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Widely considered an anthropogenic phenomenon, antimicrobial resistance (AMR) is a naturally occurring mechanism that microorganisms use to gain competitive advantage. AMR represents a significant threat to public health and has generated criticism towards the overuse of antimicrobial drugs. Livestock have been proposed as important reservoirs for AMR accumulation. Here, we show that assemblages of AMR genes in cattle and ungulates from natural environments (Yellowstone and Rocky Mountain National Parks) are all dominated by genes conferring resistance to tetracyclines. However, cattle feces contained higher proportions of erm(A-X) genes conferring resistance to macrolide antibiotics. Medically important AMR genes differed between cattle and natural ungulates, but cumulatively were more predominant in natural soils. Our findings suggest that the commonly described predominance of tetracycline resistance in cattle feces is a natural phenomenon among multiple ungulate species and not solely a result of antimicrobial drug exposure. Yet, the virtual absence of macrolide resistance genes in natural ungulates suggests that macrolide usage in agriculture may enrich these genes in cattle. Our results show that antimicrobial use in agriculture may be promoting a potential reservoir for specific types of AMR (i.e., macrolide resistance) but that a significant proportion of the ungulate resistome appears to have natural origins.

Significance The Green Wave Hypothesis (GWH) says the green wave—the progression of spring green-up from low to high elevations or latitudes—dictates the pace of herbivore migrations worldwide. Animals move in sync with the wave because young vegetation provides the best forage. We show the GWH needs to be revised to include group-forming grazers that not only move to find forage, but create forage by how they move. Bison, by moving and grazing en masse, release themselves from the need to “surf the wave.” Their movements and grazing stimulate plant growth and delay plant maturation, which allows them to eat high-quality foods despite falling behind the wave while also modifying the progression of the green wave itself., Newly emerging plants provide the best forage for herbivores. To exploit this fleeting resource, migrating herbivores align their movements to surf the wave of spring green-up. With new technology to track migrating animals, the Green Wave Hypothesis has steadily gained empirical support across a diversity of migratory taxa. This hypothesis assumes the green wave is controlled by variation in climate, weather, and topography, and its progression dictates the timing, pace, and extent of migrations. However, aggregate grazers that are also capable of engineering grassland ecosystems make some of the world’s most impressive migrations, and it is unclear how the green wave determines their movements. Here we show that Yellowstone’s bison (Bison bison) do not choreograph their migratory movements to the wave of spring green-up. Instead, bison modify the green wave as they migrate and graze. While most bison surfed during early spring, they eventually slowed and let the green wave pass them by. However, small-scale experiments indicated that feedback from grazing sustained forage quality. Most importantly, a 6-fold decadal shift in bison density revealed that intense grazing caused grasslands to green up faster, more intensely, and for a longer duration. Our finding broadens our understanding of the ways in which animal movements underpin the foraging benefit of migration. The widely accepted Green Wave Hypothesis needs to be revised to include large aggregate grazers that not only move to find forage, but also engineer plant phenology through grazing, thereby shaping their own migratory movements.

1.Decades of grazing studies have identified a number of key plant and soil processes affected by large herbivores and how those grazer effects vary among different grassland types. However, there remains little mechanistic understanding about how the effects of grazers on plants and soils may be biogeochemically linked in regulating grassland processes. 2.Here we measured monthly plant and soil variables, including soil moisture, soil nitrogen (N) availability, plant biomass, shoot N concentration and plant production, in grazed and ungrazed (fenced) grasslands during the 2012-2014 growing seasons. Measurements were used to assess direct and indirect biogeochemical pathways by which grazers influenced net aboveground plant production (NAP) in dry and mesic grasslands in Yellowstone National Park (YNP). 3.Herbivores only had direct effects on plant variables at the dry grassland compared to direct and indirect effects on both plant and soil variables at the mesic grassland. By enhancing leaf N content at both grasslands, grazers shifted the resource controlling NAP from N in ungrazed grassland to moisture, and potentially phosphorus and/or other soil nutrients, in grazed grassland. 4.Synthesis. These results indicate the mechanistic linkage between top-down (herbivore) and bottom-up (soil resource) control of grassland production. Changing the resources that limit NAP likely has a profound impact on how grazed vs ungrazed YNP grasslands respond to environmental (e.g., climate, atmospheric N deposition) variability. Because grazing enhances leaf N among many types of grasslands, increasing the sensitivity of plant production to the availability of moisture and nutrients other than N may be a general response of grasslands to grazing. This article is protected by copyright. All rights reserved.

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.

