Spreading speed of chronic wasting disease across deer groups with overlapping home ranges.

Chronic wasting disease (CWD) is a fatal disease of cervid species that continues to spread across North America and now in Europe. It poses a threat to cervid populations and the local ecological and economic communities that depend on them. Although empirical studies have shown that host home range overlap and male dispersal are important in the spread of disease, there are few mechanistic models explicitly considering those factors. We built a spatio-temporal, differential equation model for CWD spreading with restricted movement of hosts within home ranges. The model incorporates both direct and environmental transmission within and between groups as well as male dispersal. We compared the relative influence of host density, sex ratio, home range size, and male dispersal distance on the spreading speed using sensitivity analysis. We also assessed the effect of landscape heterogeneity, quantified as edge density, on the spreading speed of CWD because it jointly alters the host density and home range size. Our model binds the theoretical study of CWD spreading speed together with empirical studies on deer home ranges and sets a base for models in 2D space to evaluate management and control strategies.

Behavioural flexibility in migratory behaviour in a long-lived large herbivore.

Migratory animals are predicted to enhance lifetime fitness by obtaining higher quality forage and/or reducing predation risk compared to non-migratory conspecifics. Despite evidence for behavioural flexibility in other taxa, previous research on large mammals has often assumed that migratory behaviour is a fixed behavioural trait. Migratory behaviour may be plastic for many species, although few studies have tested for individual-level flexibility using long-term monitoring of marked individuals, especially in large mammals such as ungulates. We tested variability in individual migratory behaviour using a 10-year telemetry data set of 223 adult female elk (Cervus elaphus) in the partially migratory Ya Ha Tinda population in Alberta, Canada. We used net squared displacement (NSD) to classify migratory strategy for each individual elk-year. Individuals switched between migrant and resident strategies at a mean rate of 15% per year, and migrants were more likely to switch than residents. We then tested how extrinsic (climate, elk/wolf abundance) and intrinsic (age) factors affected the probability of migrating, and, secondly, the decision to switch between migratory strategies. Over 630 individual elk-years, the probability of an individual elk migrating increased following a severe winter, in years of higher wolf abundance, and with increasing age. At an individual elk level, we observed 148 switching events of 430 possible transitions in elk monitored at least 2 years. We found switching was density-dependent, where migrants switched to a resident strategy at low elk abundance, but residents switched more to a migrant strategy at high elk abundance. Precipitation during the previous summer had a weak carryover effect, with migrants switching slightly more following wetter summers, whereas residents showed the opposite pattern. Older migrant elk rarely switched, whereas resident elk switched more frequently to migrate at older ages. Our results show migratory behaviour in ungulates is an individually variable trait that can respond to intrinsic, environmental and density-dependent forces. Different strategies had opposing responses to density-dependent and intrinsic drivers, providing a stabilizing mechanism for the maintenance of partial migration and demographic fitness in this population.

REVIEW: Can habitat selection predict abundance?

Habitats have substantial influence on the distribution and abundance of animals. Animals' selective movement yields their habitat use. Animals generally are more abundant in habitats that are selected most strongly. Models of habitat selection can be used to distribute animals on the landscape or their distribution can be modelled based on data of habitat use, occupancy, intensity of use or counts of animals. When the population is at carrying capacity or in an ideal-free distribution, habitat selection and related metrics of habitat use can be used to estimate abundance. If the population is not at equilibrium, models have the flexibility to incorporate density into models of habitat selection; but abundance might be influenced by factors influencing fitness that are not directly related to habitat thereby compromising the use of habitat-based models for predicting population size. Scale and domain of the sampling frame, both in time and space, are crucial considerations limiting application of these models. Ultimately, identifying reliable models for predicting abundance from habitat data requires an understanding of the mechanisms underlying population regulation and limitation.

How many routes lead to migration? Comparison of methods to assess and characterize migratory movements.

