Edge Effects and Isolation: Red-Backed Voles on Forest Remnants

Negative effects of habitat edge have been advanced as an important proximate cause of extinction, and a growing literature calls attention to the matrix surrounding habitat remnants as a critical factor determining population persistence. I examined spatial distribution of California red-backed voles ( Clethrionomys californicus ) on 13 forest remnants and five control sites in southwestern Oregon. The species was virtually isolated on remnants, making little use of the regenerating clearcuts surrounding the remnants. The effects of the clearcut also impinged on the remnants as edge effects: six times more voles were captured per trap in the interior of remnants than on the edge. Consequently, the density of voles per unit area on remnants increased with remnant size, despite the potential buildup of population density in small isolates due to limited emigration. I explored potential mechanisms of the negative edge effect on voles and found that the biomass of coarse woody debris, per se, did not explain the vole distribution because both number and volume of logs increased from the interior to the edge of remnants. However, the distribution of the vole's primary food item, hypogeous sporocarps of mycorrhizal fungi, did correspond to the vole edge effect     

Effects of Habitat Loss and Fragmentation on the Behavior and Demography of Gray-Tailed Voles

We monitored the short term behavioral and demographic responses of gray-tailed voles (Microtus canicaudus) to the reduction and fragmentation of their habitat. Our objectives were (1) to test whether animals perished or moved into remaining fragments after 70% of their habitat was removed; and (2) to test the null hypothesis that the social structure and demography of animals would not differ between habitats consisting of one large continuous fragment (625 m²), a mosaic of 25 small fragments (each 25 m²) separated by 4 m of bare ground, and control, unmanipulated habitats (1850 m²). We conducted the experiment in 12, 0.2-ha enclosures planted with alfalfa with four replicates for each of two manipulated treatments and a control. A 70% reduction in habitat did not adversely affect adult survival, reproductive rate, juvenile recruitment, or population size. However, an influx of unrelated females into habitat fragments resulted in decreased juvenile recruitment in those fragments. Voles from cleared habitat moved into the remaining habitat and did not measurably affect the resident population. Similarly, the demography of voles did not differ significantly among the large-fragment, small-fragment, and control enclosures. Peak density estimates based on the amount of habitat in each enclosure were 545 animals per hectare in control, 1056 in large-fragment, and 2880 in small-fragment enclosures. Reduced movement of animals among the small fragments was the most obvious effect of habitat fragmentation. Six percent of females and 15% of males moved among small fragments within a week compared to approximately 60% moving comparable distances in large-fragment and control enclosures. Rates of juvenile dispersal and sexual maturation declined throughout the summer on all treatments, were associated with season and density, and were only marginally associated with habitat loss and fragmentation. We conclude that at the time of habitat removal and fragmentation, populations were small enough to accommodate a 70% reduction in habitat and still continue to increase in numbers. The social system of gray-tailed voles was sufficiently flexible to accommodate an influx of animals to withstand densities> 1000 voles per ha. The behavioral and demographic features of gray-tailed voles are similar to those reported for other small mammals, thus confirming the use of voles for ecological model systems in habitat fragmentation studies.     

Spatiotemporal dynamics of habitat use by koalas: the checkerboard model

Animal movement patterns and use of space depend upon food and nonfood resources, as well as conspecific and heterospecific interactions, but models of habitat use often neglect to examine multiple factors and rarely include marsupials. We studied habitat use in an Australian population of koalas (Phascolarctos cinereus) over a 6-year period in order to determine how koalas navigate their environment and partition limited patchy food and nonfood resources. Tree selection among koalas appears to be mediated by folar chemistry, but nonfood tree selection exerts a major impact on home range use due to thermoregulatory constraints. Koalas moved on a daily basis, during both day and night, but daytime resting site was not necessarily in the same location as nighttime feeding site. Koalas had substantial home range overlap in the near absence of resource sharing with less than 1% of trees located in areas of overlap used by multiple koalas. We suggest that koala spatiotemporal distribution and habitat use are probably based upon a community structure of individuals, with a checkerboard model best describing overlap in home range area but not in resource use. Nonfood refugia and social networks should be incorporated into models of animal range and habitat use.     

