Black-tailed Prairie Dog Colonies Research Articles

Efficacy of electronet fencing for excluding coyotes: A case study for enhancing production of black-footed ferrets

Reducing coyote (Canis latrans) predation can be an important management objective. Here, we evaluated the efficacy of electronet fencing for excluding coyotes from focal areas on black tailed prairie dog(Cynomysludovicianus)colonies,measuredtheeffectoffencing on wild-born black-footed ferret (Mustela nigripes) kit survival, and modeled costs and benefits of fencing. From 27 July to 2 October 2010 in north- central Montana, USA, we erected and maintained 7.7km of electronet that enclosed 108ha on portions of 2 prairie dog colonies. We monitored 2 female ferrets and 6 kits inside exclosures and 3 females and 12 kits outside of exclosures. Percent of coyote sightings in the protected areas was 6 times less than expected during the exclosure period (42% pre-exclosure, 7% exclosure, 47% post-exclosure). We conclude that the electronet fencing was effective for dramatically decreasing coyote activity in focal areas where black-footed ferret litters were being raised. We found evidence that survival of kits living primarily in protected areas was 22% higher, but we qualify this finding because of low sample sizes and because our monitoring activity on the study site may have influenced coyote activity. We estimated one-time costs for fencing to be US$4,464/km and operation and/or maintenance costs for the 68 days of fence operation to be US$641/km. If fencing increased survival by 20-30%, then total cost per ferret kit not lost to coyote predation would range between US$5,400 and $3,600, or US$2,550 and $1,700 if fence set-up-take-down labor and use of an all-terrain vehicle were donated. Published 2013. This article is a U.S. Government work and is in the public domain in the USA.

Duration of Plague (Yersinia pestis) Outbreaks in Black-Tailed Prairie Dog (Cynomys ludovicianus) Colonies of Northern Colorado

Plague, caused by the bacterium Yersinia pestis, triggers die-offs in colonies of black-tailed prairie dogs (Cynomys ludovicianus), but the time-frame of plague activity is not well understood. We document plague activity in fleas from prairie dogs and their burrows on three prairie dog colonies that suffered die-offs. We demonstrate that Y. pestis transmission occurs over periods from several months to over a year in prairie dog populations before observed die-offs.

Bison Selection of Prairie Dog Colonies on Shortgrass Steppe

American bison (Bison bison L.) preferentially use black-tailed prairie dog (Cynomys ludovicianus L.) colonies over uncolonized range in the mixed-grass prairie of North America. To assess bison use of prairie dog colonies in a different ecosystem, the shortgrass steppe, this study was conducted at the Vermejo Park Ranch, New Mexico. Driving surveys were conducted in summer 2007 to determine the number of bison on and off prairie dog colonies in 2 pastures. Prairie dogs occupied 25.5% and 48.5% of the 2 pastures surveyed. Bison of both sexes used prairie dog colonies more than expected compared to a scenario of bison using colonies based on colony availability (P < 0.001). With the exception of bulls in one pasture, bison observed off-colony were more likely to be grazing than bison observed oncolony (P < 0.04). When selection of prairie dog colonies was assessed for only those bison observed grazing, bulls in both pastures and cows in one pasture used prairie dog colonies more than expected based on availability (P ≤ 0.001). Forage quality was superior on prairie dog colonies, with crude protein higher (P < 0.001) and acid detergent fiber lower (P < 0.001) in blue grama (Bouteloua gracilis [Willd. ex Kunth] Lag. ex Griffiths) collected on-colony than offcolony. Further studies under a variety of conditions are still needed, but selection for prairie dog colonies by bison at Vermejo, combined with findings from previous studies, suggests that during the growing season, bison and cattle (Bos taurus L.) might select prairie dog colonies over uncolonized range in both the northern and southern extents of North American grasslands.

