Winter browsing on Alaska feltleaf willow twigs improves leaf nutritional value for snowshoe hares in summer

In boreal forests, browsing by mammals on winter-dormant twigs increases leaf nitrogen, leaf greenness, and leaf size. This suggests browsing reduces competition among meristems for mineral nutrients, and in particular, competition for nitrogen. Winter browsing also reduces the shoot carbohydrate reserves used by leaves to produce condensed tannin. These effects of winter browsing are predicted to improve the nutritional value of leaves for mammals because they increase the mass of digestible nitrogen in leaves. This hypothesis was tested using Alaska feltleaf willow and the snowshoe hare as the experimental system. Six in vivo indicators of leaf nutritional quality were used to compare leaves from winter-browsed plants with leaves from unbrowsed plants. The indicators used were dry matter intake, nitrogen intake, condensed tannin intake, dry matter digestibility, apparent digestibility of nitrogen and nitrogen retention. The results obtained were in agreement with the above hypothesis. In early summer, at the time snowshoe hares and other northern herbivores reproduce, hares fed leaves from browsed plants consumed more nitrogen, digested more of the nitrogen they consumed, and retained more of the nitrogen they digested than did hares fed leaves from unbrowsed plants. The high nitrogen content and low tannin content of leaves from browsed plants may explain this browsing caused increase in leaf nutritional value. How these positive effects of winter browsing on snowshoe hare nutrition at the time of reproduction might affect hare population dynamics are briefly discussed.     

Multiscale habitat relationships of snowshoe hares (Lepus americanus) in the mixed conifer landscape of the Northern Rockies,USA: Cross‐scale effects of horizontal cover with implications for forest management

Snowshoe hares (Lepus americanus) are an ecologically important herbivore because they modify vegetation through browsing and serve as a prey resource for multiple predators. We implemented a multiscale approach to characterize habitat relationships for snowshoe hares across the mixed conifer landscape of the northern Rocky Mountains, USA. Our objectives were to (1) assess the relationship between horizontal cover and snowshoe hares, (2) estimate how forest metrics vary across the gradient of snowshoe hare use and horizontal cover, and (3) model and map snowshoe hare occupancy and intensity of use. Results indicated that both occupancy and intensity of use by snowshoe hares increased with horizontal cover and that the effect became stronger as intensity of use increased. This underscores the importance of dense horizontal cover to achieve high use, and likely density, of snowshoe hares. Forest structure in areas with high snowshoe hare use and horizontal cover was characterized as multistoried with dense canopy cover and medium‐sized trees (e.g., 12.7–24.4 cm). The abundance of lodgepole pine (Pinus contorta) was associated with snowshoe hare use within a mixed conifer context, and the only species to increase in abundance with horizontal cover was Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa). Our landscape‐level modeling produced similar patterns in that we observed a positive effect of lodgepole pine and horizontal cover on both occupancy and use by snowshoe hares, but we also observed a positive yet parabolic effect of snow depth on snowshoe hare occupancy. This work is among the first to characterize the multiscale habitat relationships of snowshoe hares across a mixed conifer landscape as well as to map their occupancy and intensity of use. Moreover, our results provide stand‐ and landscape‐level insights that directly relate to management agencies, which aids in conservation efforts of snowshoe hares and their associated predators.     

UV-B-induced plant stress as a possible cause of ten-year hare cycles

Predation has been assumed to be a necessary factor in the ten-year population cycle of the snowshoe hare (Lepus americanus) and Canadian lynx (Lynx canadensis). The UV-B-induced plant stress hypothesis, in contrast, predicts that hare performance, especially reproduction, is negatively related to sunspot numbers, because production of UV-B-protective phenolics in food plants in periods of low sunspot activity, when the ozone layer is thin, increases the availability of amino acids and reduces the amount of phenolics that protect against herbivores. In accordance with the UV-B-induced plant stress hypothesis, and despite the absence of predators that have been assumed to be necessary for hare cycles, mountain hare (Lepus timidus) populations in Norway fluctuate in close synchrony with snowshoe hare populations in Alberta and the Yukon, Canada. When adjusting for the phase of the hare cycle, the natality of snowshoe hare in Alberta 1962–1976 was negatively related to sunspot numbers with a time lag of two years. It is concluded that delayed responses to UV-B-induced changes in plant chemistry during the sunspot cycle is a possible cause of ten-year cycles of hares and other herbivores, for example grouse and forest moths.     

