Evidence for a survival-driven traveling wave in a keystone boreal predator population.

Cyclical population dynamics are a common phenomenon in populations worldwide, yet the spatial organization of these cycles remains poorly understood. In this study, we investigated the spatial form and timing of a population collapse from 2018 to 2022 in Canada lynx (Lynx canadensis) across the northwest boreal forest. We analyzed survival, reproduction, and dispersal data from 143 individual global positioning system (GPS) collared lynx from populations across five study sites spanning interior Alaska to determine whether lynx displayed characteristics of a population wave following a concurrent wave in snowshoe hare (Lepus americanus) abundance. Reproductive rates declined across the study sites; however, site-level reproduction declined first in our easternmost study sites, supporting the idea of a population wave. Despite a clear increase in percent of dispersing lynx, there was no evidence of directional bias in dispersal following a hare population wave. Analysis did show increasingly poor survival for lynx dispersing to the east compared to combined resident and westward dispersal. This pattern is consistent with a survival-mediated population wave in lynx as the driver of the theorized population wave. The combination of these factors supports the idea of a hierarchical response to snowshoe hare population declines with a drop in lynx reproduction followed by increased dispersal, and finally reduced survival. All of this evidence is consistent with the expected characteristics of a population undergoing a traveling wave and supports the hypothesis that lynx presence may facilitate and mirror the underlying wave patterns in snowshoe hare.

The evolution of white-tailed jackrabbit camouflage in response to past and future seasonal climates.

The genetic basis of adaptive traits has rarely been used to predict future vulnerability of populations to climate change. We show that light versus dark seasonal pelage in white-tailed jackrabbits (Lepus townsendii) tracks snow cover and is primarily determined by genetic variation at endothelin receptor type B (EDNRB), corin serine peptidase (CORIN), and agouti signaling protein (ASIP). Winter color variation was associated with deeply divergent alleles at these genes, reflecting selection on both ancestral and introgressed variation. Forecasted reductions in snow cover are likely to induce widespread camouflage mismatch. However, simulated populations with variation for darker winter pelage are predicted to adapt rapidly, providing a trait-based genetic framework to facilitate evolutionary rescue. These discoveries demonstrate how the genetic basis of climate change adaptation can inform conservation.

Behavioural adjustments of predators and prey to wind speed in the boreal forest.

Wind speed can have multifaceted effects on organisms including altering thermoregulation, locomotion, and sensory reception. While forest cover can substantially reduce wind speed at ground level, it is not known if animals living in forests show any behavioural responses to changes in wind speed. Here, we explored how three boreal forest mammals, a predator and two prey, altered their behaviour in response to average daily wind speeds during winter. We collected accelerometer data to determine wind speed effects on activity patterns and kill rates of free-ranging red squirrels (n = 144), snowshoe hares (n = 101), and Canada lynx (n = 27) in Kluane, Yukon from 2015 to 2018. All 3 species responded to increasing wind speeds by changing the time they were active, but effects were strongest in hares, which reduced daily activity by 25%, and lynx, which increased daily activity by 25%. Lynx also increased the number of feeding events by 40% on windy days. These results highlight that wind speed is an important abiotic variable that can affect behaviour, even in forested environments.

Integrating plant stoichiometry and feeding experiments: state-dependent forage choice and its implications on body mass.

