Tawny owl prey remains indicate differences in the dynamics of coastal and inland vole populations in southern Finland

Some studies suggest that mild winters decrease overwinter survival of small mammals or coincide with decreased cyclicity in vole numbers, whereas other studies suggest non-significant or positive relationships between mild winter conditions and vole population dynamics. We expect for the number of voles to be higher in the rich and low-lying habitats of the coastal areas than in the less fertile areas inland. We assume that this geographical difference in vole abundances is diminished by mild winters especially in low-lying habitats. We examine these relationships by generalized linear mixed models using prey remains of breeding tawny owls Strix aluco as a proxy for the abundance of voles. The higher number of small voles in the coastal area than in the inland area suggest that vole populations were denser in the coastal area. Vole populations of both areas were affected by winter weather conditions particularly in March, but these relationships differed between areas. The mild ends of winter with frequent fluctuations of the ambient temperature around the freezing point (“frost seesaw”) constrained significantly the coastal vole populations, while deep snow cover, in general after hard winters, was followed by significantly lowered number of voles only in the inland populations. Our results suggest that coastal vole populations are more vulnerable to mild winters than inland ones. We also show that tawny owl prey remains can be used in a meaningful way to study vole population dynamics.     

Snow evens fragmentation effects and food determines overwintering success in ground-dwelling voles

Climate instability strongly affects overwintering conditions in organisms living in a strongly seasonal environment and consequently their survival and population dynamics. Food, predation and density effects are also strong during winter, but the effect of fragmentation of ground vegetation on ground-dwelling small mammals is unknown. Here, we report the results of a winter experiment on the effects of habitat fragmentation and food on experimental overwintering populations of bank voles Myodes glareolus. The study was conducted in large outdoor enclosures containing one large, two medium-sized or four small habitat patches or the total enclosure area covered with protective tall-grass habitat. During the stable snow cover of midwinter, only food affected the overwintering success, body condition, trappability and earlier onset of breeding in bank voles. However, after the snow thaw in spring, habitat fragmentation gained importance again, and breeding activities increased the movements of voles in the most fragmented habitat. The use of an open, risky matrix area increased along the habitat fragmentation. Our experiment showed that long-lasting stable snow cover protects overwintering individuals in otherwise exposed and risky ground habitats. We conclude that a stable winter climate and snow cover should even out habitat fragmentation effects on small mammals. However, along prolonged snow-free early winter and in an earlier spring thaw, this means loss of protection by snow cover both in terms of thermoregulation and predation. Thus, habitat cover is important for the survival of small ground-dwelling boreal mammals also during the non-breeding season.     

What triggers facultative winter migration of Great Bustard (Otis tarda) in Central Europe?

The Great Bustard is on the brink of extinction in Central Europe. Its population is known to suffer high mortality during hard winters, particularly when severe weather conditions cause migration. Long-term winter food management in two populations in Germany did not prevent migration events. To identify migration-triggering factors we tested the potential influence of snow, temperature, phase of winter and development of a tradition of migration. Comparing migratory behaviour with long-term local weather records, we found that snow cover is a much stronger trigger for migration than frost and low temperatures. We conclude that snow heavily affects the Great Bustard’s energy balance mediated not only by limited food access but also by the particular properties of its plumage. This could explain migration events despite food availability and is consistent with our results concerning a tendency for females to undertake facultative winter migration more than males. Available data are currently insufficient to confirm or reject the idea that Great Bustard populations develop a tradition of migratory behaviour following a previous winter migration, and we found no evidence for a decrease in the disposition of the Great Bustard to migrate during the course of winter.     