Estimation of black-tailed prairie dog colonies in Montana

Black-tailed prairie dog (Cynomys ludovicianus) is an influential species in prairie ecosystems. Accurate estimates of active prairie dog colony area are needed to assess the status of the species and evaluate the effects of management actions. In 2008, Montana Fish, Wildlife and Parks conducted a survey of potential black-tailed prairie dog habitat. Using fixed-wing aircraft and an aerial line-intercept method, we surveyed 771 transects totaling 56,530 km in 32 counties in central and eastern Montana, USA, excluding tribal lands. We recorded 667 black-tailed prairie dog colony intercepts totaling 336,636 m in 21 counties. Ground intercepts were 1.091 (95% credible intervals = 1.087–1.094) times longer than air intercepts. The estimated percent of colonies classified as active from the air that were active on the ground was 86.8% (95% credible intervals = 77.9–93.5%). Corrected estimates resulted in 77,430 ha (95% credible intervals = 69,480–83,380) of active and 12,990 ha (95% credible intervals = 7,039–20,970) of inactive black-tailed prairie dog colonies. We conducted a sensitivity analysis of the estimated area of active prairie dog colonies by reducing a percentage of long intercepts assumed to be entirely active. More than 30% of active intercepts >750 m in length would need to, in fact, be inactive in order for our active colony area estimates to differ >10,000 ha (13%; i.e., outside of our margin of error) from our estimates. Aerial line-intercept methods provide a reliable and repeatable method for obtaining estimates of active and inactive prairie dog colony area over large areas. Our estimates provide the basis for long-term monitoring of prairie dogs on a landscape scale. © 2013 The Wildlife Society.

COMPETITION AND FACILITATION BETWEEN PRAIRIE DOGS AND LIVESTOCK

All photographs by David Augustine. Black-tailed prairie dog on a colony in the Conata Basin of Buffalo Gap National Grassland, South Dakota, USA. Cattle grazing on a black-tailed prairie dog colony at the Buffalo Gap National Grassland, which is adjacent to the Badlands National Park visible in the background. Cattle grazing on a black-tailed prairie dog colony at the Pawnee National Grassland, Colorado. In the Great Plains of North America, black-tailed prairie dogs are often thought to compete with cattle for forage. We examined how prairie dogs affect the amount and quality of forage growing on their colonies in Colorado and South Dakota. At a site experiencing below-average rainfall, forage both on and off colonies was so short and sparse that cattle were likely to be limited by the rate at which they could consume forage, rather than the rate at which they could digest it. In contrast, at sites experiencing above-average rainfall, forage both on and off colonies was tall and dense enough for cattle to be limited by digestion rather than the rate of forage consumption. Overall, our findings show prairie dogs can both enhance and suppress the weight gains of cattle depending on local weather conditions and the proportion of a pasture that is occupied by prairie dogs. Cattle grazing off of a black-tailed prairie dog colony at the Buffalo Gap National Grassland, South Dakota, with Badlands National Park as the backdrop. These photographs illustrate the article “Competition and facilitation between a native and a domestic herbivore: trade-offs between forage quantity and quality,” by David J. Augustine and Tim L. Springer, scheduled for publication in Ecological Applications 23(4), June 2013.

Mountain Plover responses to deltamethrin treatments on prairie dog colonies in Montana

Pyrethroid insecticides containing deltamethrin provide broad spectrum insect control that can adversely affect food supplies of insectivorous birds. I hypothesized that this could result in lowered nest survival for a ground-nesting insectivorous bird, the Mountain Plover (Charadrius montanus), which preferentially nests on prairie dog colonies. I studied Mountain Plover nest survival in 2003-2010 at a small cluster of black-tailed prairie dog (Cynomys ludovicianus) colonies in north-central Montana. Three colonies were treated with deltamethrin to control fleas and limit the spread of plague; four untreated colonies served as controls. I monitored 412 plover nests during the 8 year study (264 on treatment colonies and 148 on control colonies) and found a strong negative effect of deltamethrin treatments on nest survival (β(Dust) = -1.24, 95 % CI was -2.00 to -0.48) in the years following the actual treatment (2004-2006). I conclude that the observed treatment effect most likely occurred because of changes in insect (food) availability for the plover, and this in turn lowered nest survival because adults spent more time off nests or switched to less desirable insect prey. These results lend support to the need to consider the indirect effects of insecticide treatments on non-target species and suggest a potential conflict in current plague management strategies for prairie dogs.

Chlorophacinone residues in mammalian prey at a black-tailed prairie dog colony

Black-tailed prairie dogs (BTPDs), Cynomys ludovicianus, are an important prey for raptors; therefore, the use of the rodenticide Rozol (0.005% chlorophacinone active ingredient) to control BTPDs raises concern for secondary poisonings resulting from the consumption of contaminated prey by raptors. In the present study, the authors observed Rozol exposure and adverse effects to mammalian prey on 11 of 12 search days of the study. Mammalian hepatic chlorophacinone residues ranged from 0.44 to 7.56 µg/g. Poisoned prey availability was greater than previously reported.