Intraspecific feeding choices comprise a large portion of herbivore foraging decisions. Plant resource quality is heterogeneously distributed, affected by nutrient availability and growing conditions. Herbivores navigate landscapes, foraging not only according to food qualities, but also energetic and nutritional demands. We test three non-exclusive foraging hypotheses using the snowshoe hare (Lepus americanus): (1) herbivore feeding choices and body conditions respond to intraspecific plant quality variation; (2) high energetic demands mitigate feeding responses; and (3) feeding responses are inflated when nutritional demands are high. We measured black spruce (Picea mariana) nitrogen, phosphorus and terpene compositions, as indicators of quality, within a snowshoe hare trapping grid and found plant growing conditions to explain spruce quality variation (R2 < 0.36). We then offered two qualities of spruce (H1) from the trapping grid to hares in cafeteria-style experiments and measured their feeding and body condition responses (n = 75). We proxied energetic demands (H2) with ambient temperature and coat insulation (% white coat) and nutritional demands (H3) with the spruce quality (nitrogen and phosphorus content) in home ranges. Hares with the strongest preference for high-quality spruce lost on average 2.2% less weight than hares who ate the least high-quality spruce relative to low-quality spruce. The results supported our energetic predictions as follows: hares in colder temperatures and with less-insulative coats (lower % white) consumed more spruce and were less selective towards high-quality spruce. Collectively, we found variation in plant growing conditions within herbivore home ranges substantial enough to affect herbivore body conditions, but energetic stats mediate plant-herbivore interactions.

A Mathematical Model Supporting a Hyperpredation Effect in the Apparent Competition Between Invasive Eastern Cottontail and Native European Hare.

In this work a mathematical model is built in order to validate on theoretical grounds field study results on a three-species system made of two prey, of which one is native and another one invasive, together with a native predator. Specifically, our results mathematically describe the negative effect on the native European hare after the introduction of the invasive Eastern cottontail, mediated by an increased predation rate by foxes. Two nonexclusive assumptions can be made: an increase in cottontail abundance would lead to a larger fox population, magnifying their predatory impact ("hyperpredation") on hares; alternatively, cottontails attract foxes in patches where they live, which are also important resting sites for hares and consequently the increased presence of foxes results in a higher predation rates on hares. The model results support hyperpredation of increasing fox populations on native hares.

Sex and Age-Specific Hematology and Biochemistry Reference Intervals of Live Iberian Hares (Lepus granatensis) and Comparison with Postmortem Sampling.

Hematology and serum biochemistry reference intervals were estimated for the Iberian hare (Lepus granatensis). Most parameters differed significantly between hunted and livetrapped Iberian hares. Significant differences were found for sex (red blood cell count, mean corpuscular volume, total protein, albumin, uric acid, triglycerides, cholesterol, chloride) and age classes (red blood cell count, hematocrit, mean corpuscular hemoglobin, glucose, calcium, and sodium). Sex- and age-specific reference intervals were estimated for these parameters. Red blood cell count, hemoglobin concentration, mean corpuscular hemoglobin, urea, and potassium show seasonal variations, with the lowest values in summer and the highest in winter. Creatinine, calcium, sodium, and phosphorus achieve their highest values in summer and stable baseline values throughout the rest of the year. These reference intervals can be used as baseline to monitor health, physiology, ecology, and nutrition of Iberian hare populations.

Convergent evolution of seasonal camouflage in response to reduced snow cover across the snowshoe hare range.

Determining how different populations adapt to similar environments is fundamental to understanding the limits of adaptation under changing environments. Snowshoe hares (Lepus americanus) typically molt into white winter coats to remain camouflaged against snow. In some warmer climates, hares have evolved brown winter camouflage-an adaptation that may spread in response to climate change. We used extensive range-wide genomic data to (1) resolve broad patterns of population structure and gene flow and (2) investigate the factors shaping the origins and distribution of winter-brown camouflage variation. In coastal Pacific Northwest (PNW) populations, winter-brown camouflage is known to be determined by a recessive haplotype at the Agouti pigmentation gene. Our phylogeographic analyses revealed deep structure and limited gene flow between PNW and more northern Boreal populations, where winter-brown camouflage is rare along the range edge. Genome sequencing of a winter-brown snowshoe hare from Alaska shows that it lacks the winter-brown PNW haplotype, reflecting a history of convergent phenotypic evolution. However, the PNW haplotype does occur at low frequency in a winter-white population from Montana, consistent with the spread of a locally deleterious recessive variant that is masked from selection when rare. Simulations of this population further show that this masking effect would greatly slow the selective increase of the winter-brown Agouti allele should it suddenly become beneficial (e.g., owing to dramatic declines in snow cover). Our findings underscore how allelic dominance can shape the geographic extent and rate of convergent adaptation in response to rapidly changing environments.