Small rodent winter survival: snow conditions limit access to food resources

1. In Fennoscandia during winter small rodents spend most of their time in the subnivean space, between the snow cover and the ground. The subnivean space is probably not a uniform habitat, but broken into accessible and inaccessible patches by ice covering the vegetation. This might reduce access to otherwise available food resources. 2. To test whether ice formations reduce access to food and thus limit winter survival of small rodents, we conducted an experiment where we increased subnivean space by adding corrugated aluminium sheets on the ground before onset of winter. The sheets prevented ice formation, thus mimicking natural occurring subnivean space, and providing more room for animals living in the subnivean space to forage. 3. During the experiment 142 Microtus oeconomus were passive induced transponder (PIT)-tagged, and a system consisting of fixed tube-shaped antennas and PIT-tag readers were used to provide data to analyse winter survival and individual subnivean space use. The extent of winter grazing was measured after snow melt by examining percentage area grazed. 4. The treatment resulted in increased survival which corresponded well with significantly higher space use and more grazing under the sheets. 5. Females showed a positive correlation between probability of survival and body mass while no such effect was observed in males. 6. The results suggest that the snow cover reduces survival in winter by physically enclosing the vegetation in ice and thus reducing access to otherwise available food resources. The amount of ice and its configuration might vary between years due to changing weather patterns. Our results offer a mechanistic explanation for variations in winter survival and suggest incorporating climate variables in future small rodent models. 7. Directional and long-term changes in climate might result in increased ice formation in the subnivean system. Such deterioration may lead to reduced winter survival and act by stabilizing population dynamics and dampening vole cyclicity.     

Lemming winter habitat choice: a snow-fencing experiment

The insulative value of early and deep winter snow is thought to enhance winter reproduction and survival by arctic lemmings (Lemmus and Dicrostonyx spp). This leads to the general hypothesis that landscapes with persistently low lemming population densities, or low amplitude population fluctuations, have a low proportion of the land base with deep snow. We experimentally tested a component of this hypothesis, that snow depth influences habitat choice, at three Canadian Arctic sites: Bylot Island, Nunavut; Herschel Island, Yukon; Komakuk Beach, Yukon. We used snow fencing to enhance snow depth on 9-ha tundra habitats, and measured the intensity of winter use of these and control areas by counting rodent winter nests in spring. At all three sites, the density of winter nests increased in treated areas compared to control areas after the treatment, and remained higher on treated areas during the treatment. The treatment was relaxed at one site, and winter nest density returned to pre-treatment levels. The rodents’ proportional use of treated areas compared to adjacent control areas increased and remained higher during the treatment. At two of three sites, lemmings and voles showed significant attraction to the areas of deepest snow accumulation closest to the fences. The strength of the treatment effect appeared to depend on how quickly the ground level temperature regime became stable in autumn, coincident with snow depths near the hiemal threshold. Our results provide strong support for the hypothesis that snow depth is a primary determinant of winter habitat choice by tundra lemmings and voles.     

Surviving winter: Food,but not habitat structure,prevents crashes in cyclic vole populations

Vole population cycles are a major force driving boreal ecosystem dynamics in northwestern Eurasia. However, our understanding of the impact of winter on these cycles is increasingly uncertain, especially because climate change is affecting snow predictability, quality, and abundance. We examined the role of winter weather and snow conditions, the lack of suitable habitat structure during freeze‐thaw periods, and the lack of sufficient food as potential causes for winter population crashes. We live‐trapped bank voles Myodes glareolus on 26 plots (0.36 ha each) at two different elevations (representing different winter conditions) in southeast Norway in the winters 2013/2014 and 2014/2015. We carried out two manipulations: supplementing six plots with food to eliminate food limitation and six plots with straw to improve habitat structure and limit the effect of icing in the subnivean space. In the first winter, all bank voles survived well on all plots, whereas in the second winter voles on almost all plots went extinct except for those receiving supplemental food. Survival was highest on the feeding treatment in both winters, whereas improving habitat structure had no effect. We conclude that food limitation was a key factor in causing winter population crashes.     

Fading out of vole and predator cycles?

Northern voles and lemmings are famous for their spectacular multiannual population cycles with high amplitudes. Such cyclic vole populations in Scandinavia have shown an unexpected and marked long-term decline in density since the early 1970s, particularly with a marked shift to lower spring densities in the early 1980s. The vole decline, mainly characterized by a strongly decreased rate of change in numbers over winter, is associated with an increased occurrence of mild and wet winters brought about by a recent change in the North Atlantic Oscillation. This has led to a decrease in winter stability and has shortened the period with protective snow cover, the latter considered as an important prerequisite for the occurrence of multiannual, high-amplitude cycles in vole populations. Although the vole decline is predicted to be negative for predators' reproduction and abundance, empirical data showing this are rare. Here we show that the dynamics of a predator-prey system (Tengmalm's owl, Aegolius funereus, and voles), have in recent years gradually changed from 3-4 yr, high-amplitude cycles towards more or less annual fluctuations only.     

