Rick L. Wallen

Covariates affecting spatial variability in bison travel behavior in Yellowstone National Park

Understanding mechanisms influencing the movement paths of animals is essential for comprehending behavior and accurately predicting use of travel corridors. In Yellowstone National Park (USA), the effects of roads and winter road grooming on bison (Bison bison) travel routes and spatial dynamics have been debated for more than a decade. However, no rigorous studies have been conducted on bison spatial movement patterns. We collected 121 380 locations from 14 female bison with GPS collars in central Yellowstone to examine how topography, habitat type, roads, and elevation affected the probability of bison travel year-round. We also conducted daily winter bison road use surveys (2003-2005) to quantify how topography and habitat type influenced spatial variability in the amount of bison road travel. Using model comparison techniques, we found the probability of bison travel and spatial distribution of travel locations were affected by multiple topographic and habitat type attributes including slope, landscape roughness, habitat type, elevation, and distances to streams, foraging areas, forested habitats, and roads. Streams were the most influential natural landscape feature affecting bison travel, and results suggest the bison travel network throughout central Yellowstone is spatially defined largely by the presence of streams that connect foraging areas. Also, the probability of bison travel was higher in regions of variable topography that constrain movements, such as in canyons. Pronounced travel corridors existed both in close association with roads and distant from any roads, and results indicate that roads may facilitate bison travel in certain areas. However, our findings suggest that many road segments used as travel corridors are overlaid upon natural travel pathways because road segments receiving high amounts of bison travel had similar landscape features as natural travel corridors. We suggest that most spatial patterns in bison road travel are a manifestation of general spatial travel trends. Our research offers novel insights into bison spatial dynamics and provides conceptual and analytical frameworks for examining movement patterns of other species.

Predicting Bison Migration out of Yellowstone National Park Using Bayesian Models

Long distance migrations by ungulate species often surpass the boundaries of preservation areas where conflicts with various publics lead to management actions that can threaten populations. We chose the partially migratory bison (Bison bison) population in Yellowstone National Park as an example of integrating science into management policies to better conserve migratory ungulates. Approximately 60% of these bison have been exposed to bovine brucellosis and thousands of migrants exiting the park boundary have been culled during the past two decades to reduce the risk of disease transmission to cattle. Data were assimilated using models representing competing hypotheses of bison migration during 1990–2009 in a hierarchal Bayesian framework. Migration differed at the scale of herds, but a single unifying logistic model was useful for predicting migrations by both herds. Migration beyond the northern park boundary was affected by herd size, accumulated snow water equivalent, and aboveground dried biomass. Migration beyond the western park boundary was less influenced by these predictors and process model performance suggested an important control on recent migrations was excluded. Simulations of migrations over the next decade suggest that allowing increased numbers of bison beyond park boundaries during severe climate conditions may be the only means of avoiding episodic, large-scale reductions to the Yellowstone bison population in the foreseeable future. This research is an example of how long distance migration dynamics can be incorporated into improved management policies.

Vaccination strategies for managing brucellosis in Yellowstone bison.

Concerns over migratory bison (Bison bison) at Yellowstone National Park transmitting brucellosis (Brucella abortus) to cattle herds on adjacent lands led to proposals for bison vaccination. We developed an individual-based model to evaluate how brucellosis infection might respond under alternate vaccination strategies, including: (1) vaccination of female calves and yearlings captured at the park boundary when bison move outside the primary conservation area; (2) combining boundary vaccination with the remote delivery of vaccine to female calves and yearlings distributed throughout the park; and (3) vaccinating all female bison (including adults) during boundary capture and throughout the park using remote delivery of vaccine. Simulations suggested Alternative 3 would be most effective, with brucellosis seroprevalence decreasing by 66% (from 0.47 to 0.16) over a 30-year period resulting from 29% of the population receiving protection through vaccination. Under this alternative, bison would receive multiple vaccinations that extend the duration of vaccine protection and defend against recurring infection in latently infected animals. The initial decrease in population seroprevalence will likely be slow due to high initial seroprevalence (40-60%), long-lived antibodies, and the culling of some vaccinated bison that were subsequently exposed to field strain Brucella and reacted positively on serologic tests. Vaccination is unlikely to eradicate B. abortus from Yellowstone bison, but could be an effective tool for reducing the level of infection. Our approach and findings have applicability world-wide for managers dealing with intractable wildlife diseases that cross wildlife-livestock and wildlife-human interfaces and affect public health or economic well-being.

