela nivalis on British game estates

The mean year-class ratio in 6 samples of weasels (total n = 477) ranged from 59% to 84% young, and mean age from 0.79 to 1.16 yr. There was no difference in age structure between samples from 5 game estates and 1 reserve. Mean annual mortality of weasels of both sexes and all ages was 75-90%. In 12 pregnant weasels the mean number of embryos, observed from April to June, was 5.6. There was a definite anoestrous season in winter. Records of the number of weasels killed on game estates show that weasels and stoats reacted differently to the first outbreak of myxomatosis in England in 1954; and that weasel populations are capable of enormous sudden increases (usually associated with "mouse years"). These increases are probably due largely to summer litters produced by precocious juveniles, and also to some adults breeding a second time (both possibilities confirmed from ovarian histology on this material). Traditional gamekeeping practice may influence the seasonal pattern of mortality of weasels (highest in spring) but its effects are probably short-lived.     

Estimating net primary production of boreal forests in Finland and Sweden from field data and remote sensing

We calculated annual mean stem volume increment (AMSVI) and total litter fall to produce forest net primary production (NPP) maps at 1‐km² and half‐degree resolutions in Finland and Sweden. We used a multi‐scale methodology to link field inventory data reported at plot and forestry district levels through a remotely sensed total plant biomass map derived from 1‐km² AVHRR image. Total litter fall was estimated as function of elevation and latitude. Leaf litter fall, a surrogate for fine root production, was estimated from total litter fall by forest type. The gridded NPP estimates agreed well with previously reported NPP values, based on point measurements. Regional NPP increases from northeast to southwest. It is positively related to annual mean temperature and annual mean total precipitation (strongly correlated with temperature) and is negatively related to elevation at broad scale. Total NPP (TNPP) values for representative cells selected based on three criteria were highly correlated with simulated values from a process‐based model (CEVSA) at 0.5°× 0.5° resolution. At 1‐km² resolution, mean above‐ground NPP in the region was 408 g/m²/yr ranging from 172 to 1091 (standard deviation (SD) = 134). Mean TNPP was 563 (252 to 1426, SD = 176). Ranges and SD were reduced while the mean values of the estimated NPP stayed almost constant as cell size increased from 1‐km² to 0.5°× 0.5°, as expected. Nordic boreal forests seem to have lower productivity among the world boreal forests.     

Testing Bergmann’s rule and the Rosenzweig hypothesis with craniometric studies of the South American sea lion

We tested the validity of Bergmann’s rule and Rosenzweig’s hypothesis through an analysis of the geographical variation of the skull size of Otaria flavescens along the entire distribution range of the species (except Brazil). We quantified the sizes of 606 adult South American sea lion skulls measured in seven localities of Peru, Chile, Uruguay, Argentina, and the Falkland/Malvinas Islands. Geographical and environmental variables included latitude, longitude, and monthly minimum, maximum, and mean air and ocean temperatures. We also included information on fish landings as a proxy for productivity. Males showed a positive relationship between condylobasal length (CBL) and latitude, and between CBL and the six temperature variables. By contrast, females showed a negative relationship between CBL and the same variables. Finally, female skull size showed a significant and positive correlation with fish landings, while males did not show any relationship with this variable. The body size of males conformed to Bergmann’s rule, with larger individuals found in southern localities of South America. Females followed the converse of Bergmann’s rule at the intraspecific level, but showed a positive relationship with the proxy for productivity, thus supporting Rosenzweig’s hypothesis. Differences in the factors that drive body size in females and males may be explained by their different life-history strategies. Our analyses demonstrate that latitude and temperature are not the only factors that explain spatial variation in body size: others such as food availability are also important for explaining the ecogeographical patterns found in O. flavescens.     

Cranial variation in British mustelids

Nineteen measurements were made on 136 skulls belonging to seven mustelid species: Meles meles (Eurasian badger), Mustela nivalis, (weasel), Mustela erminea (stoat), Mustela putorius (polecat), Lutra lutra (otter), Mustela furo (ferret), and Mustela vison (American mink), and polecat-ferret hybrids. To investigate shape, size-related effects were eliminated by dividing all measurements by their geometric means. Canonical variate analysis was used to reveal major interspecies distinctions. Excluding the ferrets and polecat-ferrets from the analysis, only 3.2% of the skulls misclassified (one mink, one weasel, and two stoats). Three groups separated on the first canonical axis: 1) badgers, 2) polecats, mink, and otters, and 3) stoats and weasels. The important variables were width of zygomatic arch and height of sagittal crest opposed to the postorbital distance, condylobasal length, and basilar length. Otters separated out on the second canonical axis; the most important variables were postorbital breadth and width of the postorbital constriction opposed to the basioccipital width. There was reasonable separation of polecats from mink on a combination of the second and third canonical axes. On the latter the most important variables were postorbital breadth opposed to postorbital distance. Addition of the ferret data showed that they lay closest to, and overlapped with, the polecats. The stoat and weasel data alone gave complete separation, with height of sagittal crest and width of zygomatic arch opposed to basioccipital width. However, using size-in data the best separation was the relationship between postorbital breadth and either basioccipital width or postorbital distance. Sexual dimorphism was demonstrated in the skulls of badgers but was shown to be relatively insignificant when compared to the interspecific differences.     

