Ryan K. Brook

Impacts of wildlife baiting and supplemental feeding on infectious disease transmission risk: A synthesis of knowledge

Baiting and supplemental feeding of wildlife are widespread, yet highly controversial management practices, with important implications for ecosystems, livestock production, and potentially human health. An often underappreciated threat of such feeding practices is the potential to facilitate intra- and inter-specific disease transmission. We provide a comprehensive review of the scientific evidence of baiting and supplemental feeding on disease transmission risk in wildlife, with an emphasis on large herbivores in North America. While the objectives of supplemental feeding and baiting typically differ, the effects on disease transmission of these practices are largely the same. Both feeding and baiting provide wildlife with natural or non-natural food at specific locations in the environment, which can result in large congregations of individuals and species in a small area and increased local densities. Feeding can lead to increased potential for disease transmission either directly (via direct animal contact) or indirectly (via feed functioning as a fomite, spreading disease into the adjacent environment and to other animals). We identified numerous diseases that currently pose a significant concern to the health of individuals and species of large wild mammals across North America, the spread of which are either clearly facilitated or most likely facilitated by the application of supplemental feeding or baiting. Wildlife diseases also have important threats to human and livestock health. Although the risk of intra- and inter-species disease transmission likely increases when animals concentrate at feeding stations, only in a few cases was disease prevalence and transmission measured and compared between populations. Mostly these were experimental situations under controlled conditions, limiting direct scientific evidence that feeding practices exacerbates disease occurrence, exposure, transmission, and spread in the environment. Vaccination programs utilizing baits have received variable levels of success. Although important gaps in the scientific literature exist, current information is sufficient to conclude that providing food to wildlife through supplemental feeding or baiting has great potential to negatively impact species health and represents a non-natural arena for disease transmission and preservation. Ultimately, this undermines the initial purpose of feeding practices and represents a serious risk to the maintenance of biodiversity, ecosystem functioning, human health, and livestock production. Managers should consider disease transmission as a real and serious concern in their decision to implement or eliminate feeding programs. Disease surveillance should be a crucial element within the long-term monitoring of any feeding program in combination with other available preventive measures to limit disease transmission and spread.

Transdisciplinary habitat models for elk and cattle as a proxy for bovine tuberculosis transmission risk.

Zoonotic diseases such as bovine tuberculosis (TB) that infect wildlife and livestock are particularly difficult to eradicate where wild animals make extensive use of agricultural landscapes. Transmission of TB between cattle (Bos taurus) and wild elk (Cervus elaphus) in southwestern Manitoba, Canada remains poorly understood but there is a risk when commingling occurs on summer pasture. Elk use of cattle summer pastures was assessed using ecological data (187 VHF and 25 GPS collared elk monitored over four years representing 8% of the elk population). Local knowledge was documented by conducting interviews and participatory mapping exercises with 86 cattle producers (98% of those within the study area). Of the 294 cattle pastures mapped by farmers, 13% were used by radio-collared elk, 38% were reported by farmers as being used by elk, and 42% were identified as used by elk when both when all datasets were combined. Cattle pastures that had been used by elk and those that had no elk were compared using binary logistic regression based on each of the three datasets (i.e. farmer observations, radio-collared elk on pasture, and combined dataset). For all three datasets, distance to protected area and proportion of forest cover on the cattle pasture were identified as the most and second most important predictor variables, respectively. There was strong agreement among the relative probabilities of elk occurrence on each pasture derived from the resource selection function (RSF) models developed using farmer interviews and elk collaring data. The farmer interview and collar datasets were then combined to generate a final integrated RSF map summarizing the probability of elk-cattle commingling and were contrasted over each of four cattle grazing seasons (spring, early summer, late summer, and autumn). These predictive maps indicate that use of cattle pastures by elk is extensive, particularly in spring and early summer. Farmer observations indicate that elk and cattle share water sources and livestock mineral supplements on pasture. Local knowledge and conventional ecological data complement and validate one another and help us better understand the temporospatial aspects of shared space use among wildlife and livestock and more generally the risks of disease transmission in agricultural landscapes.