Modeling the dynamics of woody plant–herbivore interactions with age-dependent toxicity

In this paper we study the effects that woody plant chemical defenses may have on interactions between boreal hares that in winter feed almost entirely on twigs. We focus particularly on the fact that toxin concentration often varies with the age of twig segments. The model incorporates the fact that the woody internodes of the youngest segments of the twigs of the deciduous angiosperm species that these hares prefer to eat are more defended by toxins than the woody internodes of the older segments that subtend and support the younger segments. Thus, the per capita daily intake of the biomass of the older segments of twigs by hares is much higher than their intake of the biomass of the younger segments of twigs. This age-dependent toxicity of twig segments is modeled using age-structured model equations which are reduced to a system of delay differential equations involving multiple delays in the woody plant-hare dynamics. A novel aspect of the modeling was that it had to account for mortality of non-consumed younger twig segment biomass when older twig biomass was bitten off and consumed. Basic mathematical properties of the model are established together with upper and lower bounds on the solutions. Necessary and sufficient conditions are found for the linear stability of the equilibrium in which the hare is extinct, and sufficient conditions are found for the global stability of this equilibrium. Numerical simulations confirmed the analytical results and demonstrated the existence of limit cycles over ranges of parameters reasonable for hares browsing on woody vegetation in boreal ecosystems. This showed that age dependence in plant chemical defenses has the capacity to cause hare-plant population cycles, a new result.     

Instability of the snowshoe hare and woody plant interaction

Summary We show that Alaskan woody plants respond to browsing in two ways that might destabilize a plantherbivore interaction and account for snowshoe hare population cycles. (1) Browse production of preferred, earliersuccessional woody plants increases in response to moderate levels of browsing. Such yield increases are potentially destabilizing. Later successional woody plants show decreases in yield after moderate browsing, which is consistent with the persistence of snowshoe hares in late successional refuge habitats (Keith 1966, Wolff 1980). (2) Many woody plants are destructively overbrowsed or girdled at the peak of the snowshoe hare cycle. The more palatable and plastic, early to mid successional plants respond by sprouting accompanied by juvenile reversion. Sprouts are markedly less palatable than mature shoots. We show here that sprout palatability and twig biomass are restored in 2–3 years for earlier successional plants, but palatability may not recover for 4–10 years in sprouts of some mid to late successional plants. The decrease in palatability helps to account for the snowshoe hare crash (assuming that damage to more palatable plants is widespread during the peak), and the 2–3 year time lag for recovery of more palatable species could account for (May 1974) the observed 8–11 year period of the hare cycles. Browse yield increases acting during the snowshoe hare population nadir and increase, and sprouting with juvenile reversion acting during the hare peak and decline can in principle account for the oscillatory nature and the observed 8–11 year periodicity of the snowshoe hare cycle.     

A natural feeding experiment on a declining snowshoe hare population

Summary A snowshoe hare (Lepus americanus) population on a 9-ha area was supplied with extra natural food by chopping down large white spruce (Picea glauca) and aspen (Populus tremuloides) trees throughout 3 winters from 1981 to 1984. Hares fed vigorously on the downed trees, but the phase of decline of the ten-year cycle occurred equally on control and experimental grids from 1981–1983, and we could detect no improvement in survival or reproduction on the food area. Growth rates were improved on the food grid during the first winter of the decline (1981–82). We concluded that food shortage is not necessary for the cyclic decline of snowshoe hares in the southern Yukon.     

Survival of dispersing versus philopatric juvenile snowshoe hares: do dispersers die?

We used radio-telemetry to monitor the survival of dispersing and philopatric juvenile snowshoe hares ( Lepus americanus ) in southwestern Yukon Territory, Canada, during a cyclic population increase. Neither 28-d survival nor the proportion of hares surviving to breed differed significantly between juvenile hares that dispersed and those that did not, nor was there a significant relationship between dispersal distance and fate (dead or alive). Our results indicate that the overall survival cost associated with natal dispersal is low for snowshoe hares during the early increase of the hare cycle.     