The Rarest of Darwin's Finches

The Mangrove Finch (Cactospiza heliobates) is unique among Darwin's finches in its specialized habits and naturally fragmented distribution on individual islands. It has been reported to occupy six patches of mangroves separated to varying degrees on two of the Galápagos islands, Isabela and Fernandina. We present the results of a field study in three of the patches. The species appears to have become extinct on Fernandina and to be restricted now to a maximum of four habitat islands on Isabela. In one of them, Playa Tortuga Negra, we estimate the population of breeding adults to be 10–20 pairs. At another, west of Villamil, the population size is probably now small due to the cutting of mangroves. The other two populations, close to each other east of Punta Moreno, probably hold the largest demes. Their status is unknown. We estimate that the genetically effective size of the combined populations is 30–380 individuals. Because the populations are so small and restricted, they are vulnerable to any disturbance of the environment. This has probably been so throughout their history because they inhabit a fragmented habitat in a tectonically active region. New anthropogenic threats this century are an introduced wasp (Polistes versicolor), which may be a predator on a major food item (insect larvae) of the finches, and destruction of the habitat. Protection of the occupied habitat against human exploitation is required for the preservation of the species because no alternative habitat is known.     

Can we measure carrying capacity with foraging behavior?

Carrying capacity is one of the most important, yet least understood and rarely estimated, parameters in population management and modeling. A simple behavioral metric of carrying capacity would advance theory, conservation, and management of biological populations. Such a metric should be possible because behavior is finely attuned to variation in environment including population density. We connect optimal foraging theory with population dynamics and life history to develop a simple model that predicts this sort of adaptive density-dependent change in food consumption. We then confirm the model's unexpected and manifold predictions with field experiments. The theory predicts reproductive thresholds that alter the marginal value of energy as well as the value of time. Both effects cause a pronounced discontinuity in quitting-harvest rate that we revealed with foraging experiments. Red-backed voles maintained across a range of high densities foraged at a lower density-dependent rate than the same animals exposed to low-density treatments. The change in harvest rate is diagnostic of populations that exceed their carrying capacity. Ecologists, conservation biologists, and wildlife managers may thus be able to use simple and efficient foraging experiments to estimate carrying capacity and habitat quality.     

Food regulates reproduction differently in different habitats: experimental evidence in the Goshawk

Food supplementation experiments have been widely used to get detailed insight into how food supply contributes to the reproductive performance of wild animals. Surprisingly, even though food seldom is distributed evenly in space, variation in local habitat quality has usually not been controlled for in food supplementation studies. With results from a two-year feeding experiment involving a habitat-sensitive avian top predator, the Northern Goshawk Accipiter gentilis, we show that treatment effects on goshawk reproductive performance are habitat dependent. Extra food reduced nestling mortality in low-quality territories where prime habitat (forest) is scarce, but not in high-quality territories where prime habitat is abundant. Consequently, brood size did not differ between treatment categories in heavily forested territories, but fledgling numbers differed between unfed and fed goshawk pairs breeding in territories where forest is scarce. However, because extra food was not superabundant, this artificial increase in offspring number induced a dramatic decrease in nestling condition in low-quality territories. Treatment effects were detected even after controlling statistically for other potentially confounding effects (year, territory identity) and strongly covaried with territory-specific abundances of the most important summer prey species. These results highlight the importance of acknowledging the effect that small-scale variation in habitat quality and availability of natural food may have on the results of food supplementation experiments. In order to assess the generality of food supplementation effects, the integration of habitat heterogeneity and variation in food abundance is thus needed, especially among species in which small-scale variation in habitat quality influences demographic patterns.     

Response of a Group of Sichuan Snub-Nosed Monkeys to Commercial Logging in the Qinling Mountains, China

Abstract:  Habitat destruction is one of the greatest threats to primates worldwide. To understand the impact of forest logging on the habitat use of primates in temperate mixed forest, we compared the range, habitat used, population size, and diet of a troop (ERT) of Sichuan snub-nosed monkeys ( Rhinopithecus roxellana ) in the Qinling Mountains before (1989), during (1997), and after (2002–2003) commercial logging. Logging significantly changed the composition of the forest and the food supply for the troop. Some areas were heavily logged and formed patches in the forest that lacked canopy cover. The troop moved 7 km away from their original range when logging took place and returned to their original range after logging stopped, but they avoided heavily logged areas that lacked canopy cover. Their movement indicated some degree of site fidelity in this species. Diet and home range changed after logging, but the population size remained stable, which suggests that this species has some ability to adapt to habitat changes. Our results may reflect a natural flexibility in primates to adapt to the changing food resources in temperate areas with marked seasonal variations in food availability and distribution. This flexibility may have contributed to their higher degree of resilience to habitat alterations caused by human activities compared with tropical forest primates that have a more specialized diet. Our findings provide important baseline information that will help decision makers in their efforts to conserve primates, especially in temperate regions, and to sustainably manage primate habitat.     