Patterns of Surface Burrow Plugging in a Colony of Black-Tailed Prairie Dogs Occupied by Black-Footed Ferrets

Black-tailed prairie dogs (Cynomys ludovicianus) can surface-plug openings to a burrow occupied by a black-footed ferret (Mustela nigripes). At a coarse scale, surface plugs are more common in colonies of prairie dogs occupied by ferrets than in colonies without ferrets. However, little is known about spatial and temporal patterns of surface plugging in a colony occupied by ferrets. In a 452-ha colony of black-tailed prairie dogs in South Dakota, we sampled burrow openings for surface plugs and related those data to locations of ferrets observed during spotlight surveys. Of 67,574 burrow openings in the colony between June and September 2007, 3.7% were plugged. In a colony-wide grid of 80 m × 80 m cells, the occurrence of surface plugging (≥1 opening plugged) was greater in cells used by ferrets (93.3% of cells) than in cells not observably used by ferrets (70.6%). Rates of surface plugging (percentages of openings plugged) were significantly higher in cells used by ferrets (median = 3.7%) than in cells without known ferret use (median = 3.2%). Also, numbers of ferret locations in cells correlated positively with numbers of mapped surface plugs in the cells. To investigate surface plugging at finer temporal and spatial scales, we compared rates of surface plugging in 20-m-radius circle-plots centered on ferret locations and in random plots 1–4 days after observing a ferret (Jun–Oct 2007 and 2008). Rates of surface plugging were greater in ferret-plots (median = 12.0%) than in random plots (median = 0%). For prairie dogs and their associates, the implications of surface plugging could be numerous. For instance, ferrets must dig to exit or enter plugged burrows (suggesting energetic costs), and surface plugs might influence microclimates in burrows and consequently influence species that cannot excavate soil (e.g., fleas that transmit the plague bacterium Yersinia pestis).

Black-Footed Ferret Digging Activity in Summer

Black-footed ferrets (Mustela nigripes) excavate soil from prairie dog (Cynomys spp.) burrows, thereby creating characteristic soil deposits at burrow openings. These soil deposits have been observed only rarely in summer. We monitored adult ferrets during June–October of the years 2007 and 2008 on a 452-ha colony of black-tailed prairie dogs (Cynomys ludovicianus) in the Conata Basin, South Dakota. We located and identified ferret excavations during nighttime spotlight surveys for ferrets and daytime sampling of prairie dog burrow openings around locations where ferrets were located via spotlight. We accumulated 48 observations of in-process or recently completed ferret excavations during spotlight surveys (21 in 2007, 27 in 2008) and located 51 diggings during daytime burrow sampling (25 in 2007, 26 in 2008). We located diggings during 5.5% of spotlight observations, most frequently in July–August. These results collectively suggest ferrets may frequently excavate soil in summer, because prairie dogs frequently use soil to plug burrow openings and tunnels in defense against ferrets. Prairie dogs might frequently destroy soil deposits left by ferrets during summer, thereby reducing detection of diggings by biologists.

Resource Selection Models are Useful in Predicting Fine-Scale Distributions of Black-Footed Ferrets in Prairie Dog Colonies

Wildlife-habitat relationships are often conceptualized as resource selection functions (RSFs)—models increasingly used to estimate species distributions and prioritize habitat conservation. We evaluated the predictive capabilities of 2 black-footed ferret (Mustela nigripes) RSFs developed on a 452-ha colony of black-tailed prairie dogs (Cynomys ludovicianus) in the Conata Basin, South Dakota. We used the RSFs to project the relative probability of occurrence of ferrets throughout an adjacent 227-ha colony. We evaluated performance of the RSFs using ferret space use data collected via postbreeding spotlight surveys June–October 2005–2006. In home ranges and core areas, ferrets selected the predicted “very high” and “high” occurrence categories of both RSFs. Count metrics also suggested selection of these categories; for each model in each year, approximately 81% of ferret locations occurred in areas of very high or high predicted occurrence. These results suggest usefulness of the RSFs in estimating the distribution of ferrets throughout a black-tailed prairie dog colony. The RSFs provide a fine-scale habitat assessment for ferrets that can be used to prioritize releases of ferrets and habitat restoration for prairie dogs and ferrets. A method to quickly inventory the distribution of prairie dog burrow openings would greatly facilitate application of the RSFs.