Snowshoe hare demography during a cyclic population low

1. Snowshoe hare ( Lepus americanus Erxleben) populations were studied in south-west Yukon during the low phase of the 10-year population cycle. Food availability and predator abundance were manipulated in a factorial design to determine the importance of each factor in hare dynamics during this phase.
2. Food was abundant during the low phase, and snowshoe hares were not food limited.
3. Survival of hares was higher than at any other phase of the cycle, and predators were scarce, but >75% of hare deaths resulted from predation.
4. Food addition resulted in higher hare densities and better body condition than on control sites. There were no observable effects of food addition on population rate of increase, recruitment, survival or age structure.
5. Mammalian predator reduction resulted in higher hare densities, higher survival, better body condition and an older age structure. Relative to control populations, recruitment was lower and population rates of increase similar.
6. The joint manipulation of food addition + predator reduction had greater positive effects on hare density and body condition than either single factor manipulation. Survival was better than on control sites, and the age structure was older than on control sites. Population rates of increase were similar, but recruitment was higher on the control areas.
7. We conclude that snowshoe hare dynamics at the low of the cycle are dominated by the interaction of food and predation. Risk of predation also had indirect effects on snowshoe hare age structure and body condition.     

Relationship Between Fecal Pellet Counts and Snowshoe Hare Density in Western Wyoming

ABSTRACT Snowshoe hares (Lepus americanus) are an important prey species for Canada lynx (Lynx canadensis) and are considered critical for lynx population persistence. Determination of snowshoe hare distribution and abundance is needed by land management agencies for lynx conservation. An accepted approach for estimating snowshoe hare abundance is the use of fecal-pellet plot counts. Locally derived regression equations are preferred for accurate calibration of pellet counts to snowshoe hare density due to local differences in pellet deposition and decomposition. We used linear regression to examine correlations between snowshoe hare density, as determined by mark–recapture estimates, and pellet plot counts on both uncleared plots and annually cleared plots on the Bridger-Teton National Forest, western Wyoming, USA. We found significant correlations between snowshoe hare density estimates and fecal pellet counts for both uncleared and annually cleared pellet counts; however, the relationship was stronger (higher r) when using pellet counts from annually cleared plots. In addition, we found that adjusting the buffer size by omitting hard habitat edges (not used by hares) around trapping grids improved correlations between snowshoe hare density and fecal pellet counts for both uncleared plots and annually cleared plots. Though precision is sacrificed when using uncleared plots, they may be useful as a coarse index of habitat use by snowshoe hares. Our derived regression equations may be useful to identify important foraging habitat for Canada lynx in western Wyoming. Land managers responsible for conserving snowshoe hare habitat in western Wyoming may use these equations to monitor changes in hare populations among habitats and during prescribed management actions.     

Novel wildlife in the Arctic: the influence of changing riparian ecosystems and shrub habitat expansion on snowshoe hares

Warming during the 20th century has changed the arctic landscape, including aspects of the hydrology, vegetation, permafrost, and glaciers, but effects on wildlife have been difficult to detect. The primary aim of this study is to examine the physical and biological processes contributing to the expanded riparian habitat and range of snowshoe hares (Lepus americanus) in northern Alaska. We explore linkages between components of the riparian ecosystem in Arctic Alaska since the 1960s, including seasonality of stream flow, air temperature, floodplain shrub habitat, and snowshoe hare distributions. Our analyses show that the peak discharge during spring snowmelt has occurred on average 3.4 days per decade earlier over the last 30 years and has contributed to a longer growing season in floodplain ecosystems. We use empirical correlations between cumulative summer warmth and riparian shrub height to reconstruct annual changes in shrub height from the 1960s to the present. The effects of longer and warmer growing seasons are estimated to have stimulated a 78% increase in the height of riparian shrubs. Earlier spring discharge and the estimated increase in riparian shrub height are consistent with observed riparian shrub expansion in the region. Our browsing measurements show that snowshoe hares require a mean riparian shrub height of at least 1.24–1.36 m, a threshold which our hindcasting indicates was met between 1964 and 1989. This generally coincides with observational evidence we present suggesting that snowshoe hares became established in 1977 or 1978. Warming and expanded shrub habitat is the most plausible reason for recent snowshoe hare establishment in Arctic Alaska. The establishment of snowshoe hares and other shrub herbivores in the Arctic in response to increasing shrub habitat is a contrasting terrestrial counterpart to the decline in marine mammals reliant on decreasing sea ice.     