Age-specific social dominance affects habitat use by breeding American redstarts (Setophaga ruticilla): a removal experiment

Summary By removing older males from their breeding territories, we tested the hypothesis that age-related dominance behavior influenced the pattern of habitat selection by breeding American redstarts Setophaga ruticilla (Aves: Parulinae). Fifteen older male redstarts removed in five experimental replicates during three breeding seasons were replaced by ten yearling and five older males; no redstart males of either age colonized the control sites during the same time periods, although two yearlings disappeared. Significantly more yearling males (67%, n=9) colonized the vacated areas than were present in the redstart population at large (26.8%, n=209). We reject the alternative hypothesis that yearling male redstarts occupy different habitats from older males because of age-related (innate) habitat preferences. Redstarts that colonized the territories made vacant by our removals (i.e., floaters) were a behaviorally heterogeneous group of animals. The presence of both yearling and older male floaters indicates that suitable habitat is limiting for this species and that intraspecific competitive interactions are important in habitat distribution, and potentially in population regulation.     

Response of Red-Backed Voles to Recent Patch Cutting in Subalpine Forest

We examined the response of southern red-backed voles ( Clethrionomys gapperi ) to patch cutting in a forested landscape in which 23% of the forest cover had been removed by timber harvest. We live trapped voles in and around 18 patchcuts in one watershed of southern Wyoming. Although we found a significant difference in capture rates between patchcut interior and forest habitats in 1 of 2 years, voles did not strongly avoid the interior of patchcuts. This result contrasts with results from most studies of voles in clearcut ecosystems, which report that red-backed voles are generally rare or absent from clearcuts. Capture rates were highest on both sides of the patchcut edge, which also contrasts with studies of voles at the edges of forest remnants. The use of patchcut interior and edge habitats could not be explained as a consequence of juvenile voles dispersing to those habitats or males moving through the habitat in search of mates. We suggest that, despite similar physiognomy in patchcut and clearcut sites, the differences in landscape structure in perforated versus fragmented landscapes lead to very different patterns of vole movement. Understanding the scales at which voles perceive landscapes as coarse- or fine-grained will be key to developing predictive models to aid managers in designing timber sales that maintain high vole populations. Our results emphasize the importance of the spatial pattern and scale of disturbance in determining the response of vertebrates to landscape change and the need for more refined investigations of the consequences of deforestation.     

Movements of vervets (Cercopithecus aethiops) and patas monkeys (Erythrocebus patas) as estimators of food resource size, density, and distribution

The effect of food resources on behavior has been difficult to measure. Here we use animals themselves to describe “effective” food abundance and distribution by comparing, relative to where individuals stopped to eat, movements of (1) adult females living in a small group of vervet monkeys (Cercopithecus aethiops) with those living in a large group and (2) vervets and patas monkeys (Erythrocebus patas). Although females in the large vervet group travelled farther and stopped to eat more often than females in the small vervet group, these differences resulted from foraging in Acacia drepanolobium habitat. In A. xanthophloea habitat, females in the large group travelled less far, travelled shorter distances between foods, and stopped as often as females in the small group. Greater foraging costs of females in larger vervet groups may be offset by access to home ranges of better quality. Compared to patas, vervets travelled shorter distances, moved shorter distances between food sites, stopped less often, and had longer feeding bouts, suggesting that foods of vervets are denser and larger, overall, than foods of patas. When vervets foraged in A. drepanolobium habitat, also the habitat of patas, their foraging behavior became more like that of patas. Vervets travelled farther, stopped more often, and spent less time at food sites in A. drepanolobium habitat than in A. xanthophloea habitat, suggesting that foods are smaller and less usurpable in A. drepanolobium habitat. Distance between foods, a component of food distribution, did not increase, however. The critical variable underlying usurpability of foods may be food site depletion time, a temporal measure. Received: 14 March 1997 / Accepted after revision: 19 October 1997     

Natal dispersal and philopatry in prairie voles (Microtus ochrogaster) in relation to population density,season, and natal social environment