Intranasal Administration of Xylazine to Reduce Stress in Elk Captured by Net Gun

Forty free-ranging elk (Cervus elaphus manitobensis) were captured by net gun in Riding Mountain National Park (Manitoba, Canada) during February 2002 and were administered either saline (control) or xylazine by the intranasal route, to evaluate the efficacy and benefit of intranasal xylazine to reduce stress. Elk that received xylazine had higher relaxation scores than control elk, and the onset of sedation occurred quickly, often <1 min. Serum concentrations of cortisol, creatine kinase, and γ-glutamyltransferase were lower in elk that received xylazine than in control elk. At the conclusion of handling, the intravenous administration of yohimbine quickly abolished the sedative effect of xylazine, which allowed elk to be released without concern of physical injury due to ataxia. The intranasal administration of xylazine can be used to reduce stress in wild animals under situations where they are being handled while physically restrained.

Temporal variation in site fidelity: scale-dependent effects of forage abundance and predation risk in a non-migratory large herbivore

Large herbivores are typically confronted by considerable spatial and temporal variation in forage abundance and predation risk. Although animals can employ a range of behaviours to balance these limiting factors, scale-dependent movement patterns are expected to be an effective strategy to reduce predation risk and optimise foraging opportunities. We tested this prediction by quantifying site fidelity of global positioning system-collared, non-migratory female elk (Cervus canadensis manitobensis) across multiple nested temporal scales using a long-established elk–wolf (Canis lupus) system in Manitoba, Canada. Using a hierarchical analytical approach, we determined the combined effect of forage abundance and predation risk on variation in site fidelity within four seasons across four nested temporal scales: monthly, biweekly, weekly, daily. Site fidelity of female elk was positively related to forage-rich habitat across all seasons and most temporal scales. At the biweekly, weekly and daily scales, elk became increasingly attached to low forage habitat when risk was high (e.g. when wolves were close or pack sizes were large), which supports the notion that predator-avoidance movements lead to a trade-off between energetic requirements and safety. Unexpectedly, predation risk at the monthly scale increased fidelity, which may indicate that elk use multiple behavioural responses (e.g. movement, vigilance, and aggregation) simultaneously to dilute predation risk, especially at longer temporal scales. Our study clearly shows that forage abundance and predation risk are important scale-dependent determinants of variation in site fidelity of non-migratory female elk and that their combined effect is most apparent at short temporal scales. Insight into the scale-dependent behavioural responses of ungulate populations to limiting factors such as predation risk and forage variability is essential to infer the fitness costs incurred.

Factors driving variation in movement rate and seasonality of sympatric ungulates

Abstract Defining biologically relevant seasons is a critical issue in the interpretation of animal space-use studies. Moreover, understanding the effects of extrinsic (e.g., predation risk) and intrinsic (e.g., age and sex) factors on individual differences in seasonal transition dates will deepen our understanding of the mechanisms driving animal movement and potentially population dynamics. We used nonlinear modeling of movement rate over time using global positioning system–collared nonmigratory elk (Cervus elaphus manitobensis) and white-tailed deer (Odocoileus virginianus) in southern Manitoba, Canada, to derive species- and sex-specific seasonal transition dates. In addition, we used variables known to influence timing of migration in migratory populations to explain individual differences in seasonal transition dates. We found ecologically important differences in start and length of seasons between male and female elk and white-tailed deer. Individual differences in seasonal transition dates were large, and could be explained by a combination of intrinsic and extrinsic factors. Age-class of the individual animal and elevation influenced timing of winter, spring, and date of parturition, whereas predation risk from wolves (Canis lupus) influenced onset of spring, summer, and autumn. Our findings suggest that similar extrinsic and intrinsic factors can influence both large- (i.e., migratory) and small-scale movement patterns and can be used effectively to empirically define biologically relevant seasons for sympatric large herbivores.