Decreasing rate of migration in several species as a consequence of climate change and anthropic pressure, together with increasing evidence of space-use strategies intermediate between residency and complete migration, are very strong motivations to evaluate migration occurrence and features in animal populations. The main goal of this paper was to perform a relative comparison between methods for identifying and characterizing migration at the individual and population level on the basis of animal location data. We classified 104 yearly individual trajectories from five populations of three deer species as migratory or non-migratory, by means of three methods: seasonal home range overlap, spatio-temporal separation of seasonal clusters and the Net Squared Displacement (NSD) method. For migratory cases, we also measured timing and distance of migration and residence time on the summer range. Finally, we compared the classification in migration cases across methods and populations. All methods consistently identified migration at the population level, that is, they coherently distinguished between complete or almost complete migratory populations and partially migratory populations. However, in the latter case, methods coherently classified only about 50% of the single cases, that is they classified differently at the individual-animal level. We therefore infer that the comparison of methods may help point to 'less-stereotyped' cases in the residency-to-migration continuum. For cases consistently classified by all methods, no significant differences were found in migration distance, or residence time on summer ranges. Timing of migration estimated by NSD was earlier than by the other two methods, both for spring and autumn migrations. We suggest three steps to identify improper inferences from migration data and to enhance understanding of intermediate space-use strategies. We recommend (i) classifying migration behaviours using more than one method, (ii) performing sensitivity analysis on method parameters to identify the extent of the differences and (iii) investigating inconsistently classified cases as these may often be ecologically interesting (i.e. less-stereotyped migratory behaviours).

Selection, use, choice and occupancy: clarifying concepts in resource selection studies.

1. During the last decade, there has been a proliferation of statistical methods for studying resource selection by animals. While statistical techniques are advancing at a fast pace, there is confusion in the conceptual understanding of the meaning of various quantities that these statistical techniques provide. 2. Terms such as selection, choice, use, occupancy and preference often are employed as if they are synonymous. Many practitioners are unclear about the distinctions between different concepts such as 'probability of selection,' 'probability of use,' 'choice probabilities' and 'probability of occupancy'. 3. Similarly, practitioners are not always clear about the differences between and relevance of 'relative probability of selection' vs. 'probability of selection' to effective management. 4. Practitioners also are unaware that they are using only a single statistical model for modelling resource selection, namely the exponential probability of selection, when other models might be more appropriate. Currently, such multimodel inference is lacking in the resource selection literature. 5. In this paper, we attempt to clarify the concepts and terminology used in animal resource studies by illustrating the relationships among these various concepts and providing their statistical underpinnings.

Wildlife disease elimination and density dependence.

Disease control by managers is a crucial response to emerging wildlife epidemics, yet the means of control may be limited by the method of disease transmission. In particular, it is widely held that population reduction, while effective for controlling diseases that are subject to density-dependent (DD) transmission, is ineffective for controlling diseases that are subject to frequency-dependent (FD) transmission. We investigate control for horizontally transmitted diseases with FD transmission where the control is via culling or harvest that is non-selective with respect to infection and the population can compensate through DD recruitment or survival. Using a mathematical model, we show that culling or harvesting can eradicate the disease, even when transmission dynamics are FD. Eradication can be achieved under FD transmission when DD birth or recruitment induces compensatory growth of new, healthy individuals, which has the net effect of reducing disease prevalence by dilution. We also show that if harvest is used simultaneously with vaccination, and there is high enough transmission coefficient, application of both controls may be less efficient than vaccination alone. We illustrate the effects of these control approaches on disease prevalence for chronic wasting disease in deer where the disease is transmitted directly among deer and through the environment.

Large herbivores in a partially migratory population search for the ideal free home.