Noninvasive measures of physiological stress are confounded by exposure.

Glucocorticoids and glucocorticoid metabolites are increasingly used to index physiological stress in wildlife. Although feces is often abundant and can be collected noninvasively, exposure to biotic and abiotic elements may influence fecal glucocorticoid metabolite (FGM) concentrations, leading to inaccurate conclusions regarding wildlife physiological stress. Using captive snowshoe hares (Lepus americanus) and simulated environmental conditions, we evaluated how different realistic field conditions and temporal sampling constraints might influence FGM concentrations using an 11-oxoetiocholanolone-enzyme immunoassay. We quantified how fecal pellet age (i.e., 0-6 days), variable summer temperatures, and precipitation affected FGM concentrations. Fecal pellet age had a strong effect on FGM concentrations (ßAge = 0.395, s.d. = 0.085; ß2Age = -0.061, s.d. = 0.012), which were lowest at the beginning and end of our exposure period (e.g., meanday6 = 37.7 ng/mg) and typically highest in the middle (meanday3 = 51.8 ng/mg). The effect of fecal pellet age on FGM concentrations varied across treatments with warm-dry and cool-wet conditions resulting in more variable FGM concentrations relative to control samples. Given the confounding effects of exposure and environmental conditions, if fresh fecal pellet collection is not an option, we encourage researchers to develop a temporally consistent sampling protocol to ensure all samples are exposed to similar environmental conditions.

Introgression drives repeated evolution of winter coat color polymorphism in hares.

Changing from summer-brown to winter-white pelage or plumage is a crucial adaptation to seasonal snow in more than 20 mammal and bird species. Many of these species maintain nonwhite winter morphs, locally adapted to less snowy conditions, which may have evolved independently. Mountain hares (Lepus timidus) from Fennoscandia were introduced into the Faroe Islands in 1855. While they were initially winter-white, within ∼65 y all Faroese hares became winter-gray, a morph that occurs in the source population at low frequency. The documented population history makes this a valuable model for understanding the genetic basis and evolution of the seasonal trait polymorphism. Through whole-genome scans of differentiation and single-nucleotide polymorphism (SNP) genotyping, we associated winter coat color polymorphism to the genomic region of the pigmentation gene Agouti, previously linked to introgression-driven winter coat color variation in the snowshoe hare (Lepus americanus). Lower Agouti expression in the skin of winter-gray individuals during the autumn molt suggests that regulatory changes may underlie the color polymorphism. Variation in the associated genomic region shows signatures of a selective sweep in the Faroese population, suggesting that positive selection drove the fixation of the variant after the introduction. Whole-genome analyses of several hare species revealed that the winter-gray variant originated through introgression from a noncolor changing species, in keeping with the history of ancient hybridization between the species. Our findings show the recurrent role of introgression in generating winter coat color variation by repeatedly recruiting the regulatory region of Agouti to modulate seasonal coat color change.

Living on the edge - circadian habitat usage in pre-weaning European hares (Lepus europaeus) in an intensively used agricultural area.