Fine-Scale Habitat Use of Reintroduced Black-Footed Ferrets on Prairie Dog Colonies in New Mexico

ABSTRACT. Black-footed ferrets (Mustela nigripes) are among the most endangered animals in North America. Reintroductions of captive-born ferrets onto prairie dog (Cynomys spp.) colonies are crucial to the conservation of the species. In September 2007, captive-born ferrets were released on a black-tailed prairie dog (Cynomys ludovicianus) colony at the Vermejo Park Ranch, New Mexico. Ferret kits experimentally released in areas of comparatively low and high prairie dog burrow densities were located via spotlight surveys. Some maturing ferret kits were subsequently translocated to areas of low and high burrow densities on nearby prairie dog colonies. For 2 months, fine-scale habitat use was quantified by mapping all burrow openings within a 30-m radius of each ferret location. Spatial statistics accounted for autocorrelation in the burrow densities in areas used by ferrets. It was hypothesized that ferrets would select areas of high burrow densities within colonies; however, burrow densities in areas used...

Comparison of bird and mammal communities on black-tailed prairie dog (Cynomys ludovicianus) colonies and uncolonized shortgrass prairie in New Mexico

Black-tailed prairie dogs (BTPD; Cynomys ludovicianus) have often been labeled as keystone species because of their ability to strongly influence grassland ecosystems. I used line-transect surveys and distance sampling to compare breeding bird and mammal communities on shortgrass prairie occupied by BTPD colonies versus similar uncolonized habitat in New Mexico, and to identify species that were either strongly associated with, or that avoided, BTPD colonies. Overall, I detected 32 bird and 8 mammal species during three years. Mountain plover (Charadrius montanus), ferruginous hawk (Buteo regalis), burrowing owl (Athene cunicularia), curve-billed thrasher (Toxostoma curvirostre), desert cottontail (Sylvilagus audubonii), and American badger (Taxidea taxus) were more abundant on, or at least strongly associated with, colonies, while long-billed curlew (Numenius americanus), horned lark (Eremophila alpestris), vesper sparrow (Poecetes gramineus), lark sparrow (Chondestes grammacus), Cassin's sparrow (Aimophila cassinii), and western meadowlark (Sturnella neglecta) were more abundant on, or strongly associated with, uncolonized prairie. Observed responses of several species differed from other studies suggesting that a species' response to BTPD activities may vary by location, grassland type, or season. Although BTPDs negatively impacted a suite of grassland bird species, biodiversity is maximized in this landscape by maintaining a mixture of colonized and uncolonized habitats.

Deltamethrin flea-control preserves genetic variability of black-tailed prairie dogs during a plague outbreak

Genetic variability and structure of nine black-tailed prairie dog (BTPD, Cynomys ludovicianus) colonies were estimated with 15 unlinked microsatellite markers. A plague epizootic occurred between the first and second years of sampling and our study colonies were nearly extirpated with the exception of three colonies in which prairie dog burrows were previously dusted with an insecticide, deltamethrin, used to control fleas (vectors of the causative agent of plague, Yersinia pestis). This situation provided context to compare genetic variability and structure among dusted and non-dusted colonies pre-epizootic, and among the three dusted colonies pre- and post-epizootic. We found no statistical difference in population genetic structures between dusted and non-dusted colonies pre-epizootic. On dusted colonies, gene flow and recent migration rates increased from the first (pre-epizootic) year to the second (post-epizootic) year which suggested dusted colonies were acting as refugia for prairie dogs from surrounding colonies impacted by plague. Indeed, in the dusted colonies, estimated densities of adult prairie dogs (including dispersers), but not juveniles (non-dispersers), increased from the first year to the second year. In addition to preserving BTPDs and many species that depend on them, protecting colonies with deltamethrin or a plague vaccine could be an effective method to preserve genetic variability of prairie dogs.