State‐space modeling to support management of brucellosis in the Yellowstone bison population

The bison (Bison bison) of the Yellowstone ecosystem, USA, exemplify the difficulty of conserving large mammals that migrate across the boundaries of conservation areas. Bison are infected with brucellosis (Brucella abortus) and their seasonal movements can expose livestock to infection. Yellowstone National Park has embarked on a program of adaptive management of bison, which requires a model that assimilates data to support management decisions. We constructed a Bayesian state-space model to reveal the influence of brucellosis on the Yellowstone bison population. A frequency-dependent model of brucellosis transmission was superior to a density-dependent model in predicting out-of-sample observations of horizontal transmission probability. A mixture model including both transmission mechanisms converged on frequency dependence. Conditional on the frequency-dependent model, brucellosis median transmission rate was 1.87 yr−1. The median of the posterior distribution of the basic reproductive ratio (R0) was...

Integrating population‐ and individual‐level information in a movement model of Yellowstone bison

Throughout the world, fragmentation of landscapes by human activities has constrained the opportunity for large herbivores to migrate. Conflict between people and wildlife results when migrating animals transmit disease to livestock, damage property, and threaten human safety. Mitigating this conflict requires understanding the forces that shape migration patterns. Bison Bos bison migrating from Yellowstone National Park into the state of Montana during winter and spring concern ranchers on lands surrounding the park because bison can transmit brucellosis (Brucella abortus) to cattle. Migrations have been constrained, with bison being lethally removed or moved back into the park. We developed a state-space model to support decisions on bison management aimed at mitigating conflict with landowners outside the park. The model integrated recent GPS observations with 22 years (1990-2012) of aerial counts to forecast monthly distributions and identify factors driving migration. Wintering areas were located along decreasing elevation gradients, and bison accumulated in wintering areas prior to moving to areas progressively lower in elevation. Bison movements were affected by time since the onset of snowpack, snowpack magnitude, standing crop, and herd size. Migration pathways were increasingly used over time, suggesting that experience or learning influenced movements. To support adaptive management of Yellowstone bison, we forecast future movements to evaluate alternatives. Our approach of developing models capable of making explicit probabilistic forecasts of large herbivore movements and seasonal distributions is applicable to managing the migratory movements of large herbivores worldwide. These forecasts allow managers to develop and refine strategies in advance, and promote sound decision-making that reduces conflict as migratory animals come into contact with people.

Simulating sterilization, vaccination, and test-and-remove as brucellosis control measures in bison

Brucella abortus, the causative agent of bovine brucellosis, infects wildlife, cattle, and humans worldwide, but management of the disease is often hindered by the logistics of controlling its prevalence in wildlife reservoirs. We used an individually based epidemiological model to assess the relative efficacies of three management interventions (sterilization, vaccination, and test-and-remove). The model was parameterized with demographic and epidemiological data from bison in Yellowstone National Park, USA. Sterilization and test-and-remove were most successful at reducing seroprevalence when they were targeted at young seropositive animals, which are the most likely age and sex category to be infectious. However, these approaches also required the most effort to implement. Vaccination was less effective (even with a perfect vaccine) but also required less effort to implement. For the treatment efforts we explored (50-100 individuals per year or 2.5-5% of the female population), sterilization had little impact upon the bison population growth rate when selectively applied. The population growth rate usually increased by year 25 due to the reduced number of Brucella-induced abortions. Initial declines in seroprevalence followed by rapid increases (.15% increase in 5 years) occurred in 3-13% of simulations with sterilization and test-and-remove, but not vaccination. We believe this is due to the interaction of superspreading events and the loss of herd immunity in the later stages of control efforts as disease prevalence declines. Sterilization provided a mechanism for achieving large disease reductions while simultaneously limiting population growth, which may be advantageous in some management scenarios. However, the field effort required to find the small segment of the population that is infectious rather than susceptible or recovered will likely limit the utility of this approach in many free-ranging wildlife populations. Nevertheless, we encourage scientists and policy makers to consider sterilization as part of a suite of available brucellosis management tools.