Winter Survival of Wild Turkey Females in Central Minnesota

Abstract: There is interest in expanding eastern wild turkey (Meleagris gallopavo silvestris) populations north of their current range. We hypothesized that winter survival and food availability are primary determinants in setting the northern extent of wild turkey distribution. To test our hypothesis, we translocated wild turkey females north of their present range into central Minnesota, USA, and compared survival in areas with supplemental food in the form of corn food plots versus areas with no supplemental food. During 2 winters with below-average snow, winter survival was higher for females with supplemental food. In one winter with above-average snow depths, survival was extremely low even with supplemental food. Supplemental food could augment survival during mild winters if wildlife managers arrange with farmers to, annually, retain standing corn near roosting habitat, but food plots may only partially offset effects of deep snow. Managers should critically evaluate northern habitats, long-term costs of sustained feeding, and potential outcomes of concentrating animals and introducing wild animals into new ecosystems. Winter survival may delimit the northern range of wild turkeys, though annual survival rates may also be important and need further research.     

Survival Rates of Female Greater Sage-Grouse in Autumn and Winter in Southeastern Oregon

ABSTRACT We estimated survival rates of 135 female greater sage-grouse (Centrocercus urophasianus) on 3 study areas in southeastern Oregon, USA during autumn and winter for 3 years. We used known-fate models in Program MARK to test for differences among study areas and years, investigate the potential influence of weather, and compute estimates of overwinter survival. We found no evidence for differences in survival rates among study areas, which was contrary to our original hypothesis. There also were no declines in survival rates during fall-winter, but survival rates varied among years and time within years. Average survival rate from October through February was 0.456 (SE = 0.062). The coefficient of variation for this estimate was 13.6% indicating good precision in our estimates of survival. We found strong evidence for an effect of weather (i.e., mean daily min. temp, extreme min. temp, snow depth) on bi-weekly survival rates of sage-grouse for 2 of the study areas in one year. Extremely low (<-15°C) temperatures over an 8-week period and accumulation of snow had a negative effect on survival rates during the winter of 1990–1991 on the 2 study areas at the higher (>1,500 m) elevations. In contrast, we found no evidence for an influence of weather on the low-elevation study area or during the winters of 1989–1990 and 1991–1992. Extreme weather during winter can cause lower survival of adult female sage-grouse, so managers should be aware of these potential effects and reduce harvest rates accordingly.     

Does maternal stress influence winter survival of offspring in root voles Microtus oeconomus? A field experiment

Maternal stress can have long‐term adverse consequences on immunocompetence and disease risk of offspring, and winter survival is a crucial demographic parameter in the life‐history of an individual that can substantially affect northern rodent population dynamics. An understanding of the effects of maternal stress on winter survival of offspring may help identify mechanisms driving population fluctuations of northern small mammals. Thus, we assessed the effects of maternal stress, resulting from high population densities, on winter survival of first generation (F₁) and second generation (F₂) in root voles Microtus oeconomus. Replicate high‐ and low‐density enclosed parental populations were established, from which we obtained F₁ generation that were used to establish new enclosed, equal‐density populations. The adults of the high‐density parental populations had higher corticosterone levels, an indication of physiological stress, than did those of the low‐density parental populations. Over‐winter survival of the F₁ generation voles from the low‐density parental populations was greater than that of those from the high‐density parental populations. Over‐winter survival of F₂ generation voles did not differ between the two treatments. Our results suggest that maternal stress affected over‐winter survival of first generations but not second generations. Reduced immunocompetence, resulting from high population density stresses, transferred to offspring may be a factor in annual (winter) population declines. Because the effect is transitory, i.e. immunocompetence of F₂ voles is not affected, reduced immunocompetence resulting from high density stresses would not contribute to lengthy periods of low population densities that are characteristic of multi‐annual population fluctuations.     

Survival of cohorts in a fluctuating population of the vole Microtus townsendii

Survival patterns of cohorts are described during a population cycle of the vole Microtus townsendii near Vancouver, British Columbia, Canada. A two–year live–trapping study on both enclosed and unfenced populations showed that cohorts during the increase phase of growth lived longest and had the best survival. Smaller voles in the peak density spring cohort had poor survival, but survival increased during the peak density summer. Survival of cohorts in the decline phase breeding season was very poor. The suggestions are made that changes in spacing behaviour may cause changes in cohort survival and that the causes of rapid changes in survival need to be determined.     