Mortality of Siberian polecats and black-footed ferrets released onto prairie dog colonies

Black-footed ferrets (Mustela nigripes) likely were extirpated from the wild in 1985–1986, and their repatriation depends on captive breeding and reintroduction. Postrelease survival of animals can be affected by behavioral changes induced by captivity. We released neutered Siberian polecats (M. eversmanii), close relatives of ferrets, in 1989–1990 on black-tailed prairie dog (Cynomys ludovicianus) colonies in Colorado and Wyoming initially to test rearing and reintroduction techniques. Captive-born polecats were reared in cages or cages plus outdoor pens, released from elevated cages or into burrows, and supplementally fed or not fed. We also translocated wild-born polecats from China in 1990 and released captive-born, cage-reared black-footed ferrets in 1991, the 1st such reintroduction of black-footed ferrets. We documented mortality for 55 of 92 radiotagged animals in these studies, mostly due to predation (46 cases). Coyotes (Canis latrans) killed 31 ferrets and polecats. Supplementally fed polecats survived longer than nonprovisioned polecats. With a model based on deaths per distance moved, survival was highest for wild-born polecats, followed by pen-experienced, then cage-reared groups. Indexes of abundance (from spotlight surveys) for several predators were correlated with mortality rates of polecats and ferrets due to those predators. Released black-footed ferrets had lower survival rates than their ancestral population in Wyoming, and lower survival than wild-born and translocated polecats, emphasizing the influence of captivity. Captive-born polecats lost body mass more rapidly postrelease than did captive-born ferrets. Differences in hunting efficiency and prey selection provide further evidence that these polecats and ferrets are not ecological equivalents in the strict sense.

Postbreeding resource selection by adult black-footed ferrets in the Conata Basin, South Dakota

We investigated postbreeding resource selection by adult black-footed ferrets (Mustela nigripes) on a 452-ha black-tailed prairie dog (Cynomys ludovicianus) colony in the Conata Basin of South Dakota during 2007–2008. We used resource selection functions (RSFs) to evaluate relationships between numbers of ferret locations and numbers of prairie dog burrow openings (total or active), distances to colony edges, and connectivity of patches of burrow openings. In both years ferrets selected areas near edges of the prairie dog colony where active burrow openings were abundant. In the interior of the colony ferrets selected areas with low abundance of active burrow openings. At times, prairie dog productivity (i.e., pup abundance) might be greatest at colony edges often characterized by grasses; ferrets are likely to select areas where refuge and vulnerable prey are abundant. Ferrets could have used interior areas with few active burrow openings as corridors between edge areas with many active burrow openings. ...

Nighttime Aboveground Movements by Prairie Dogs on Colonies Inhabited by Black-Footed Ferrets

We describe observations of black-tailed prairie dogs (Cynomys ludovicianus) emerging aboveground at night, apparently in response to wild-born and captive-born black-footed ferrets (Mustela nigripes) in South Dakota and New Mexico, respectively, We also discuss other similar observations accumulated on black-tailed prairie dog colonies as well as observations of white-tailed prairie dogs (Cynomys leucurus) making nighttime movements, apparently in response to pre-reintroduction ferrets in Wyoming. Our observations suggest that, in addition to documented daytime defenses against ferrets, prairie dogs reduce vulnerability to predation by ferrets by using evasive movements at night.

Population genetic structure of black-tailed prairie dogs, a highly interactive species, in fragmented urban habitat

Fragmentation of wildlife populations can have detrimental effects, including genetic differentiation of populations, loss of genetic diversity, and inbreeding depression. We evaluated the genetic structure among isolated colonies of black-tailed prairie dogs (Cynomys ludovicianus) along an urban gradient in southern Denver, Colorado. Urban colonies are important ecologically and for educational purposes, and they serve as source populations for relocation efforts. Levels of genetic differentiation between colonies were high relative to colonies in natural habitat at comparable or greater distances. Prairie dog colonies depend on dispersal among colonies for long-term persistence, and we found evidence for reduced but measurable rates of movement of individual prairie dogs among urban fragments. We observed a trend for smaller and more isolated colonies to exhibit lower genetic diversity, but we did not detect inbreeding in any of the colonies sampled. Isolation-bydistance measures, including measures based on permeability of various features of urban habitat such as roads and development, did not explain genetic differences. Our system represents a possible end point in the genetic consequences of continued loss and isolation of prairie dog colonies as fragmentation increases in both urban and natural landscapes. Urban development could affect dispersal in unexpected and complex ways and requires further study, but prairie dog colonies and their associated wildlife communities in urban areas have the potential for long-term persistence if not extirpated by human activity. DOI: 10.1644/09-MAMM-A-019.1.