Density-Dependent Effects on Group Size Are Sex-Specific in a Gregarious Ungulate

Density dependence can have marked effects on social behaviors such as group size. We tested whether changes in population density of a large herbivore (elk, Cervus canadensis) affected sex-specific group size and whether the response was density- or frequency-dependent. We quantified the probability and strength of changes in group sizes and dispersion as population density changed for each sex. We used group size data from a population of elk in Manitoba, Canada, that was experimentally reduced from 1.20 to 0.67 elk/km2 between 2002 and 2009. Our results indicated that functional responses of group size to population density are sex-specific. Females showed a positive density-dependent response in group size at population densities ≥0.70 elk/km2 and we found evidence for a minimum group size at population density ≤0.70 elk/km2. Changes in male group size were also density-dependent; however, the strength of the relationship was lower than for females. Density dependence in male group size was predominantly a result of fusion of solitary males into larger groups, rather than fusion among existing groups. Our study revealed that density affects group size of a large herbivore differently between males and females, which has important implications for the benefits e.g., alleviating predation risk, and costs of social behaviors e.g., competition for resources and mates, and intra-specific pathogen transmission.

Spatial and Temporal Factors Influencing Sightability of Elk

ABSTRACT Few tracking studies consider seasonal changes in ability to re-sight wildlife, despite potential for biases in sightability to mislead our interpretation of models of movement and abundance. We developed seasonal sightability models based on visual observations of radio-collared elk (Cervus elaphus) in Manitoba, Canada, through 6 seasons. We located 377 elk 8,862 times using aerial telemetry from 2002 to 2009. We tested the hypothesis that sites where we were able to visually observe radio-collared elk during aerial telemetry differed from sites where collared elk were known to be present but could not be sighted. Relationships varied with season and elk sightability was influenced by forest type, habitat openness, distance to edge, and time of day. Our results confirm that observers have the highest probability of detecting elk in early and late winter. However, factors such as day length, which increases by 64% during this period, suggest that fewer impediments to detection exist in late winter. Our findings reinforce the need to account for seasonal as well as spatial changes in habitat-specific sightability models.

Juxtaposition between host population structures: implications for disease transmission in a sympatric cervid community

Sympatric populations of phylogenetically related species are often vulnerable to similar communicable diseases. Although some host populations may exhibit spatial structure, other hosts within the community may have unstructured populations. Thus, individuals from unstructured host populations may act as interspecific vectors among discrete subpopulations of sympatric alternate hosts. We used a cervid‐bovine tuberculosis (Mycobacterium bovis) system to investigate the landscape‐scale potential for bovine tuberculosis transmission within a nonmigratory white‐tailed deer (Odocoileus virginianus) and elk (Cervus canadensis) community. Using landscape population genetics, we tested for genetic and spatial structure in white‐tailed deer. We then compared these findings with the sympatric elk population that is structured and which has structure that correlates spatially and genetically to physiognomic landscape features. Despite genetic structure that indicates the white‐tailed deer population forms three sympatric clusters, the absence of spatial structure suggested that intraspecific pathogen transmission is not likely to be limited by physiognomic landscape features. The potential for intraspecific transmission among subpopulations of elk is low due to spatial population structure. Given that white‐tailed deer are abundant, widely distributed, and exhibit a distinct lack of spatial population structure, white‐tailed deer likely pose a greater threat as bovine tuberculosis vectors among elk subpopulations than elk.