Migration is a tactic used across taxa to access resources in temporally heterogenous landscapes. Populations that migrate can attain higher abundances because such movements allow access to higher quality resources, or reduction in predation risk resulting in increased fitness. However, most migratory species occur in partially migratory populations, a mix of migratory and non-migratory individuals. It is thought that the portion of migrants in a partial migration population is maintained either through (1) a population-level evolutionary stable state where counteracting density-dependent vital rates act on migrants and residents to balance fitness or (2) conditional migration, where the propensity to migrate is influenced by the individual's state. However, in many respects, migration is also a form of habitat selection and the proportion of migrants and residents may be the result of density-dependent habitat selection. Here, we test whether the theory of Ideal Free Distribution (IFD) can explain the coexistence of different migratory tactics in a partially migratory population. IFD predicts individuals exhibit density-dependent vital rates and select different migratory tactics to maximize individual fitness resulting in equal fitness (λ) between tactics. We tested the predictions of IFD in a partially migratory elk population that declined by 70% with 19 years of demographic data and migratory tactic switching rates from >300 individuals. We found evidence of density dependence for resident pregnancy and adult female survival providing a fitness incentive to switch tactics. Despite differences in vital rates between migratory tactics, mean λ (fitness) was equal. However, as predicted by the IFD, individuals switched tactics toward those of higher fitness. Our analysis reveals that partial migration may be driven by tactic selection that follows the ideal free distribution. These findings reinforce that migration across taxa may be a polymorphic behavior in large herbivores where migratory tactic selection is determined by differential costs and benefits, mediated by density dependence.

Beyond the encounter: Predicting multi-predator risk to elk (Cervus canadensis) in summer using predator scats.

There is growing evidence that prey perceive the risk of predation and alter their behavior in response, resulting in changes in spatial distribution and potential fitness consequences. Previous approaches to mapping predation risk across a landscape quantify predator space use to estimate potential predator-prey encounters, yet this approach does not account for successful predator attack resulting in prey mortality. An exception is a prey kill site that reflects an encounter resulting in mortality, but obtaining information on kill sites is expensive and requires time to accumulate adequate sample sizes.We illustrate an alternative approach using predator scat locations and their contents to quantify spatial predation risk for elk (Cervus canadensis) from multiple predators in the Rocky Mountains of Alberta, Canada. We surveyed over 1300 km to detect scats of bears (Ursus arctos/U. americanus), cougars (Puma concolor), coyotes (Canis latrans), and wolves (C. lupus). To derive spatial predation risk, we combined predictions of scat-based resource selection functions (RSFs) weighted by predator abundance with predictions that a predator-specific scat in a location contained elk. We evaluated the scat-based predictions of predation risk by correlating them to predictions based on elk kill sites. We also compared scat-based predation risk on summer ranges of elk following three migratory tactics for consistency with telemetry-based metrics of predation risk and cause-specific mortality of elk.We found a strong correlation between the scat-based approach presented here and predation risk predicted by kill sites and (r = .98, p < .001). Elk migrating east of the Ya Ha Tinda winter range were exposed to the highest predation risk from cougars, resident elk summering on the Ya Ha Tinda winter range were exposed to the highest predation risk from wolves and coyotes, and elk migrating west to summer in Banff National Park were exposed to highest risk of encountering bears, but it was less likely to find elk in bear scats than in other areas. These patterns were consistent with previous estimates of spatial risk based on telemetry of collared predators and recent cause-specific mortality patterns in elk.A scat-based approach can provide a cost-efficient alternative to kill sites of quantifying broad-scale, spatial patterns in risk of predation for prey particularly in multiple predator species systems.

Spatio-temporal changes in chronic wasting disease risk in wild deer during 14 years of surveillance in Alberta, Canada.

Disease risk modeling is a key first step to understand the spatio-temporal dynamics of wildlife disease and to direct cost-effective surveillance and management. In Alberta, active surveillance for chronic wasting disease (CWD) in wild cervids began in 1998 with the first case detected in free-ranging cervids in 2005. Following the detection, a herd reduction program was implemented during 2005-2008 and in 2006 the ongoing hunter-based CWD Surveillance Program became mandatory in high-risk Wildlife Management Units (WMU). We used data collected during the CWD surveillance program to 1) document growth in sex-specific CWD prevalence (proportion of deer in sample that is CWD-positive) in hunter-harvest deer in 6 WMUs consistently monitored from 2006 to 2018, 2) document landscape features associated with where CWD-positive compared to CWD-negative deer were removed during hunter harvest and herd reduction in an early (2005-2012) and in a late period (2013-2017), and 3) to map the spatial risk of harvesting a deer infected with CWD in the prairie parklands of Alberta. In the 6 continuously monitored WMUs, risk of a harvested deer being CWD positive increased from 2006 to 2018 with CWD prevalence remaining highest in male mule deer whereas overall growth rate in CWD prevalence was greater in female mule deer, but similar to male white-tailed deer. We found no evidence that the 3-year herd reduction program conducted immediately after CWD was first detected affected the rate at which CWD grew over the course of the invasion. Risk of deer being CWD-positive was the highest in animals taken near small stream drainages and on soils with low organic carbon content in the early period, whereas risk became highest in areas of agriculture especially when far from large river drainages where deer often concentrate in isolated woody patches. The change in the influence of proximity to known CWD-positive cases suggested the disease was initially patchy but became more spatially homogeneous over time. Our results indicate that a targeted-removal program will remove more CWD positive animals compared to hunter harvest. However, the discontinuation of targeted removals during our research program, restricted our ability to assess its long term impact on CWD prevalence.