Over the last decades, the European hare (Lepus europaeus) has become the subject of many interdisciplinary studies due to the sharp Europe-wide population decline. In European hares, the first stage of life until weaning and the subsequent dispersal have been sparsely studied, in particular, habitat selection, movements and survival rate, as juveniles´ precocial lifestyle is dominated by concealment, motionlessness and inconspicuousness. In this study, free-living juvenile European hares (leverets) were detected systematically by thermography (n = 394), radio-tagged or marked (n = 122) from birth until the fifth week of life to research their habitat usage and pre-dispersal movements. The day-resting places and night locations, as well as the distance moved by leverets with aging, were evaluated by generalized linear mixed effect models. In addition, the habitat preference was assessed by a conservative use-availability analysis. Up to the fifth week of life, 30.5% of all leverets used cultivated areas in the daytime. In contrast, the remaining 69.4% animals inhabitated linear or small planar structures in the daytime, with the edges of field tracks, hedges and some ruderal structures clearly being preferred. At nighttime, 93% of all juveniles, which occupied linear structures in the daytime, used the adjoining fields up to 20 m away from the next linear structure. Nocturnal distances of more than 60 m to the next edge rarely occurred before the end of the pre-weaning phase. The time of day and age have a significant influence on the distance moved by juvenile hares. With increasing age, leverets moved less during the day and roamed further at night. The results are largely consistent with the behavioral patterns found in the few previous studies on pre-weaning European hares and show the importance of hiding places for leverets in early life stages. This study should contribute to a better understanding of behavior in juvenile life-history stages of European hares that may help to identify vulnerable phases in their lifecycle. In addition, the findings can refine existing population models and improve conservation efforts.

The Cooccurrence of Hypolagus and Lepus (Leporidae, Lagomorpha) in the Early Pleistocene of Crimea.

New fossil leporid materials from the Lower Pleistocene deposits of the Taurida Cave in the central Crimea (Zuya village, Belogorsk district) are referred to Hypolagus brachygnathus (Kormos, 1930) and Lepus sp. The cooccurrence of Hypolagus and Lepus in the Early Pleistocene of Crimea is significant in terms of the question of the first appearance of Lepus in Europe, as well as the hypothesis of the competition with hares as one of the possible causes of extinction of Hypolagus.

The importance of wildlife in the ecology and epidemiology of the TBE virus in Sweden: incidence of human TBE correlates with abundance of deer and hares.

BACKGROUND: Tick-borne encephalitis (TBE) is one tick-transmitted disease where the human incidence has increased in some European regions during the last two decades. We aim to find the most important factors causing the increasing incidence of human TBE in Sweden. Based on a review of published data we presume that certain temperature-related variables and the population densities of transmission hosts, i.e. small mammals, and of primary tick maintenance hosts, i.e. cervids and lagomorphs, of the TBE virus vector Ixodes ricinus, are among the potentially most important factors affecting the TBE incidence. Therefore, we compare hunting data of the major tick maintenance hosts and two of their important predators, and four climatic variables with the annual numbers of human cases of neuroinvasive TBE. Data for six Swedish regions where human TBE incidence is high or has recently increased are examined by a time-series analysis. Results from the six regions are combined using a meta-analytical method. RESULTS: With a one-year time lag, the roe deer (Capreolus capreolus), red deer (Cervus elaphus), mountain hare (Lepus timidus) and European hare (Lepus europaeus) showed positive covariance; the Eurasian elk (moose, Alces alces) and fallow deer (Dama dama) negative covariance; whereas the wild boar (Sus scrofa), lynx (Lynx lynx), red fox (Vulpes vulpes) and the four climate parameters showed no significant covariance with TBE incidence. All game species combined showed positive covariance. CONCLUSIONS: The epidemiology of TBE varies with time and geography and depends on numerous factors, i.a. climate, virus genotypes, and densities of vectors, tick maintenance hosts and transmission hosts. This study suggests that the increased availability of deer to I. ricinus over large areas of potential tick habitats in southern Sweden increased the density and range of I. ricinus and created new TBEV foci, which resulted in increased incidence of human TBE. New foci may be established by TBE virus-infected birds, or by birds or migrating mammals infested with TBEV-infected ticks. Generally, persistence of TBE virus foci appears to require presence of transmission-competent small mammals, especially mice (Apodemus spp.) or bank voles (Myodes glareolus).

Adaptive introgression underlies polymorphic seasonal camouflage in snowshoe hares.