Background The collection and analysis of fecal DNA is a common practice, especially when dealing with wildlife species that are difficult to track or capture. While fecal DNA is known to be lower quality than traditional sources of DNA, such as blood or other tissues, few investigations have verified fecal samples as a valid source of DNA by directly comparing the results to high quality DNA samples from the same individuals. Our goal was to compare DNA from fecal and blood samples from the same 50 American plains bison (Bison bison) from Yellowstone National Park, analyze 35 short tandem repeat (STR) loci for genotyping efficiency, and compare heterozygosity estimates. Results We discovered that some of the fecal-derived genotypes obtained were significantly different from the blood-derived genotypes from the same bison. We also found that fecal-derived DNA samples often underestimated heterozygosity values, in some cases by over 20%. Conclusions These findings highlight a potential shortcoming inherent in previous wildlife studies that relied solely on a multi-tube approach, using exclusively low quality fecal DNA samples with no quality control to account for false alleles and allelic dropout. Herein, we present a rigorous marker selection protocol that is applicable for a wide range of species and report a set of 15 STR markers for use in future bison studies that yielded consistent results from both fecal and blood-derived DNA. Electronic supplementary material The online version of this article (10.1186/s12863-019-0722-3) contains supplementary material, which is available to authorized users.

Whole-genome sequencing has provided fundamental insights into infectious disease epidemiology, but has rarely been used for examining transmission dynamics of a bacterial pathogen in wildlife. In the Greater Yellowstone Ecosystem (GYE), outbreaks of brucellosis have increased in cattle along with rising seroprevalence in elk. Here we use a genomic approach to examine Brucella abortus evolution, cross-species transmission and spatial spread in the GYE. We find that brucellosis was introduced into wildlife in this region at least five times. The diffusion rate varies among Brucella lineages (∼3 to 8 km per year) and over time. We also estimate 12 host transitions from bison to elk, and 5 from elk to bison. Our results support the notion that free-ranging elk are currently a self-sustaining brucellosis reservoir and the source of livestock infections, and that control measures in bison are unlikely to affect the dynamics of unrelated strains circulating in nearby elk populations., The role of wild elk in the spread and persistence of bovine brucellosis in the Great Yellowstone area is unclear. Here, Kamath et al. analyse the genomic sequences of 245 Brucella abortus isolates from elk, bison and cattle, supporting the idea that elk is an important reservoir and source of livestock infections.

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...

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.

Yellowstone National Park is home to one of the only plains bison populations that have continuously existed on their present landscape since prehistoric times without evidence of domestic cattle introgression. Previous studies characterized the relatively high levels of nuclear genetic diversity in these bison, but little is known about their mitochondrial haplotype diversity. This study assessed mitochondrial genomes from 25 randomly selected Yellowstone bison and found 10 different mitochondrial haplotypes with a haplotype diversity of 0.78 (± 0.06). Spatial analysis of these mitochondrial DNA (mtDNA) haplotypes did not detect geographic population subdivision (FST = -0.06, p = 0.76). However, we identified two independent and historically important lineages in Yellowstone bison by combining data from 65 bison (defined by 120 polymorphic sites) from across North America representing a total of 30 different mitochondrial DNA haplotypes. Mitochondrial DNA haplotypes from one of the Yellowstone lineages represent descendants of the 22 indigenous bison remaining in central Yellowstone in 1902. The other mitochondrial DNA lineage represents descendants of the 18 females introduced from northern Montana in 1902 to supplement the indigenous bison population and develop a new breeding herd in the northern region of the park. Comparing modern and historical mitochondrial DNA diversity in Yellowstone bison helps uncover a historical context of park restoration efforts during the early 1900s, provides evidence against a hypothesized mitochondrial disease in bison, and reveals the signature of recent hybridization between American plains bison (Bison bison bison) and Canadian wood bison (B. b. athabascae). Our study demonstrates how mitochondrial DNA can be applied to delineate the history of wildlife species and inform future conservation actions.

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.

Loss of genetic variation through genetic drift can reduce population viability. However, relatively little is known about loss of variation caused by the combination of fluctuating population size and variance in reproductive success in age structured populations. We built an individual-based computer simulation model to examine how actual culling and hunting strategies influence the effective population size (Ne) and allelic diversity in Yellowstone bison over 200 years (∼28 generations). The Ne for simulated populations ranged from 746 in stable populations of size 2000 up to 1165 in fluctuating populations whose census size fluctuates between 3000 and 3500 individuals. Simulations suggested that ∼93% of allelic diversity, for loci with five alleles will be maintained over 200 years if the population census size remains well above ∼2000 bison (and if variance in male reproductive success is high). However for loci with 20 alleles, only 83% of allelic diversity will be maintained over 200 years. Removal of only juveniles (calves and yearlings) resulted in longer generation intervals which led to higher maintenance of allelic diversity (96%) after 200 years compared to the culling of adults (94%) when the mean census size was 3250 (for loci with five alleles). These simulations suggest that fluctuations in population census size do not necessarily accelerate the loss of genetic variation, at least for the relatively large census size and growing populations such as in Yellowstone bison. They also suggest that the conservation of high allelic diversity (>95%) at loci with many alleles (e.g., ⩾5) will require maintenance of a populations size greater than approximately 3250 and removal of mainly or only juveniles.

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.