Are migrant and resident elk (Cervus elaphus) exposed to similar forage and predation risk on their sympatric winter range?

Partially migratory populations, where one portion of a population conducts seasonal migrations (migrants) while the other remains on a single range (residents), are common in ungulates. Studies that assess trade-offs between migratory strategies typically compare the amount of predation risk and forage resources migrants and residents are exposed to only while on separate ranges and assume both groups intermix completely while on sympatric ranges. Here we provide one of the first tests of this assumption by comparing the amount of overlap between home ranges of GPS-collared migrant and resident elk and fine-scale exposure to wolf predation risk and forage biomass at telemetry locations on a sympatric winter range in west-central Alberta, Canada. Overlap between migrant and resident home ranges increased throughout the winter, and both groups were generally intermixed and exposed to equal forage biomass. During the day, both migrants and residents avoided predation risk by remaining in areas far from timber with high human activity, which wolves avoided. However, at night wolves moved onto the grasslands close to humans and away from timber. Resident elk were consistently closer to areas of human activity and further from timber than migrants, possibly because of a habituation to humans. As a result, resident elk were exposed to higher night-time predation risk than migrants. Our study does not support the assumption that migrant and resident elk are exposed to equal predation risk on their sympatric range when human presence alters predation risk dynamics and habituation to humans is unequal between migratory strategies.

Living with liver flukes: Does migration matter?

Migration is typically thought to be an evolved trait driven by responses to forage or predation, but recent studies have demonstrated avoidance of parasitism can also affect success of migratory tactics within a population. We evaluated hypotheses of how migration alters parasite exposure in a partially migratory elk (Cervus canadensis) population in and adjacent to Banff National Park, Alberta, Canada. Equal numbers of elk remain year-round on the winter range or migrate to summer range. We quantified diversity and abundance of parasites in faecal elk pellets, and prevalence (number of infected individuals) and intensity (egg counts) of giant liver fluke eggs (Fascioloides magna) in faeces across migratory tactics. We tested whether giant liver fluke intensity in faeces was affected by elk use of wetlands, elevation, forage biomass, and elk concentration in the previous summer. We rejected the "migratory escape" hypothesis that suggests migration allowed elk to escape parasite exposure because migrant elk had the highest richness and evenness of parasite groups. We also rejected the hypothesis that prevalence was highest at highest summer densities because higher-density resident elk had the lowest diversity and giant liver fluke egg presence and intensity. Instead, the high prevalence and intensity of giant liver flukes in migrants was consistent with both the hypothesis of "environmental tracking", because elk that migrated earlier may expose themselves to favourable parasite conditions, and with the "environmental sampling" hypothesis, because giant liver fluke intensity increased with increased exposure to secondary host habitat (i.e., wetland). Our results indicate that differential exposure of different migratory tactics that leave the winter range has a greater influence on parasites than the concentration of elk that reside on the winter range year-round.

Within-population diversity of bacterial microbiomes in winter ticks (Dermacentor albipictus).