Effects of ice duration on plankton succession during spring in a shallow polymictic lake

1. Long-term records of air temperature and ice phenology (ice duration), and phyto- and zooplankton time series (1979–1997) were used to study the effects of ice duration on the successional pattern within plankton communities during spring in a shallow polymictic lake. 2. Water temperature in March was significantly lower after cold winters when compared to average or mild winters. Mean water temperature in April was not significantly different after mild, average or cold winters, but showed an overall significant negative correlation with ice duration. 3. Ice duration affected the timing and the magnitude of the peak abundance of diatoms, rotifers and daphnids during spring, but had no direct effects on the timing and maximum of chlorophytes, cryptophytes, cyanobacteria, bosminids and cyclopoid copepods. 4. Plankton groups which appeared first in the seasonal succession (i.e. diatoms, rotifers and daphnids) reached maximum abundance earlier after mild and average winters. The peak abundance of diatoms was negatively correlated with ice duration, whereas that of rotifers and daphnids was independent of the conditions during the preceding winter. 5. Temperature alone was generally a poor predictor of the timing and magnitude of both phyto- and zooplankton maxima. Turbulence may be important in the timing and the magnitude of peaks in diatoms, while total algal biomass was the most important determinant for the timing of the rotifer maximum. The magnitude of the daphnid maxima were significantly influenced by water temperature in March and April, and by rotifer abundance. The magnitude of the bosminid maximum was correlated with food availability and predation, whereas the timing of the maximum was more closely related to water temperature in May. 6. We conclude that, as a result of the low heat storage capacity of shallow lakes, the effects of winter on planktonic communities are short lived, and soon overtaken by the prevailing weather and by biotic interactions.     

Winter regulation of tundra litter carbon and nitrogen dynamics

Mass and nitrogen (N) dynamics of leaf litter measured in Alaskan tussock tundra differed greatly from measurements of these processes made in temperate ecosystems. Nearly all litter mass and N loss occurred during the winter when soils were mostly frozen. Litter lost mass during the first summer, but during the subsequent two summers when biological activity was presumably higher than it is during winter, litter mass remained constant and litter immobilized N. By contrast, litter lost significant mass and N over both winters of measurement. Mass loss and N dynamics were unaffected by microsite variation in soil temperature and moisture. Whether wintertime mass and N loss resulted from biological activity during winter or from physical processes (e.g., fragmentation or leaching) associated with freeze-thaw is unknown, but has implications for how future climate warming will alter carbon (C) and N cycling in tundra. We hypothesize that spring runoff over permafrost as soils melt results in significant losses of C and N from litter, consistent with the observed influx of terrestrial organic matter to tundra lakes and streams after snow melt and the strong N limitation of terrestrial primary production.     

Lower body mass and higher metabolic rate enhance winter survival in root voles,Microtus oeconomus

Although the biological significance of individual variation in physiological traits is widely recognized, studies of their association with fitness in wild populations are surprisingly scarce. We investigated the effect of individual phenotypic variation in body mass, resting (RMR) and peak metabolic rates (PMR) on mortality of the root vole Microtus oeconomus. Body mass and metabolic rates varied significantly among consecutive years and were also age dependent, as individuals born in late summer and autumn were characterized by significantly lower body mass and metabolic rates than animals born earlier. At the beginning of winter voles born in spring and early summer exhibited reduced body mass and metabolic rates, whereas animals born later maintained lower body mass and RMR, which may be interpreted as phenotypic plasticity enhancing the probability of survival. Body mass had no significant effect on vole survival during summer. In contrast, smaller individuals were characterized by lower mortality during early winter, whereas higher body mass was positively associated with survival later in the season. High body‐mass‐corrected RMR positively affected survival in both summer and winter. The effect of PMR was apparent only during winter, though its direction (and correlation with RMR) varied among years. Deep snow cover negatively affected the survival of voles in both early and late winter. Ambient temperature was positively associated with winter survival, except for late winter, when rising temperature caused flooding of vole habitat. We conclude that the lack of consistency in the directionality and strength of the effects of body mass and metabolic rates on winter survival does not undermine their importance, but rather demonstrates the ability of individuals to adjust metabolic rate to changing environmental conditions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 297–309.     