Summary We used intensive livetrapping to examine natal dispersal and philopatry in prairie voles (Microtus ochrogaster). The majority of male (70.0%) and female prairie voles (75.1 %) remained at the natal nest until death. Those males and females that did disperse left home at about the same age (45–55 days) and moved similar distances (28–33 m). Dispersal was more common (1) from small natal groups than from large natal groups, (2) following disappearance of parents, (3) during the breeding period than during the nonbreeding period, and (4) at low population densities than at high densities. Dispersal was not associated with level of competition for mates within natal groups, and dispersers did not differ from nondispersers in body weight. Our data do not support competition for mates or resources as important factors influencing natal dispersal in prairie voles. The absence of sex differences in dispersal tendency or distance, and our fording that more than half of dispersers had become reproductive before leaving the natal nest, lead us to suggest that inbreeding avoidance is not a primary function of dispersal in this species. Dispersal was, however, more common when potential mates within the natal group were relatives than when they were nonrelatives. Although not tested here, if family members avoid mating with one another through patterns of mate choice, then some animals may leave home in search of mates. The precise benefits associated with philopatry in prairie voles remain to be identified. Correspondence to: B. McGuire at her present address     

Frequency-dependent male mate harassment and intra-specific variation in its avoidance by females of the damselfly Ischnura elegans

We focused on male harassment on different female color morphs of the damselfly Ischnura elegans and on variation in morph-specific mating avoidance tactics by females. In I. elegans, one of the female morphs is colored like the conspecific male (andromorphs) while the other morphs are not (gynomorphs). Our first goal was to quantify morph-specific male mating attempts, hence male harassment, in populations with manipulated population parameters (densities, sex ratios, and proportion of andromorphs). Second, we examined the female's perspective by looking for potential differences in morph-specific mating avoidance tactics and success of those tactics in a natural population. Differences in population conditions did influence the number of male mating attempts per morph. The less frequent female morph was always subject to fewer mating attempts, which contradicts earlier hypotheses on mimicry, but supports those that assume that males learn to recognize female morphs. Gynomorphs occupy less open habitat and often fly away when a male approaches, while andromorphs use more open habitat, do not fly large distances and directly face approaching males. Female morphs did not differ in the proportion of successful mating-avoidance attempts. Our results suggest that the maintenance of the color polymorphism is most probably the result of interactive selective forces depending on variation in all population conditions, instead of solely density- or frequency-dependent selection within populations.     

Different escape tactics of two vole species affect the success of the hunting predator, the least weasel

In the ongoing evolutionary arms race between predators and their prey, successful escape from the predator leads to the evolution of improved escape tactics in prey, but also predators become more effective in following and attacking the prey. Antipredatory behavior of prey is considered to be the strongest towards their most dangerous predators. However, prey species can differ both in vulnerability and efficiency of escape to a shared predator. We studied escape reactions of two vole species, the bank vole (Myodes glareolus) and the field vole (Microtus agrestis), under a simulated predation risk of the least weasel (Mustela nivalis nivalis). We conducted a laboratory experiment where a vole was given a possibility to escape from a weasel by fleeing to a horizontal tunnel or climbing the tree. Subsequently to the vole escape decision, we released a weasel to the same tunnel system to test how the weasel succeeded in following the vole. Weasel presence changed the behavior of voles as especially bank voles escaped by climbing. Instead, the majority of field voles fled into the ground-layer tunnel. The different escape tactics of the voles affected the success of the weasel, because climbing voles were less often successfully followed. We suggest that the difference in escape tactics has evolved as an adaptation to different habitats; meadow-exploiting field voles using ground-level escape while bank voles living in three-dimensional forest habitat frequently use arboreal escape tactics. This is likely to lead to different habitat-dependent vulnerabilities to predation in Microtus and Myodes vole species.     

Quantitative recommendations for amphibian terrestrial habitat conservation derived from habitat selection behavior

Conservation scientists have noted that conservation managers rarely use scientific information when making decisions. One of the reasons why managers rarely use scientific information may be that conservation scientists rarely provide their knowledge in a way that can directly be used by conservation practitioners. Here we show how quantitative recommendations for conservation can be derived. Previous research on terrestrial habitat selection behavior of toads (Bufo bufo and Bufo viridis) showed that wood deposits are a key resource in the terrestrial habitat. We used habitat-dependence analysis to estimate the amount of this key resource, wood deposits, that individual toads require. Based on these estimates we then quantify the requirements for wood deposits for a population. Additionally, we quantified the area that a population requires. Although wood deposits vary strongly in size, we found little evidence for size preferences: only one species preferred smallest sizes of wood deposits. We report all the estimates in a way that can be directly used by conservation managers. Habitat-dependence analysis is a simple and useful tool to quantify habitat requirements. Provisioning of wood deposits may improve the quality of terrestrial habitat for amphibians. Thereby, managers may increase the carrying capacity of terrestrial habitats and support elevated population densities.     