The bacterial microbiome of ticks is notoriously diverse, but the factors leading to this diversity are poorly understood. We sequenced bacterial 16S rRNA amplicons from individual winter ticks, Dermacentor albipictus, to assess whether their one-host life cycle is associated with reduced bacterial diversity. On average, about 100 bacterial genera were found for individual ticks. Francisella-like endosymbiont (FLE) dominated bacterial communities, particularly in female ticks and in ticks that had fed. The remainder of the winter tick microbiome was highly variable. In addition to FLE, the main bacterial genera associated with winter ticks on elk were Pseudomonas, Ehrlichia, Asinibacterium, Acinetobacter and Streptococcus, although sequences associated with hundreds of other minor bacterial genera were detected. A complex interaction between richness and evenness was revealed in comparisons among tick life stages, using the Hill number series to show trends in diversity with decreasing emphasis on rare members of the assemblage. Male ticks had a significantly greater number of bacterial genera than females or nymphs, while males had greater evenness than females and similar evenness to nymphs. We intentionally sampled ticks from a single host species, North American elk, from a single location in Alberta, Canada, to constrain the ecological and blood meal variation that individuals experience through their life cycle. In spite of this, we found that the number of bacterial genera detected in this one-host tick system was remarkably diverse. The high taxonomic variability of the minor components of the winter tick microbiome suggests that this part of their microbiome diversity should be examined for functional significance.

Wave-like Patterns of Plant Phenology Determine Ungulate Movement Tactics.

Animals exhibit a diversity of movement tactics [1]. Tracking resources that change across space and time is predicted to be a fundamental driver of animal movement [2]. For example, some migratory ungulates (i.e., hooved mammals) closely track the progression of highly nutritious plant green-up, a phenomenon called "green-wave surfing" [3-5]. Yet general principles describing how the dynamic nature of resources determine movement tactics are lacking [6]. We tested an emerging theory that predicts surfing and the existence of migratory behavior will be favored in environments where green-up is fleeting and moves sequentially across large landscapes (i.e., wave-like green-up) [7]. Landscapes exhibiting wave-like patterns of green-up facilitated surfing and explained the existence of migratory behavior across 61 populations of four ungulate species on two continents (n = 1,696 individuals). At the species level, foraging benefits were equivalent between tactics, suggesting that each movement tactic is fine-tuned to local patterns of plant phenology. For decades, ecologists have sought to understand how animals move to select habitat, commonly defining habitat as a set of static patches [8, 9]. Our findings indicate that animal movement tactics emerge as a function of the flux of resources across space and time, underscoring the need to redefine habitat to include its dynamic attributes. As global habitats continue to be modified by anthropogenic disturbance and climate change [10], our synthesis provides a generalizable framework to understand how animal movement will be influenced by altered patterns of resource phenology.

Trade-offs between predation risk and forage differ between migrant strategies in a migratory ungulate.

Trade-offs between predation risk and forage fundamentally drive resource selection by animals. Among migratory ungulates, trade-offs can occur at large spatial scales through migration, which allows an "escape" from predation, but trade-offs can also occur at finer spatial scales. Previous authors suggest that ungulates will avoid predation risk at the largest scale, although few studies have examined multi-scale trade-offs to test for the relative benefits of risk avoidance across scales. Building on previously developed spatial models of forage and wolf predation risk, we tested for trade-offs at the broad landscape scale and at a finer, within-home-range scale for migratory and non-migratory resident elk (Cervus elaphus) during summer in the Canadian Rockies in Banff National Park (BNP) and adjacent Alberta, Canada. Migration reduced exposure to wolf predation risk by 70% relative to residents at the landscape scale; at the fine scale, migrants used areas that were, on average, 6% higher in forage digestibility. In contrast, by forgoing migration, resident elk were exposed to higher predation risk, but they reduced predation risk at fine scales to only 15% higher than migrants by using areas close to human activity, which wolves avoided. Thus, residents paid for trying to avoid predation risk with lower forage quality. Residents may have been able to compensate, however, by using areas of abundant forage close to human activity where they may have been able to forage more selectively while avoiding predation risk. Human activity effectively decoupled the positive correlation between high forage quality and wolf predation, providing an effective alternate strategy for residents, similar to recent findings in other systems. Although ungulates appear capable of balancing risk and forage at different spatial scales, risk avoidance at large landscape scales may be more effective in the absence of human-caused refugia from predation.