Snowshoe hares (Lepus americanus) maintain seasonal camouflage by molting to a white winter coat, but some hares remain brown during the winter in regions with low snow cover. We show that cis-regulatory variation controlling seasonal expression of the Agouti gene underlies this adaptive winter camouflage polymorphism. Genetic variation at Agouti clustered by winter coat color across multiple hare and jackrabbit species, revealing a history of recurrent interspecific gene flow. Brown winter coats in snowshoe hares likely originated from an introgressed black-tailed jackrabbit allele that has swept to high frequency in mild winter environments. These discoveries show that introgression of genetic variants that underlie key ecological traits can seed past and ongoing adaptation to rapidly changing environments.

Brown Hare's (Lepus europaeus) Histone H1 Variant H1.2 as an Indicator of Anthropogenic Stress.

From the liver tissues of brown hare individuals that lived in two various habitats, i.e., the agricultural region with the predominant farms and the industrial area near a metallurgical plant, histones H1 were analyzed to compare their within and between population variability. Furthermore, because agricultural production emits mainly organic pollutants and metallurgical industry is a primarily source of inorganic contaminations, we wanted to check how the brown hare individuals are sensitive for both agents. Among brown hare H1 histones, the histone H1.2 was determined as heterogeneous due to its varied mobility in two-dimensional SDS-polyacrylamide gel. The obtained electrophoretic patterns contained differently moving single spots of histone H1.2 and also its double spots have a similar rate of electrophoretic mobility. Based on this, two homozygous phenotypes (slowly migrating 2a and faster moving 2b) and a heterozygous phenotype (2a2b) was distinguished. The relatively low variable (CV < 0.25) and comparably abundant (p > 0.05) histone H1.2 homozygous phenotypes form a heterozygous phenotype in a similar proportion, at a ratio approximating 0.5. Although the brown hare population originating from agricultural area displayed a slight excess of heterozygous individuals 2a2b (F = - 0.04), it was conformed to the Hardy-Weinberg assumption (χ2 = 0.035, p = 0.853). Compared with this population, a sevenfold reduced frequency of the phenotype 2b and above tenfold increase of a heterozygosity (F = - 0.53) was observed in the brown hare population inhabiting the vicinity of metallurgical plant. Therefore, this population did not fit to the Hardy-Weinberg law (χ2 = 5.65, p = 0.017). Despite the negligible genetic differentiation (FST = 0.026) between brown hare populations inhabiting areas with different anthropogenic pressure, a statistically significant difference in the distribution of their phenotypes (χ2 = 6.01, p = 0.049) and alleles (χ2 = 6.50, p = 0.013) was noted. The collected data confirm that the brown hare species is sensitive for environmental quality and may serve as a good indicator of habitat conditions related to both organic pollution emitted by agricultural activities (PIC = 0.48) and inorganic contamination originating from metallurgical processes (PIC = 0.49). These difference in the environmental quality might be assessed by estimation of genetic variability among the brown hare populations, based on the phenotypes distribution of histone H1 variant H1.2, the protein that was not so far employed as a molecular marker of anthropogenic stress.

Functional responses of white spruce to snowshoe hare herbivory at the treeline.

Herbivores can modify the rate of shrub and treeline advance. Both direct and indirect effects of herbivory may simultaneously interact to affect the growth rates of plants at this ecotone. We investigated the effect of snowshoe hare herbivory on the height of white spruce at two treeline locations in Alaska, USA. White spruce is expanding its distribution both upwards in elevation and northward in latitude because of climate warming, and snowshoe hares are already present in areas likely to be colonized by spruce. We hypothesized that herbivory would result in browsed individuals having reduced height, suggesting herbivory is a direct, negative effect on spruce treeline advance. We found an interactive effect between browsing history and spruce age. When young (under 30 years old), individuals that were browsed tended to be taller than unbrowsed individuals. However, older seedlings (over 30 years old) that had been browsed were shorter than unbrowsed individuals of the same age. Hares suppress faster growing individuals that are initially taller by preferentially browsing them as they emerge above the winter snowpack. This reduced height, in combination with increased mortality associated with browsing, is predicted to slow the advance of both latitudinal and altitudinal treeline expansions and alter the structure of treeline forests.