Timing of parturition events in Yellowstone bison Bison bison: implications for bison conservation and brucellosis transmission risk to cattle

Abstract Yellowstone bison Bison bison are chronically infected with brucellosis (caused by the bacterium Brucella abortus), which raises concerns about possible transmission to cattle when they migrate to winter ranges outside the Yellowstone National Park. We monitored bison from April to mid-June during 2004-2007 to estimate the timing and location of parturition events that may shed tissues infected by B. abortus. Observed abortions (N = 29) occurred from January through 19 May, while peak calving (80% of births) occurred from 25 April to 26 May, and calving was finished by 5 June. Observed parturition events (N = 115) occurred in the Park and on the Horse Butte peninsula in Montana, USA, where cattle were not present at any time of the year. Allowing bison to occupy public lands outside the Park where cattle are never present (e.g. Horse Butte peninsula) until most bison calving is completed (late May or early June) is not expected to significantly increase the risk of brucellosis transmission from bison to cattle because: 1) bison parturition is essentially completed weeks before cattle occupy nearby ranges, 2) female bison meticulously consume birthing tissues, 3) ultraviolet light and heat degrade B. abortus on tissues, vegetation and soil, 4) scavengers remove fetuses and remaining birth tissues and 5) management maintains separation between bison and cattle on nearby ranges. Allowing bison to occupy public lands outside the Park through their calving season will help conserve bison migratory behaviour and reduce stress on pregnant females and their newborn calves, while still minimizing the risk of brucellosis transmission to cattle.

Yellowstone Bison—Should We Preserve Artificial Population Substructure or Rely on Ecological Processes?

Halbert et al. (2012) analyzed microsatellite genotypes collected from 661 Yellowstone bison sampled during winters from 1999 to 2003 and identified 2 genetically distinct subpopulations (central, northern) based on genotypic diversity and allelic distributions. On the basis of these findings, they raised concerns about the management and long-term conservation of Yellowstone bison because of disproportionate culling of the 2 subpopulations in some winters. The data and findings of Halbert et al. (2012) are significant and useful for managers charged with conserving these iconic wildlife. However, their article provides information regarding the behavior and management of Yellowstone bison that does not accurately portray historic or current conditions. This response clarifies those conditions and challenges some of their apparent deductions and recommendations. Halbert et al. (2012, p. 1) indicate that Yellowstone bison provide an opportunity to examine a “. . . natural population substructure, which could have important implications for the long-term evolution of these populations.” They assume “. . . the Yellowstone population was not subdivided before 1936” and that “these 2 subpopulations [central, northern] have differentiated in a relatively short period of approximately 8 generations [64 years]” (Halbert et al. 2012, p. 5, 7). However, these statements ignore that humans contributed to the observed population and genetic substructure in Yellowstone bison by nearly extirpating them in the late 19th century (except for approximately 23 bison that survived in central Yellowstone) and then by creating another breeding herd in northern Yellowstone at the turn of the 20th century from 21 bison of unrelated breeding descent and divergent genetic stock that were relocated from northern Montana and Texas (Meagher 1973). A few individuals from the endemic central herd were introduced into the northern herd in the early 1900s, whereas 71 bison from the northern herd were relocated to central Yellowstone during 1935–1936 (Cahalane 1944). The northern herd was not released from traditional livestock management practices and allowed to evolve natural patterns of distribution until the 1950s—which likely contributed to some geographic separation between the herds (Meagher 1973). Further, each herd was sporadically culled from the 1950s to present (Meagher 1973, White et al. 2011b). Thus, the history of Yellowstone bison suggests the population substructure and genetic differentiation was substantially influenced by a human-induced bottleneck in the late 1800s and the effects of human stewardship thereafter. As a result, there is evidence that the existing genetic substructure was artificially created. Halbert et al. (2012, p. 2,5) state that “Radiotelemetry data indicate the [central and northern] herds remain isolated during the summer breeding season” and “the number of migrants into and out of each subpopulation each generation is about 2 (Nm = 2.3) or approximately 1 every fourth year.” This statement and estimate may generally reflect conditions during the period of intense human stewardship (1900–1968) and subsequent increase in bison abundance and distribution during the period of ecological process management (Plumb et al. 2009). However, extensive monitoring of the movements and productivity of radio-collared bison since 2005, when the population reached an abundance of approximately 5000 bison, suggests that emigration and gene flow is now much higher. Since 2007 (one half of one generation), biologists have detected 17 radio-collared bison emigrating between the central and northern herds and remaining through one or more breeding seasons (see Supplementary Table 1 online). Female bison rarely travel alone, so dispersal by these marked females likely represents emigration in groups of 25–40 bison each time, which increases the probability that gene flow occurred. Eleven of these 17 radio-collared bison produced calves on their new range (e.g., northern) that were conceived on the range they left (e.g., central). At least 23 calves were produced by these dispersing bison through mating and calving on their new JHEREDOUPJournal Of HeredityJHERED0022-15031465-7333OUPUS