Movement ecology: size-specific behavioral response of an invasive snail to food availability

Immigration, emigration, migration, and redistribution describe processes that involve movement of individuals. These movements are an essential part of contemporary ecological models, and understanding how movement is affected by biotic and abiotic factors is important for effectively modeling ecological processes that depend on movement. We asked how phenotypic heterogeneity (body size) and environmental heterogeneity (food resource level) affect the movement behavior of an aquatic snail (Tarebia granifera), and whether including these phenotypic and environmental effects improves advection-diffusion models of movement. We postulated various elaborations of the basic advection diffusion model as a priori working hypotheses. To test our hypotheses we measured individual snail movements in experimental streams at high- and low-food resource treatments. Using these experimental movement data, we examined the dependency of model selection on resource level and body size using Akaike's Information Criterion (AIC). At low resources, large individuals moved faster than small individuals, producing a platykurtic movement distribution; including size dependency in the model improved model performance. In stark contrast, at high resources, individuals moved upstream together as a wave, and body size differences largely disappeared. The model selection exercise indicated that population heterogeneity is best described by the advection component of movement for this species, because the top-ranked model included size dependency in advection, but not diffusion. Also, all probable models included resource dependency. Thus population and environmental heterogeneities both influence individual movement behaviors and the population-level distribution kernels, and their interaction may drive variation in movement behaviors in terms of both advection rates and diffusion rates. A behaviorally informed modeling framework will integrate the sentient response of individuals in terms of movement and enhance our ability to accurately model ecological processes that depend on animal movement.     

Habitat-specific impacts of multiple consumers on plant population dynamics

Multiple consumers often attack seeds, seedlings, and adult plants, but their population-level consequences remain uncertain. We examined how insect and small mammal consumers influenced the demography and abundance of the perennial shrub, bush lupine (Lupinus arboreus). In grassland and dune habitats we established replicate experimental lupine populations in 81-m2 plots that were either protected from, or exposed to, herbivorous voles and granivorous mice (via fencing) and/or root feeding insects (via insecticide treatment). Populations were initiated with transplanted seedlings in 1999 and 2000. We followed the demography of these cohorts, subsequent generations, and the seed bank for 5.5 years. Voles and insects killed many seedlings in dune (1999 only) and grassland (1999 and 2000) habitats. After 2000, insects and voles had minimal effects on seedling or adult survival. Seed predation by granivorous mice, however, greatly depressed seedling recruitment, resulting in lower adult lupine abundance in control plots vs. those protected from rodents. In grasslands, initial effects of voles and insects on seedling survival produced large differences among treatments in adult plant density and the cumulative number of seeds produced throughout the experiment. Differences among grassland populations in seed rain, however, had little influence on the magnitude of seedling recruitment into this habitat. Instead, recruitment out of a preexisting seed bank compensated for the lack of seed production in populations exposed to consumers. Shading by dense adults in plots protected from consumers limited seedling establishment within these populations. Although differences among populations in cumulative seed rain did not influence adult establishment, populations protected from consumers accumulated substantially larger seed banks than controls. These results illustrate how density dependence, habitat-specific seed dynamics, and particular demographic impacts of consumers interact to shape plant population responses to consumers.     

Interference competition between introduced black rats and endemic Galápagos rice rats

Replicated field experiments were used to quantify and to describe the mechanism of competition between the introduced black rat Rattus rattus and the endemic Santiago rice rat Nesoryzomys swarthi on Santiago Island, Galápagos Islands, Ecuador. The removal of R. rattus significantly slowed the rate of seasonal population decline in N. swarthi. This effect was particularly evident for female, relative to male, N. swarthi and appeared to be driven solely by enhanced immigration; no other fitness or space use parameters were affected. The candidate hypotheses to explain the mechanism of competition were exploitation competition, interference by resource defense, and interference by aggressive encounter. To distinguish between hypotheses, we conducted a replicated resource supplementation experiment with patchy food, scattered food, and no food (control) treatments. The opportunistic R. rattus responded to the extra resources with increased adult immigration and juvenile recruitment, resulting in a significant abundance boost of sevenfold on patchy grids and fourfold on scattered grids. Females increased in body mass, and the breeding season was lengthened. In contrast, there was no change in the abundance of N. swarthi and no obvious benefit to reproduction. Instead, the costs of interference apparently outweighed the benefits of extra food: female N. swarthi increased in mass with supplementary food, but female (relative to male) immigration and residency were repressed on all supplemented areas. This response supported the hypothesis of interference by aggressive encounter, and we were able to rule out the alternative hypotheses. Although periodic population crashes of R. rattus on the arid north coast of Santiago may ameliorate its competitive impact, climate change may tip the balance. Control or eradication of R. rattus should improve future survival prospects for N. swarthi, but wildlife managers must be prepared for the potential eruption of the introduced house mouse Mus musculus, because this species experienced a release from interference competition and immigrated to removal areas.     