First passage time analysis of animal movement and insights into the functional response.

Movement plays a role in structuring the interactions between individuals, their environment, and other species. Although movement models coupled with empirical data are widely used to study animal distribution, they have seldom been used to study search time. This paper proposes first passage time as a novel approach for understanding the effect of the landscape on animal movement and search time. In the context of animal movement, first passage time is the time taken for an animal to reach a specified site for the first time. We synthesize current first passage time theory and derive a general first passage time equation for animal movement. This equation is related to the Fokker-Planck equation, which is used to describe the distribution of animals in the landscape. We illustrate the first passage time method by analyzing the effect of territorial behavior on the time required for a red fox to locate prey throughout its home range. Using first passage time to compute search times, we consider the effect of two different searching modes on a functional response. We show that random searching leads to a Holling type III functional response. First passage time analysis provides a new tool for studying how animal movement may influence ecological processes.

Polymorphisms at the PRNP gene influence susceptibility to chronic wasting disease in two species of deer (Odocoileus Spp.) in western Canada.

Chronic wasting disease (CWD) is increasingly prevalent in multiple wild mule (Odocoileus hemionus) and white-tailed deer (O. virginianus) herds in North America. CWD was first found in Canadian wild mule deer in Saskatchewan in 2000 and has since spread into the neighboring province of Alberta. The infectious agent for CWD is a misfolded prion protein encoded by the PRNP gene. Previous studies revealed association between PRNP genotype and susceptibility to CWD in both mule and white-tailed deer in other regions. To investigate this link in Canadian populations, PRNP gene sequence was examined in 166 CWD- and 83 CWD+ mule deer, and 197 CWD- and 30 CWD+ white-tailed deer from Saskatchewan and Alberta. Two variable sites were found in mule deer, and 15 in white-tailed deer. In both species PRNP genotype was associated with CWD status. Mule deer possessing at least one copy of the common allele at codon 20 were less likely to test positive for CWD than expected, given the frequency of this allele in the population. A variant at codon 96 in white-tailed deer was also linked with reduced incidence of CWD. A greater knowledge of the genetic sources of heterogeneity in CWD susceptibility may improve our understanding of the mechanisms underlying the CWD epidemic in western Canada.

Functional response of wolves to human development across boreal North America.

AIM: The influence of humans on large carnivores, including wolves, is a worldwide conservation concern. In addition, human-caused changes in carnivore density and distribution might have impacts on prey and, indirectly, on vegetation. We therefore tested wolf responses to infrastructure related to natural resource development (i.e., human footprint). LOCATION: Our study provides one of the most extensive assessments of how predators like wolves select habitat in response to various degrees of footprint across boreal ecosystems encompassing over a million square kilometers of Canada. METHODS: We deployed GPS-collars on 172 wolves, monitored movements and used a generalized functional response (GFR) model of resource selection. A functional response in habitat selection occurs when selection varies as a function of the availability of that habitat. GFRs can clarify how human-induced habitat changes are influencing wildlife across large, diverse landscapes. RESULTS: Wolves displayed a functional response to footprint. Wolves were more likely to select forest harvest cutblocks in regions with higher cutblock density (i.e., a positive functional response to high-quality habitats for ungulate prey) and to select for higher road density in regions where road density was high (i.e., a positive functional response to human-created travel routes). Wolves were more likely to use cutblocks in habitats with low road densities, and more likely to use roads in habitats with low cutblock densities, except in winter when wolves were more likely to use roads regardless of cutblock density. MAIN CONCLUSIONS: These interactions suggest that wolves trade-off among human-impacted habitats, and adaptively switch from using roads to facilitate movement (while also risking encounters with humans), to using cutblocks that may have higher ungulate densities. We recommend that conservation managers consider the contextual and interacting effects of footprints when assessing impacts on carnivores. These effects likely have indirect impacts on ecosystems too, including on prey species.

Functional connectivity in ruminants: A generalized state-dependent modelling approach.