From process to pattern: how fluctuating predation risk impacts the stress axis of snowshoe hares during the 10-year cycle

Predation is a central organizing process affecting populations and communities. Traditionally, ecologists have focused on the direct effects of predation—the killing of prey. However, predators also have significant sublethal effects on prey populations. We investigated how fluctuating predation risk affected the stress physiology of a cyclic population of snowshoe hares (Lepus americanus) in the Yukon, finding that they are extremely sensitive to the fluctuating risk of predation. In years of high predator numbers, hares had greater plasma cortisol levels at capture, greater fecal cortisol metabolite levels, a greater plasma cortisol response to a hormone challenge, a greater ability to mobilize energy and poorer body condition. These indices of stress had the same pattern within years, during the winter and over the breeding season when the hare:lynx ratio was lowest and the food availability the worst. Previously we have shown that predator-induced maternal stress lowers reproduction and compromises offspring’s stress axis. We propose that predator-induced changes in hare stress physiology affect their demography through negative impacts on reproduction and that the low phase of cyclic populations may be the result of predator-induced maternal stress reducing the fitness of progeny. The hare population cycle has far reaching ramifications on predators, alternate prey, and vegetation. Thus, predation is the predominant organizing process for much of the North American boreal forest community, with its indirect signature—stress in hares—producing a pattern of hormonal changes that provides a sensitive reflection of fluctuating predator pressure that may have long-term demographic consequences.     

Demographic differences in diet breadth of Canada lynx during a fluctuation in prey availability

Population dynamics of specialist carnivores are closely linked to prey availability, but the extent of variability in diet breadth of individual carnivores relative to natural variability in the abundance of their primary prey is not well understood. Canada lynx (Lynx canadensis) specialize on snowshoe hares (Lepus americanus) and exhibit cyclic fluctuations in abundance that lag 1–2 years behind those of snowshoe hares. Declining hare densities spur demographic changes in lynx, but it is unclear whether a corresponding increase in diet breadth occurs: (1) broadly across a lynx population; (2) only among individuals who are able to effectively switch to alternative prey; or (3) only among individuals who cannot capture sufficient primary prey. We measured stable isotope ratios of lynx muscle tissue spanning a cyclic increase and decline in hare density (1998–2001) in Fort Providence, NT, Canada. We found that lynx cohorts responded differently to hare population change, with yearling animals having broader diets at low hare densities, while adults and dependent juveniles maintained a constant diet through the initial decline in hare density. This result was consistent irrespective of lynx sex and indicates that yearling lynx likely are forced to adopt a broader diet when primary prey densities decline. Our results imply that select cohorts of specialist carnivores can exhibit high dietary plasticity in response to changes in primary prey abundance, prompting the need to determine whether increased diet breadth in young lynx is a successful strategy for surviving through periods of snowshoe hare scarcity. In this way, cohort‐specific niche expansion could strongly affect the dynamics of organisms exhibiting population cycles.     

Predator-Mediated Indirect Effects of Snowshoe Hares on Dall's Sheep in Alaska

ABSTRACT Indirect interactions among species can strongly influence population dynamics and community structure but are often overlooked in management of large mammals. We estimated survival of Dall's sheep (Ovis dalli) in the central Alaska Range, USA, during years of differing snowshoe hare (Lepus americanus) abundance to test whether indirect interactions with a cyclic hare population affect Dall's sheep either negatively, by subsidizing predators (apparent competition), or positively, by diverting predation (apparent commensalism). Annual survival of adult female sheep was consistently high (0.85 for all yr and age classes combined). In contrast, annual estimates of lamb survival ranged from 0.15 to 0.63. The main predators of lambs were coyotes (Canis latrans) and golden eagles (Aquila chrysaetos), which rely on hares as their primary food and prey on lambs secondarily. Coyotes and eagles killed 78% of 65 radiocollared lambs for which cause of death was known. Lamb survival was negatively related to hare abundance during the previous year, and lamb survival rates more than doubled when hare abundance declined, supporting the hypothesis of predator-mediated apparent competition between hares and sheep. However, stage-specific predation and delays in predator responses to changes in hare numbers led to a positive relationship between abundance of adult Dall's sheep and hares. Lacking reliable estimates of survival, a manager might erroneously conclude that hares benefit sheep. Thus, support for different indirect effects can be obtained from different types of data, which demonstrates the need to determine the mechanisms that create indirect interactions. Long-term survey data suggest that predation by coyotes is limiting this sheep population below levels typical when coyotes were rare or absent. Understanding the nature of indirect interactions is necessary to effectively manage complex predator–prey communities.     