Lagged effects of North Atlantic Oscillation on spittlebug Philaenus spumarius (Homoptera) abundance and survival

The North Atlantic Oscillation (NAO) is a large‐scale pattern of climate variability that has been shown to have important ecological effects on a wide spectrum of taxa. Studies on terrestrial invertebrates are, however, lacking. We studied climate‐connected causes of changes in population sizes in island populations of the spittlebug Philaenus spumarius (L.) (Homoptera). Three populations living in meadows on small Baltic Sea islands were investigated during the years 1970–2005 in Tvärminne archipelago, southern Finland. A separate analysis was done on the effects of NAO and local climate variables on spittlebug survival in 1969–1978, for which survival data existed for two islands. We studied survival at two stages of the life cycle: growth rate from females to next year's instars (probably mostly related to overwintering egg survival), and survival from third instar stage to adult. The latter is connected to mortality caused by desiccation of plants and spittle masses. Higher winter NAO values were consistently associated with smaller population sizes on all three islands. Local climate variables entering the most parsimonious autoregressive models of population abundance were April and May mean temperature, May precipitation, an index of May humidity, and mean temperature of the coldest month of the previous winter. High winter NAO values had a clear negative effect on late instar survival in 1969–1978. Even May–June humidity and mean temperature of the coldest month were associated with late instar survival. The climate variables studied (including NAO) had no effect on the growth rate from females to next year's instars. NAO probably affected the populations primarily in late spring. Cold and snowy winters contribute to later snow melt and greater spring humidity in the meadows. We show that winter NAO has a considerable lagged effect on April and May temperature; even this second lagged effect contributes to differences in humidity. The lagged effect of the winter NAO to spring temperatures covers a large area in northern Europe and has been relatively stationary for 100 years at least in the Baltic area.     

Age variation in a fluctuating population of the common vole

We analysed variation in age in a fluctuating population of the common vole (Microtus arvalis) in southern Moravia, Czech Republic, to test the assumption of the senescence hypothesis that the age of voles increases with increasing population density. Between 1996 and 1998, we monitored the demographic changes by snap-trapping and live-trapping in a field population passing through the increase, peak and decline phase of the population cycle. We used the eye lens mass method to determine the age of snap-trapped animals and those that died in live-traps. The average age of winter males was clearly higher after the peak phase breeding season than before it. No such phase-dependent shift in age, however, was observed in the female component. Male age continued to increase from autumn to spring over the pre-peak winter, and the highest age was in spring of the peak phase year. However, after the peak phase breeding season the highest age was achieved in winter, with the decline phase males during the next spring tending to be younger. The average age of females in spring populations was always lower than in winter populations. The average age of voles from live-traps was always higher than voles from snap-traps, particularly in winter and spring populations, suggesting the presence of senescent animals. Although the density-dependent changes in age are consistent with those observed for other voles, they provide only weak evidence that population cycles in the common vole are accompanied by pronounced shifts in individual age, particularly in female voles.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Does the removal of snowpack and the consequent changes in soil frost affect the physiology of Norway spruce needles?

Climate change may cause a decrease in snow cover in northern latitudes. This, on the other hand, may result in more severe soil frost even in areas where it is not common at present, and may lead to increased stress on the tree canopy. We studied the effects of snow removal and consequent changes in soil frost and water content on the physiology of Norway spruce (Picea abies [L.] Karst.) needles and implications on root biomass. The study was conducted at a 47-year-old Norway spruce stand in eastern Finland during the two winters of 2005/06 and 2006/07. The treatments in three replicates were: (i) natural snow accumulation and melting (CTRL), (ii) artificial snow removal during the winter (OPEN), and (iii) the same as OPEN, but the ground was insulated in early spring to delay soil thawing (FROST). In spite of the deeper soil frost in the OPEN than in the CTRL treatment, soil warming in spring occurred at the same time, whereas soil warming in the FROST was delayed by 2 and 1.5 months in 2006 and 2007, respectively. The soil water content was affected by snow manipulations, being at a lower level in the OPEN and FROST than CTRL in spring and early summer. The physiological measurements of the needles (e.g. starch, carbon and nitrogen content and apoplastic electrical resistance) showed differences between soil frost treatments. The differences were mostly seen between the CTRL and FROST, but also in the case of the starch content in early spring 2007 between the CTRL and OPEN. The needle responses in the FROST were more evident after the colder winter of 2006. The physiological changes seemed to be related to the soil temperature and water content in the early growing season rather than to the wintertime soil temperature. No difference was found in the fine root (diameter < 2 mm) biomass between the treatments assessed in 2007. In the future, conditions similar to the OPEN treatment may be more common than at present in areas experiencing a thick snow cover. The present experiment took place over the course of two years. It is possible that whenever thin snow cover occurs yearly, the reduced starch content during the early spring may be reflected in the tree growth itself as a result of reduced energy reserves.     