Targeting hunter distribution based on host resource selection and kill sites to manage disease risk

Endemic and emerging diseases are rarely uniform in their spatial distribution or prevalence among cohorts of wildlife. Spatial models that quantify risk-driven differences in resource selection and hunter mortality of animals at fine spatial scales can assist disease management by identifying high-risk areas and individuals. We used resource selection functions (RSFs) and selection ratios (SRs) to quantify sex- and age-specific resource selection patterns of collared (n = 67) and hunter-killed (n = 796) nonmigratory elk (Cervus canadensis manitobensis) during the hunting season between 2002 and 2012, in southwestern Manitoba, Canada. Distance to protected area was the most important covariate influencing resource selection and hunter-kill sites of elk (AICw = 1.00). Collared adult males (which are most likely to be infected with bovine tuberculosis (Mycobacterium bovis) and chronic wasting disease) rarely selected for sites outside of parks during the hunting season in contrast to adult females and juvenile males. The RSFs showed selection by adult females and juvenile males to be negatively associated with landscape-level forest cover, high road density, and water cover, whereas hunter-kill sites of these cohorts were positively associated with landscape-level forest cover and increasing distance to streams and negatively associated with high road density. Local-level forest was positively associated with collared animal locations and hunter-kill sites; however, selection was stronger for collared juvenile males and hunter-killed adult females. In instances where disease infects a metapopulation and eradication is infeasible, a principle goal of management is to limit the spread of disease among infected animals. We map high-risk areas that are regularly used by potentially infectious hosts but currently underrepresented in the distribution of kill sites. We present a novel application of widely available data to target hunter distribution based on host resource selection and kill sites as a promising tool for applying selective hunting to the management of transmissible diseases in a game species.

Island use as an anti-predator tactic by parturient elk and nursery herds in Riding Mountain National Park, Manitoba

Wildlife Society Bulletin 2004, 32(4):1321–1324 Peer edited Elk (Cervus elaphus) breed seasonally to take advantage of fluctuations in forage quality and quantity (Sadlier 1987). A single, highly developed calf is produced annually, usually in late May or early June (Greer 1966), while forage quantity is increasing and forage quality reaches a maximum (Kittams 1953, Flook 1970, Collins and Urness 1983). Despite having access to the highest-quality forage, cows must balance increased foraging demands associated with high energetic costs of lactation (Carl and Robbins 1988) with protection of calves from predators. Elk are gregarious. However, great variation in degree of sociality exists between seasons, sexes, and populations (Peek 2003). Murie (1951) observed cows seeking seclusion from other elk before and during parturition. Calves hide for the first 18–20 days of their lives but associate with the cow for short periods to nurse (Lent 1974). Calves often join nursery herds a few weeks after parturition and remain in cow–calf groups of various sizes through the summer. Nursery herds reach their maximum size within 6 weeks after birth (Franklin and Lieb 1979). The size of these aggregations appears related to vegetation density, with the largest groups appearing in the most open habitats (Clutton-Brock 1974). A possible reason for variation in aggregation size is that large groups in open terrain provide individuals security from predators (Crook 1970, Kie 1999). In other words, group size substitutes for cover as an antipredator strategy. As a calf grows, attention from its mother normally decreases. Altmann (1952) and McCullough (1969) observed that cows repel strange calves and allow only their own to suckle. Predation on elk calves by wolves (Canis lupus), coyotes (C. latrans), black bears (Ursus americanus), and other carnivores can be significant during the calving and post-calving period (Houston 1978, Carbyn 1983, Gese and Grothe 1995, Smith 1996). Thus, predation can affect population dynamics of elk, especially through its influence on calf survival (Peek 2003). In nursery groups, calves typically stay close together while cows are feeding. Calf groups appear to maintain their collective focus on a single female for long periods (Geist 2002). Altmann (1952) and Harper et al. (1967) considered this a form of “babysitting.” However, Geist (2002) suggested this was an adaptive strategy, which reduces confusion when a dangerous situation occurs. Herein, we report 23 years of observational data, documenting use of a lake island by elk for cooperative rearing of calves. Sharing responsibilities of raising calves may increase feeding time of individual cows during the post-calving period and reduce risk of predation. Use of islands as calving and calfrearing habitat has been reported for caribou From the Field: Island use as an anti-predator tactic by parturient elk and nursery herds in Riding Mountain National Park, Manitoba