Conservation implications of the evolutionary history and genetic diversity hotspots of the snowshoe hare

With climate warming, the ranges of many boreal species are expected to shift northward and to fragment in southern peripheral ranges. To understand the conservation implications of losing southern populations, we examined range‐wide genetic diversity of the snowshoe hare (Lepus americanus), an important prey species that drives boreal ecosystem dynamics. We analysed microsatellite (8 loci) and mitochondrial DNA sequence (cytochrome b and control region) variation in almost 1000 snowshoe hares. A hierarchical structure analysis of the microsatellite data suggests initial subdivision in two groups, Boreal and southwestern. The southwestern group further splits into Greater Pacific Northwest and U.S. Rockies. The genealogical information retrieved from mtDNA is congruent with the three highly differentiated and divergent groups of snowshoe hares. These groups can correspond with evolutionarily significant units that might have evolved in separate refugia south and east of the Pleistocene ice sheets. Genetic diversity was highest at mid‐latitudes of the species' range, and genetic uniqueness was greatest in southern populations, consistent with substructuring inferred from both mtDNA and microsatellite analyses at finer levels of analysis. Surprisingly, snowshoe hares in the Greater Pacific Northwest mtDNA lineage were more closely related to black‐tailed jackrabbits (Lepus californicus) than to other snowshoe hares, which may result from secondary introgression or shared ancestral polymorphism. Given the genetic distinctiveness of southern populations and minimal gene flow with their northern neighbours, fragmentation and loss of southern boreal habitats could mean loss of many unique alleles and reduced evolutionary potential.     

ASSESSMENT OF POTENTIAL BIAS WITH SNOWSHOE HARE FECAL PELLET-PLOT COUNTS

Abstract: The fecal pellet-plot method has been used extensively for snowshoe hare (Lepus americanus) population studies across the species' range, but potential biases associated with the technique have not been addressed adequately. We studied hare pellet-plots in northern Idaho to quantify pellet decomposition rates across environmental gradients, and conducted feeding trials on captive hares to assess the role of diet on pellet production rates. We found that across our study area pellet numbers tended to be higher on plots with high vegetative cover, which likely was a reflection of hare habitat choice rather than lesser pellet decomposition in such habitat. A pellet decomposition experiment indicated that pellet persistence was negatively related to moisture level, and that pellets produced by hares during summer decomposed more quickly than those from winter. We found that only 19% of fecal pellets collected from plots located across northern Idaho were produced by hares during winter. There was a correlation between pellet numbers from plots that were pre-cleared 1 year earlier and estimated numbers of hares on 6 study areas. A similar correlation was lacking for pellet counts from uncleared plots, implying that hare population estimation via pellet-plot counts should involve plot pre-clearing. In captive studies, juvenile hares produced slightly fewer pellets per day per gram of food ingested than adults, but pellet production was similar across diets comprised of 10 different browse species. We conclude that for our study area the fecal pellet-plot method may be subject to notable pellet decomposition bias, and therefore recommend that use of the method elsewhere across the species' range be preceded by assessment of both the pellet-hare density relationship and pellet decomposition rates across habitats.     

Influence of forest structure on the abundance of snowshoe hares in western Wyoming

Snowshoe hares (Lepus americanus) are a primary prey species for Canada lynx (Lynx canadensis) in western North America. Lynx management plans require knowledge of potential prey distribution and abundance in the western United States. Whether even-aged regenerating forests or multi-storied forests contain more snowshoe hares is currently unknown. During 2006–2008, we estimated snowshoe hare density in 3 classes of 30–70-year-old lodgepole pine (Pinus contorta) and 4 classes of late seral multi-storied forest with a spruce (Picea engelmannii)-fir (Abies lasiocarpa) component in the Bridger-Teton National Forest, Wyoming. We recorded physiographic variables and forest structure characteristics to understand how these factors influence abundance of snowshoe hares. In many instances, snowshoe hares were more abundant in late seral multi-storied forests than regenerating even-aged forests. Forest attributes predicting hare abundance were often more prevalent in multi-storied forests. Late seral multi-storied forests with a spruce–fir component and dense horizontal cover, as well as 30–70-year-old lodgepole pine with high stem density, were disproportionately influential in explaining snowshoe hare densities in western Wyoming. In order to promote improved habitat conditions for snowshoe hares in this region, management agencies should consider shifting their focus towards maintaining, enhancing, and promoting multi-storied forests with dense horizontal cover, as well as developing 30–70-year-old lodgepole pine stands with high stem density that structurally mimic multi-storied forests. © 2012 The Wildlife Society.     