Unique population dynamics of Japanese field vole: Winter breeding and summer population decline

It is common knowledge that winter temperatures influence the life history of small mammals. Cold temperatures necessitate increased energy requirements for survival, and recent studies indicate that snow cover can have both negative and positive influences. With each new observation, we develop a more comprehensive understanding of the mechanisms that influence small mammal populations. Here we report on our recent study on Japanese field vole Microtus montebelli, which reaches its peak in population during the early spring and its low during the autumn. To understand the population dynamics of these voles, we conducted a capture-mark-recapture survey, then estimated the seasonal abundance, recruit, capture probabilities, and survival probabilities using the Bayesian hierarchical model. We also analyzed the impact of mammalian generalist predator visits on the survival probabilities. Our data indicates that the early spring peak in population is due to intensive winter breeding and the highest survival probabilities during the periods of deep snow cover. When snow cover reaches a certain depth, the circumstances can combine to raise survival probability and favor breeding. During the breeding season in May and June, on the contrary, the survival probability reached its lowest, resulting in a decrease in population despite breeding. The low survival probability between spring and autumn could be attributed to the impact of generalist predators, and low vegetation may have amplified the effect. In summary, the deep snow cover and generalist predators were considered to be the key factors shaping this unique population dynamics in this orchard area.     

Analysis of spatial and yearly variation in winter survival of winter wheat

 Four years of winter survival data for winter wheat (Triticum aestivum L.) were collected on a loam soil located on the Central Experimental Farm at Ottawa, Ontario (45° 23′N, 75° 43′W). The site was low-lying and subject to frequent winter flooding and ice sheet formation. It appeared level although there was microtopographic variation with a range in elevation of approximately 0.15 m. The objective of the study was to gain insight into factors which might affect winter survival. Two varieties, a soft white and a hard red winter wheat, were planted in September. Crop establishment was measured in late fall and the percentage survival was measured in April of the following year. We assumed the large spatial differences in survival were not totally random, but rather were affected by spatial variation in environmental factors such as snow and ice depth, soil moisture and temperature. Hourly measurements of soil temperature at a depth of 0.05 m were recorded throughout the fall, winter and spring. Fall and spring soil moisture at the same depth were measured on the plot, as well as snow and ice depth at selected times throughout the winter. Measurements were taken on a grid covering the plot to help explain spatial variation in survival. In addition, detailed measurements of elevation were taken on a grid. Meteorological data were available from the nearby weather site. While soil temperatures were never low enough to kill plants according to the CERES model, the spatial variation in winter survival was associated with differences in elevation and the resulting surface drainage patterns. Received: 23 March 1998/Accepted: 21 October 1998     

Variations in Black Grouse Tetrao tetrix winter survival in a year with prolonged snow cover

Capsule Low Black Grouse survival rate in northern England during a severe winter with prolonged snow was attributed to limited availability and proximity of woodland.

Aims To compare the impact of the severe winter weather on Black Grouse in an open treeless landscape in northern England with more wooded landscapes in Scotland.

Methods We assessed the impact of severe winter conditions in relation to previous milder winters at a sample of leks counted annually in northern England and the Scottish Highlands between 2001 and 2012. In addition, following the severe winter in 2009/10 Black Grouse were surveyed at all leks in 2010 where they were surveyed the previous year. We assessed woodland habitat extent within 1 km of leks and related this to indices of Black Grouse over-winter survival.

Results Black Grouse in northern England were severely affected by the severe winter of 2009/10, declining by 38% in comparison to a 23% increase in the Scottish Highlands. Leks in northern England were at higher altitudes and had less tree cover than those in Scotland.

Conclusion The lack of woodland cover around leks in northern England implies that availability of above snow forage, shelter and cover from predators was too low for Black Grouse to survive during prolonged deep snow. Measures are required to provide emergency food sources in periods of prolonged snow, such as the provision of supplementary food at leks, the cultivation of seed-rich arable crops, and the establishment of pockets of woodland.