Detection of Yersinia pestis DNA in Prairie Dog–Associated Fleas by Polymerase Chain Reaction Assay of Purified DNA

We evaluated, refined, and applied well-established polymerase chain reaction (PCR) techniques for detecting Yersinia pestis DNA in fleas (mainly Oropsylla spp.) collected from prairie dog (Cynomys spp.) burrows. Based on results from PCR of avirulent Y. pestis strain A1122 DNA, we used DNA purification and primers for the plasminogen activator gene to screen field-collected fleas. We detected Y. pestis DNA in flea pools from two black-tailed prairie dog (Cynomys ludovicianus) colonies with evidence of recent plague epizootics, and from one of four white-tailed prairie dog (Cynomys leucurus) colony complexes (Wolf Creek) where evidence of epizootic plague was lacking. Relative flea abundance and occurrence of Y. pestis DNA among flea pools appeared to vary over time at Wolf Creek. Both DNA purification and primer sequences appeared to influence the likelihood of detecting Y. pestis DNA by PCR in fleas collected from prairie dog burrows in the absence of observed epizootic plague. Presence of Y. pestis plasmid DNA in fleas collected from prairie dog burrows at Wolf Creek may represent evidence that infected fleas were somehow being maintained in that system between epizootics, consistent with the hypothesized enzootic maintenance of plague in prairie dog colony complexes elsewhere.

Evaluation of Physical Barriers to Prevent Prairie Dog Colony Expansion

Expansion of black-tailed prairie dog (Cynomys ludovicianus) colonies on public and private lands can result in damage to property. Physical barriers to prairie dogs can be used to minimize human-wildlife confl icts. We evaluated 17 existing barriers in the Fort Collins and Boulder, Colorado, areas. Most barriers were made of a single row of vinyl material; these barriers sustained high levels of damage, primarily from wind, and were frequently breached by prairie dogs digging underneath them. Barriers that included a vegetation and a vinyl barrier or a double-vinyl barrier were wind damaged and breached less frequently than the single- vinyl barriers. Sturdy panels of corrugated metal or fi berglass, extending about 76 cm above and 76 cm below the ground surface, were not damaged by wind and were rarely breached by prairie dogs. These barriers were about twice the cost of the single-vinyl barriers, but were much more durable and more effective in preventing prairie dog colony expansion.

Mountain Plover Responses to Plague in Montana

Plague is a bacterial (Yersinia pestis) disease that causes epizootic die-offs in black-tailed prairie dog (Cynomys ludovicianus) populations in the North American Great Plains. Through their grazing and burrowing, prairie dogs modify vegetation and landscape structure on their colonies in ways that affect other grassland species. Plague epizootics on prairie dog colonies can have indirect effects on species associated with colonies. The mountain plover (Charadrius montanus) preferentially nests on black-tailed prairie dog colonies and is thus negatively impacted by the loss of prairie dogs. We studied the effects of plague and colony spatial characteristics on the occupancy of 81 prairie dog colonies by nesting plovers in Phillips County, Montana, during a 13-year period (1995-2007). We used a robust design patch occupancy model to investigate how colony occupancy and extinction and colonization rates were affected by plague history, colony size, and colony shape. Here extinction and colonization rates refer to the probability that a colony loses/gains plovers in a subsequent nesting season, given that it had/lacked plovers in that breeding season. Colony occupancy was best explained by a model with no annual variation or plague effects. Colony extinction rates were driven by a combination of a quadratic of colony area, a 3-year plague response, and a measure of colony shape. Conversely, colonization rates were best explained by a model with a 4-year plague response. The estimated annual proportion of colonies occupied by plovers was 0.75 (95% confidence interval = 0.57-0.87). Estimated extinction probability ranged from a low of 0.07 (standard error [SE] = 0.02) in 2002 to a high of 0.25 (SE = 0.03) in 1995; colonization probability ranged from 0.24 (SE = 0.05) in 2006 to 0.35 (SE = 0.05) in 2000. Our results highlight how a bird that depends on prairie dogs for nesting habitat responds to plague history and other spatial characteristics of the colony. Ultimately, this information will broaden our understanding of the impact of plague on species other than the host.