Animal behaviour is increasingly seen as an important component in maintaining functional connectivity between patches in fragmented landscapes. However, models that explicitly incorporate behavioural trade-offs are rarely applied to landscape planning problems like connectivity. The aim of this study was to explore how state-dependent behaviour influenced functional connectivity between patches from a theoretical perspective. We investigated how inter-patch distances influenced functional connectivity using a dynamic state variable model framework. The decision making process of an individual ruminant facing fitness trade-offs in staying in its patch of origin or moving to another patch at various distances were explicitly modelled. We incorporated energetic costs and predation costs of feeding, ruminating, and resting while in the patch and for transit between patches based on inter-patch distance. Functional connectivity was maintained with isolated patches when they offered high intake and the inactivity of rumination associated with rapid gut fill resulted in reduced predation risk. Nevertheless, individuals in high energetic state often would forgo moving to another patch, whereas individuals in poor energetic states were forced to accept the cost of movement to best meet their requirements in the distant patch. The inclusion of state-dependent behavioural models provides important insights into functional connectivity in fragmented landscapes and helps integrate animal behaviour into landscape planning. We discuss the consequences of our findings for landscape planning to show how the approach provides a heuristic tool to assess alternative scenarios for restoring landscape functional connectivity.

Willow on Yellowstone's northern range: evidence for a trophic cascade?

Reintroduction of wolves (Canis lupus) to Yellowstone National Park in 1995-1996 has been argued to promote a trophic cascade by altering elk (Cervus elaphus) density, habitat-selection patterns, and behavior that, in turn, could lead to changes within the plant communities used by elk. We sampled two species of willow (Salix boothii and S. geyeriana) on the northern winter range to determine whether (1) there was quantitative evidence of increased willow growth following wolf reintroduction, (2) browsing by elk affected willow growth, and (3) any increase in growth observed was greater than that expected by climatic and hydrological factors alone, thereby indicating a trophic cascade caused by wolves. Using stem sectioning techniques to quantify historical growth patterns we found an approximately twofold increase in stem growth-ring area following wolf reintroduction for both species of willow. This increase could not be explained by climate and hydrological factors alone; the presence of wolves on the landscape was a significant predictor of stem growth above and beyond these abiotic factors. Growth-ring area was positively correlated with the previous year's ring area and negatively correlated with the percentage of twigs browsed from the stem during the winter preceding growth, indicating that elk browse impeded stem growth. Our results are consistent with the hypothesis of a behaviorally mediated trophic cascade on Yellowstone's northern winter range following wolf reintroduction. We suggest that the community-altering effects of wolf restoration are an endorsement of ecological-process management in Yellowstone National Park.

Territory surveillance and prey management: Wolves keep track of space and time.

Identifying behavioral mechanisms that underlie observed movement patterns is difficult when animals employ sophisticated cognitive-based strategies. Such strategies may arise when timing of return visits is important, for instance to allow for resource renewal or territorial patrolling. We fitted spatially explicit random-walk models to GPS movement data of six wolves (Canis lupus; Linnaeus, 1758) from Alberta, Canada to investigate the importance of the following: (1) territorial surveillance likely related to renewal of scent marks along territorial edges, to reduce intraspecific risk among packs, and (2) delay in return to recently hunted areas, which may be related to anti-predator responses of prey under varying prey densities. The movement models incorporated the spatiotemporal variable "time since last visit," which acts as a wolf's memory index of its travel history and is integrated into the movement decision along with its position in relation to territory boundaries and information on local prey densities. We used a model selection framework to test hypotheses about the combined importance of these variables in wolf movement strategies. Time-dependent movement for territory surveillance was supported by all wolf movement tracks. Wolves generally avoided territory edges, but this avoidance was reduced as time since last visit increased. Time-dependent prey management was weak except in one wolf. This wolf selected locations with longer time since last visit and lower prey density, which led to a longer delay in revisiting high prey density sites. Our study shows that we can use spatially explicit random walks to identify behavioral strategies that merge environmental information and explicit spatiotemporal information on past movements (i.e., "when" and "where") to make movement decisions. The approach allows us to better understand cognition-based movement in relation to dynamic environments and resources.