An Inverse Problem: Trappers Drove Hares to Eat Lynx.

The Canadian lynx and snowshoe hare pelt data by the Hudson Bay Company did not fit the classical predator-prey theory. Rather than following the peak density of the hare, that of the lynx leads it, creating the hares-eat-lynx (HEL) paradox. Although trappers were suspected to play a role, no mathematical model has ever demonstrated the HEL effect. Here we show that the long-held assumption that the pelt number is a proxy of the wild populations is false and that when the data are modeled by the harvest rates by the trappers, the problem is finally resolved: both the HEL paradox and the classical theory are unified in our mechanistic hare-lynx-competitor-trapper (HLCT) model where competitor stands for all predators of the hares other than the lynx. The result is obtained by systematically fitting the data to various models using Newton's inverse problem method. Main findings of this study include: the prey-eats-predator paradox in kills by an intraguild top-predator can occur if the top-predator prefers the predator to the prey; the benchmark HLCT model is more sensitive to all lynx-trapper interactions than to the respective hare-trapper interactions; the Hudson Bay Company's hare pelt number maybe under-reported; and, the most intriguing of all, the trappers did not interfere in each other's trapping activities.

Alpine glacial relict species losing out to climate change: The case of the fragmented mountain hare population (Lepus timidus) in the Alps.

Alpine and Arctic species are considered to be particularly vulnerable to climate change, which is expected to cause habitat loss, fragmentation and-ultimately-extinction of cold-adapted species. However, the impact of climate change on glacial relict populations is not well understood, and specific recommendations for adaptive conservation management are lacking. We focused on the mountain hare (Lepus timidus) as a model species and modelled species distribution in combination with patch and landscape-based connectivity metrics. They were derived from graph-theory models to quantify changes in species distribution and to estimate the current and future importance of habitat patches for overall population connectivity. Models were calibrated based on 1,046 locations of species presence distributed across three biogeographic regions in the Swiss Alps and extrapolated according to two IPCC scenarios of climate change (RCP 4.5 & 8.5), each represented by three downscaled global climate models. The models predicted an average habitat loss of 35% (22%-55%) by 2100, mainly due to an increase in temperature during the reproductive season. An increase in habitat fragmentation was reflected in a 43% decrease in patch size, a 17% increase in the number of habitat patches and a 34% increase in inter-patch distance. However, the predicted changes in habitat availability and connectivity varied considerably between biogeographic regions: Whereas the greatest habitat losses with an increase in inter-patch distance were predicted at the southern and northern edges of the species' Alpine distribution, the greatest increase in patch number and decrease in patch size is expected in the central Swiss Alps. Finally, both the number of isolated habitat patches and the number of patches crucial for maintaining the habitat network increased under the different variants of climate change. Focusing conservation action on the central Swiss Alps may help mitigate the predicted effects of climate change on population connectivity.

A time-lagged effect of conspecific density on habitat selection by snowshoe hare.

Ideal free distribution theory predicts that increased conspecific density redistributes individuals to low-density, suboptimal habitat. However, possible lags in response to population density remain poorly documented. Snowshoe hares (Lepus americanus) may exhibit density-dependent habitat selection due to its marked variation in population density. Based on 11 years (2004-2014) of snow tracking in Quebec (Canada), we investigated snowshoe hares' short-term and delayed habitat selection responses to population density. We predicted that at high densities, hare distribution expands into low-density habitat, thus weakening the association between hares and high-density habitat. We surveyed hare tracks along 95 km of transects on average each year and georeferenced 14,240 tracks. We used Generalized Estimating Equations for track count per 100 m transect segment as a function of the proportion of different forest age classes (0-20 y, 20-40 y and 40-80 y) within 50 m of the segments. We used model coefficients for each age class as a measure of habitat preference, and modeled those coefficients as a function of a population density index in current and previous winters. Coefficients for 20- to 40-y-old forests were positive each year, indicating that this habitat was preferred. The association between track counts and 20- to 40-y-old forest significantly declined with density during the previous winter, suggesting that hare spread from preferred forest with a lagged response to density. To our knowledge, no previous empirical studies have documented a lagged habitat selection response to population density. Time lags offer possible explanation for documented deviations from ideal free distribution models.