Effects of anthropogenic developments on common raven nesting biology in the West Mojave Desert.

Subsidized predators may affect prey abundance, distribution, and demography. Common Ravens (Corvus corax) are anthropogenically subsidized throughout their range and, in the Mojave Desert, have increased in number dramatically over the last 3-4 decades. Human-provided food resources are thought to be important drivers of raven population growth, but human developments add other features as well, such as nesting platforms. From 1996 to 2000, we examined the nesting ecology of ravens in the Mojave Desert, relative to anthropogenic developrhent. Ravens nested disproportionately near point sources of food and water subsidies (such as towns, landfills, and ponds) but not near roads (sources of road-killed carrion), even though both sources of subsidy enhanced fledging success. Initiation of breeding activity was more likely when a nest from the previous year was present at the start of a breeding season but was not affected by access to food. The relative effect of environmental modifications on fledging success varied from year to year, but the effect of access to human-provided resources was comparatively consistent, suggesting that humans provide consistently high-quality breeding habitat for ravens. Anthropogenic land cover types in the desert are expected to promote raven population growth and to allow ravens to occupy parts of the desert that otherwise would not support them. Predatory impacts of ravens in the Mojave Desert can therefore be considered indirect effects of anthropogenic development.     

Reducing per capita food supply alters urchin condition and habitat

When food supply declines or population density increases, the per capita food availability is reduced causing a decline in condition of consumers. Many consumers alter their feeding behaviour and ultimately the surrounding community (e.g. overgrazing and formation of urchin barrens). This study tested the hypothesis that sea urchin populations are of greater density and poorer condition in barrens (little food) than forest habitat (lots of food). We then tested the hypothesis that a decrease in per capita food supply to urchins has a negative effect not only on their condition but also on their surrounding habitat. We experimentally assessed the effect of limited food supply and increased density of a subtidal Australian sea urchin (Heliocidaris erythrogramma) on their condition (i.e. gonad index) and surrounding benthic habitat (i.e. turf-forming algae). Our results show that a reduction in food supply led to poorer consumer condition and greater herbivory on surrounding local habitat. We provide evidence that per capita food reduction is one of the necessary conditions for the over-consumption by urchins (i.e. urchin barrens), a proposed but previously untested mechanism.     

Bark salicylates and condensed tannins reduce vole browsing amongst cultivated dark-leaved willows (<Emphasis Type="Italic">Salix myrsinifolia</Emphasis>)

Summary. Vole feeding amongst herbal willows that have a high concentration of salicylates in their bark and leaves, and may therefore be cultivated for use as raw material for herbal medicine was tested in the field and in laboratory conditions. Eight clones of dark-leaved willow (Salix myrsinifolia Salisb.) were cultivated for two years with six different methods combining three fertilisation levels (none, low and high), black plastic mulch applied for suppressing weed competition and unmulched control. Samples for the laboratory feeding trial were taken from the unfertilised plants during willow winter dormancy and twigs were fed to 16 voles as a multi-choice experiment. The bark area removed was calculated from image analysis of the material left by the voles. The diameter and the bark thickness of the twigs were measured. Concentrations of salicin, salicortin, HCH-salicortin, picein, triandrin, triandrin derivative, gallocatechin, (+)-catechin, luteolin-7-glucoside, hyperin, total condensed tannins and total nitrogen were measured from the twigs fed to voles in the laboratory. Browsing by a natural population of voles amongst winter-dormant willows was measured in the field. In the laboratory, voles browsed on 80% of the twigs and in the field voles browsed on 33% of the twigs. Vole feeding followed similar patterns in the field and in the laboratory experiment; feeding was clearly higher amongst the plants grown in unmulched control compared to those in plastic mulch. The same clones, 1, 2 and 6 were preferred in both experiments. Voles preferred thin twigs to thick ones. Feeding correlated negatively with concentrations of salicylates and tannins. As vole feeding seems to be highly affected by willow cultivation method and plant genotype, careful selection of cultivated clones and cultivation methods can enhance the reliability of herbal willow cultivation.