Summary 1. Disease management along the boundaries of wildlife reserves is a growing conservation problem worldwide, as infected wildlife can migrate outside protected areas and pose a threat to livestock and human health. The bison Bison bison population in Yellowstone National Park has long been infected with Brucella abortus, but culling of Yellowstone bison to prevent transmission to cattle has been ineffective at reducing brucellosis infection. This management strategy is negatively affecting long-term bison conservation because of difficulties in diagnosing actively infected animals. 2. We integrated age-specific serology and B. abortus culture results from slaughtered Yellowstone bison to estimate probabilities of active brucellosis infection using a Bayesian framework. Infection probabilities were associated with age in young bison (0–5 years old) and with elevated antibody levels in older bison (>5 years old). Our results indicate that Yellowstone bison acquire B. abortus infection early in life but typically recover as they grow older. 3. A tool was developed to allow bison management to better reflect the probability that particular animals are infective, with the aim of conserving Yellowstone bison while reducing the risk of brucellosis transmission to cattle. Combining selective removal of infectious bison with additional management practices, such as vaccination, has the potential to advance an effective brucellosis reduction programme. 4. Synthesis and applications. We conclude that active B. abortus infection in Yellowstone bison is age dependent, which allows true infection probabilities to be estimated based on age and quantitative diagnostic tests. These findings have important application to disease management worldwide where accurate diagnostic tests for wildlife are unavailable. Estimation of true infection probabilities can replace culling practices that conflict with wildlife conservation. The ability to identify infective individuals can improve management practices that support conservation, particularly when human health is at risk or endangered wildlife species are involved.

Yellowstone bison (Bison bison bison) are managed to reduce the risk of brucellosis (Brucella abortus) transmission to cattle while allowing some migration out of Yellowstone National Park to winter ranges in Montana. Intensive management near conservation area boundaries maintained separation between bison and cattle, with no transmission of brucellosis. However, brucellosis prevalence in the bison population was not reduced and the management plan underestimated bison abundance, distribution, and migration, which contributed to larger risk management culls (total >3000 bison) than anticipated. Culls differentially affected breeding herds and altered gender structure, created reduced female cohorts, and dampened productivity. The ecological future of plains bison could be significantly enhanced by resolving issues of disease and social tolerance for Yellowstone bison so that their unique wild state and adaptive capabilities can be used to synergize the restoration of the species. We recommend several adaptive management adjustments that could be implemented to enhance the conservation of plains bison and reduce brucellosis infection. These findings and recommendations are pertinent to wood bison (Bison bison athabascae), European bison (Bison bonasus), and other large ungulates worldwide that are managed using best practices within a risk framework.

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 ¼ 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 (N¼ 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.

The conservation of bison in Yellowstone National Park, from near extinction in the late 19th century to a recent high of 5000, has led to long-term societal conflict regarding perceived overabundance, trans-boundary movements, and potential transmission of brucellosis from bison to livestock. We synthesized available information to address two central questions in this debate: (1) has the Yellowstone bison population surpassed numbers that can be supported by the forage base in the park; and (2) why do some bison move outside the park during winter, even when numbers are below food-limited carrying capacity? A spatially-explicit model of the system that integrated abiotic variables with biotic processes indicated bison have not reached a theoretical food-limited carrying capacity of 6200 in Yellowstone National Park. However, more bison began to migrate earlier to lower-elevation winter ranges as numbers increased and climatic factors interacted with density to limit nutritional intake and foraging efficiency. A gradual expansion of the winter range as bison numbers increased enabled relatively constant population growth and increased food-limited carrying capacity. Current management actions attempt to preserve bison migration to essential winter range areas within and adjacent to the park, while actively preventing dispersal and range expansion to outlying areas via hazing and removals (i.e., dispersal sink). A population of 2500–4500 bison should satisfy collective interests concerning the park’s forage base, bison movement ecology, retention of genetic diversity, brucellosis risk management, and prevailing social conditions.

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.

The wildlife of the Greater Yellowstone Ecosystem carries brucellosis, which was first introduced to the area by cattle in the 19th century. Brucellosis transmission between wildlife and livestock has been difficult to study due to challenges in culturing the causative agent, Brucella abortus . We examined B. abortus transmission between American bison ( Bison bison ), Rocky Mountain elk ( Cervus elaphus nelsoni), and cattle ( Bos taurus ) using variable number tandem repeat (VNTR) markers on DNA from 98 B. abortus isolates recovered from populations in Idaho, Montana, and Wyoming, US. Our analyses reveal interspecies transmission. Two outbreaks (2007, 2008) in Montana cattle had B. abortus genotypes similar to isolates from both bison and elk. Nevertheless, similarity in elk and cattle isolates from the 2008 outbreak suggest that elk are the likely source of brucellosis transmission to cattle in Montana and Wyoming. Brucella abortus isolates from sampling in Montana appear to be divided in two clusters: one found in local Montana elk, cattle, and bison; and another found mainly in elk and a bison from Wyoming, which is consistent with brucellosis having entered Montana via migration of infected elk from Wyoming. Our findings illustrate complex patterns of brucellosis transmission among elk, bison, and cattle as well as the utility of VNTRs to infer the wildlife species of origin for disease outbreaks in livestock.

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

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.