Body size of the red fox Vulpes vulpes in Spain: the effect of agriculture

The body size of animals is affected by several factors, including ambient temperature and food availability. Ambient temperature is often negatively related to body size (Bergmann's rule) whereas an improved diet, especially during growth, has a positive effect. Animals commensal with man commonly exploit additional food sources (e.g. garbage dumps), thereby increasing their food supply. Using museum material, we studied morphological variation in skull size (and thus body size) among Spanish red foxes. Four measurements were taken of each skull and were related to the habitat from which the foxes were collected (agricultural and non-agricultural), and to latitude as a proxy for ambient temperature. The skull size of foxes collected in agricultural areas during the late 20th Century was significantly larger than that of those from non-agricultural areas, and was negatively related to latitude, thus contradicting Bergmann's rule. We suggest that increased food availability from animal husbandry is the cause for the observed increase in skull size (and thus body size).  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 729–734.     

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.     

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.     

Patterns in body size and melanism along a latitudinal cline in the wingless grasshopper,Phaulacridium vittatum

Aim We explore geographic variation in body size within the wingless grasshopper, Phaulacridium vittatum, along a latitudinal gradient, and ask whether melanism can help explain the existence of clinal variation. We test the hypotheses that both male and female grasshoppers will be larger and lighter in colour at lower latitudes, and that reflectance and size will be positively correlated, as predicted by biophysical theory. We then test the hypothesis that variability in size and reflectance is thermally driven, by assessing correlations with temperature and other climatic variables. Location Sixty‐one populations were sampled along the east coast of Australia between latitudes 27.63° S and 43.10° S, at elevations ranging from 10 to 2000 m a.s.l. Methods Average reflectance was used as a measure of melanism and femur length as an index of body size for 198 adult grasshoppers. Climate variables were generated by BIOCLIM for each collection locality. Hierarchical partitioning was used to identify those variables with the most independent influence on grasshopper size and reflectance. Results Overall, there was no simple relationship between size and latitude in P. vittatum. Female body size decreased significantly with latitude, while male body size was largest at intermediate latitudes. Rainfall was the most important climatic variable associated with body size of both males and females. Female body size was also associated with radiation seasonality and male body size with reflectance. The reflectance of females was not correlated with latitude or body size, while male reflectance was significantly higher at intermediate latitudes and positively correlated with body size. Analyses of climate variables showed no significant association with male reflectance, while female reflectance was significantly related to the mean temperature of the driest quarter. Main conclusions Geographic variation in the body size of the wingless grasshopper is best explained in terms of rainfall and radiation seasonality, rather than temperature. However, melanism is also a significant influence on body size in male grasshoppers, suggesting that thermal fitness does play a role in determining adaptive responses to local conditions in this sex.     

Why do European stoats Mustela erminea not follow Bergmann's rule?

Stoat Mustela erminea body size shows remarkably great variation over the species' European range. The pattern of this variation is opposite to that suggested by Bergmann's rule, i.e., stoats in central Europe are significantly larger than those in southern Sweden which, in turn are considerably larger than their conspecifics in northern Sweden. Neither winter temperature nor the length of snow cover shows any consistent correlation with stoat body size variation, and the larger body size of the southern populations could not be related to a widening of the stoat's feeding niche. A positive correlation was found between the frequency distribution, by size, of available prey and stoat body size in the different areas examined. This supports the hypothesis that body size variation in the stoat, especially females, is an adjustment to regional variations in the sizes of their available prey.     

Geographic variation in size and sexual dimorphism of North American weasels

Geographic variation in size (skull length) and sexual dimorphism in Mustela erminea, Mustela frenata and Mustela nivalis in North America is described and analysed in relation to latitude, longitude, climatic variables, and sympatry or allopatry of these species. Only erminea increases in size with latitude; it does so regardless of the presence or absence of frenata or nivalis. Latitude is a better predictor of size in erminea than available measures of climate, seasonality or prey size. There is no evidence for character displacement between any pair of species. The sexes covary in size in frenata and erminea , and probably in nivalis , although geographic variation in sexual dimorphism occurs in frenata and erminea. The principal cause of sexual dimorphism appears to be sexual selection for large size in males rather than the high energetic requirements resulting from an elongate body shape. However, prey size may constrain female size (and possibly also male size). Regional differences in the abundance of prey during the growth of young weasels may affect adult size much more in males than in females and contribute to geographic variation in sexual dimorphism.