Owl predation on snowshoe hares: consequences of antipredator behaviour

We show evidence of differential predation on snowshoe hares (Lepus americanus) by great horned owls (Bubo virginianus) and ask whether predation mortality is related to antipredator behaviour in prey. We predicted higher predation on (1) young and inexperienced hares, (2) hares in open habitats lacking cover to protect from owl predation, and (3) hares in above average condition assuming that rich food patches are under highest risk of predation. Information on killed hares was obtained at nest sites of owls and by monitoring hares using radio-telemetry. The availability of age classes within the hare population was established from live-trapping and field data on reproduction and survival. Great horned owls preferred juvenile over adult hares. Juveniles were more vulnerable to owl predation before rather than after dispersal, suggesting that displacement or increased mobility were not causes for this increased mortality. Owls killed ratio-collared hares more often in open than in closed forest types, and they avoided or had less hunting success in habitats with dense shrub cover. Also, owls took hares in above average condition, although it is unclear whether samples from early spring are representative for other seasons. In conclusion, these results are consistent with the hypothesis that variation in antipredator behaviours of snowshoe hares leads to differential predation by great horned owls.     

Phenolic compounds of willow bark as deterrents against feeding by mountain hare

Summary Overwintering mountain hares (Lepus timidus) fed selectively on the shoots of a number of northern willow (Salix spp.) species. The hares preferred certain species over others and mature shoots over juvenile ones. There was a negative correlation between the phenolic glycoside concentration in the twigs and the hares' feeding. This correlation was substantiated by feeding experiments in which oat grains treated with purified phenoglycoside and catechin fractions of willow bark were offered along with untreated control oat grains to free-ranging mountain hares. Both fractions in concentrations normally found in willow twigs inhibited hare feeding. The results suggest that these phenolic compounds play a crucial role in the building up of resistance patterns among the willows. The decline in resistance in mature shoots of tall willows indicates that the juvenile resistance can be, perhaps secondarily, an adaptation against mammals browsing from ground level. Accordingly, low willow species retain a high level of resistance also in maturity.     

Winter Prey Selection of Canada Lynx in Northwestern Montana

ABSTRACT The roles that diet and prey abundance play in habitat selection of Canada lynx (Lynx canadensis) in the contiguous United States is poorly understood. From 1998–2002, we back-tracked radiocollared lynx (6 F, 9 M) for a distance of 582 km and we located 86 kills in northwestern Montana, USA. Lynx preyed on 7 species that included blue grouse (Dendragapus obscurus), spruce grouse (Canachites canadensis), northern flying squirrel (Glaucomys sabrinus), red squirrel (Tamiasciurus hudsonicus), snowshoe hare (Lepus americanus), least weasel (Mustela nivalis), and white-tailed deer (Odocoileus virginianus). Snowshoe hares (69 kills) accounted for 96% (4-yr average, range = 94–99%) of prey biomass during the sample period. Red squirrels were the second-most-common prey (11 kills), but they only provided 2% biomass of the winter diet. Red squirrels contributed little to the lynx diet despite low hare densities. A logistic regression model of snowshoe hare, red squirrel, and grouse abundance, as indexed by the number of track crossings of use and available lynx back-tracks, was a significant (Wald statistic = 19.03, df = 3, P < 0.001) predictor of habitat use. As we expected, lynx (P < 0.001) selected use-areas with higher snowshoe hare abundance compared to random expectation. However, the red squirrel index had a weak (P = 0.087) negative relationship to lynx use, and grouse was nonsignificant (P = 0.432). Our results indicate that lynx in western Montana prey almost exclusively on snowshoe hares during the winter with little use of alternative prey. Thus, reductions in horizontal cover for hares would degrade lynx habitat.