Using experimentation to understand the 10-year snowshoe hare cycle in the boreal forest of North America.

Population cycles have long fascinated ecologists from the time of Charles Elton in the 1920s. The discovery of large population fluctuations in undisturbed ecosystems challenged the idea that pristine nature was in a state of balance. The 10-year cycle of snowshoe hares (Lepus americanus Erxleben) across the boreal forests of Canada and Alaska is a classic cycle, recognized by fur traders for more than 300 years. Since the 1930s, ecologists have investigated the mechanisms that might cause these cycles. Proposed causal mechanisms have varied from sunspots to food supplies, parasites, diseases, predation and social behaviour. Both the birth rate and the death rate change dramatically over the cycle. Social behaviour was eliminated as a possible cause because snowshoe hares are not territorial and do not commit infanticide. Since the 1960s, large-scale manipulative experiments have been used to discover the major limiting factors. Food supply and predation quickly became recognized as potential key factors causing the cycle. Experiments adding food and restricting predator access to field populations have been decisive in pinpointing predation as the key mechanism causing these fluctuations. The immediate cause of death of most snowshoe hares is predation by a variety of predators, including the Canada lynx (Lynx canadensis Kerr). The collapse in the reproductive rate is not due to food shortage as was originally thought, but is a result of chronic stress from predator chases. Five major issues remain unresolved. First, what is the nature of the predator-induced memory that results in the prolonged low phase of the cycle? Second, why do hare cycles form a travelling wave, starting in the centre of the boreal forest in Saskatchewan and travelling across western Canada and Alaska? Third, why does the amplitude of the cycle vary greatly from one cycle to the next in the same area? Fourth, do the same mechanisms of population limitation apply to snowshoe hares in eastern North American or in similar ecosystems across Siberia? Finally, what effect will climatic warming have on all the above issues? The answers to these questions remain for future generations of biologists to determine.

The effects of acaricide treatment of sheep on red grouse Lagopus lagopus scotica tick burdens and productivity in a multi-host system.

Ixodes ricinus (Ixodida: Ixodidae) ticks are of economic and pathogenic importance across Europe. Within the uplands of the U.K., management to reduce ticks is undertaken to benefit red grouse Lagopus lagopus scotica (Galliformes: Phasianidae). Management strategies focus on the acaricide treatment of domestic sheep Ovis aries (Artiodactyla: Bovidae), but the effectiveness of this is less certain in the presence of wild hosts, particularly red deer Cervus elaphus (Artiodactyla: Cervidae) and mountain hare Lepus timidus (Lagomorpha: Leporidae). This study examines the effects of sheep management on grouse tick burdens and productivity using sites with a range of wild host densities. Sites at which applications of acaricide were more frequent had lower tick burdens; this relationship was similar on sites with a range of deer densities. However, no direct link was detected between acaricide treatment interval and grouse productivity. Sites with higher deer densities had higher grouse tick burdens and lower productivity [mean ± standard error (SE) young : adult ratio: 1.2 ± 0.2] compared with sites with lower deer densities (mean ± SE young : adult ratio: 1.8 ± 0.1). Sites with higher grouse brood sizes and higher proportions of hens with broods were also those with higher mountain hare abundance indices. This study highlights the importance of the frequent treatment of sheep with acaricide to reduce tick burdens on grouse, even in the presence of wild hosts.