Yersinia enterocolitica: an unlikely cause of positive brucellosis tests in greater yellowstone ecosystem bison (Bison bison).

Yersinia enterocolitica serotype O:9 has identical O-antigens to those of Brucella abortus and has apparently caused false-positive reactions in numerous brucellosis serologic tests in elk (Cervus canadensis) from southwest Montana. We investigated whether a similar phenomenon was occurring in brucellosis antibody–positive bison (Bison bison) using Y. enterocolitica culturing techniques and multiplex PCR of four diagnostic loci. Feces from 53 Yellowstone bison culled from the population and 113 free-roaming bison from throughout the Greater Yellowstone Ecosystem (GYE) were tested. Yersinia enterocolitica O:9 was not detected in any of 53 the bison samples collected at slaughter facilities or in any of the 113 fecal samples from free-ranging bison. One other Y. enterocolitica serotype was isolated; however, it is not known to cause cross-reaction on B. abortus serologic assays because it lacks the perosamine synthetase gene and thus the O-antigens. These findings suggest that Y. enterocolitica O:9 cross-reactivity with B. abortus antigens is unlikely to have been a cause of false-positive serology tests in GYE bison and that Y. enterocolitica prevalence was low in bison in the GYE during this study.

Predator foraging response to a resurgent dangerous prey

Summary Prey switching occurs when a generalist predator kills disproportionately more of an abundant prey species and correspondingly spares a rarer species. Although this behaviour is a classic stabilizing mechanism in food web models, little is known about its operation in free-living systems which often include dangerous prey species that resist predation. We used long-term (1995–2015) data from a large mammal system in northern Yellowstone National Park, USA, to understand how prey preference of a wild, generalist predator (Canis lupus) responds to a shift in prey species evenness involving rising numbers of dangerous prey (Bison bison) and dropping numbers of relatively safer prey (Cervus elaphus). Contrary to the prey switching hypothesis, wolves attacked and killed disproportionately more of the rarer, but safer, species. Wolves maintained a strong preference against bison even when this species was nearly twice as abundant as elk. [Correction added after online publication on 26 April 2017: ‘more than’ changed to ‘nearly’]. There was also evidence that wolves were increasingly averse to hunting bison as relative bison abundance increased. Wolves seldom hunted bison because capture success was limited to a narrow set of conditions: larger packs (>11 wolves) chasing smaller herds (10–20 bison) with calves. Wolves scavenged bison carrion instead and did so more frequently as bison abundance increased. Our study demonstrates the overarching importance of prey vulnerability to understanding the prey preferences of generalist predators in ecological communities with dangerous prey. The formidable defences of such prey diminish the potential for switching and its stabilizing influence on population dynamics. In these communities, shifts from hunting to scavenging are perhaps more likely than shifts in prey preference. The assumption of switching may therefore overestimate the stability of multi-prey systems that include dangerous prey species. A lay summary is available for this article.