Digital Commons Network^™

Distribution And Prevalence Of Antibodies To Trichinella Spp. And Toxoplasma Gondii In Wild Pigs (Sus Scrofa) In The United States, Christopher A. Cleveland, Ellen Haynes, Katherine C. Callaghan, Alinde Fojtik, Sarah Coker, Emily Doub, Vienna R. Brown, Ania A. Majewska, Michael J. Yabsley

United States Department of Agriculture Wildlife Services: Staff Publications

Invasive wild pigs (Sus scrofa) are a reservoir for over 100 viral, bacterial, and parasitic pathogens that are transmissible to humans, livestock, domestic animals, and wildlife in North America. Numerous historical local surveys and results from a nation-wide survey (2006–2010) indicated that wild pigs in the United States act as reservoirs for Trichinella spp. and Toxoplasma gondii, two zoonotic pathogens of importance for human and animal health. Since that time, wild pig populations have expanded and increased in density in many areas. Population expansion of wild pigs creates opportunities for the introduction of pathogens to new areas …

Characterization Of Recent Wild Pig‑Vehicle Collisions In Georgia, Usa, Jeremiah L. Psiropoulos, Emily Howe, John J. Mayer, Sophie C. Mckee

United States Department of Agriculture Wildlife Services: Staff Publications

Vehicle collisions with wild pigs (Sus scrofa) are reported almost everywhere this species is found. However, this is one of the least studied and characterized forms of damage that these invasive animals cause in the United States (US). We analyzed 518 wild pig-vehicle collisions (WPVCs) that took place statewide in Georgia between 2015 and 2021. From that dataset, we analyzed several parameters in order to better understand and characterize these accidents on a scale that had previously not been done in the US. Wild pig-vehicle collisions were reported from 105 out of the 159 counties in Georgia, increasing …

Expanding Cwd Disease Surveillance Options Using Environmental Contamination At Deer Signposts, Miranda H. J. Huang, Steve Demarais, Alejandro Banda, Bronson K. Strickland, Anna Grace Welch, Scoty Hearst, Stuart Lichtenberg, Allan Houston, Kim M. Pepin

United States Department of Agriculture Wildlife Services: Staff Publications

1. Environmental surveillance can allow early detection of diseases, which increases management options and can improve disease trajectories. Chronic wasting disease (CWD) in cervids is a significant prion disease that has been spreading across North America since the 1960s, leading to cervid population declines and concern from hunters and state wildlife agencies. White-tailed deer have a unique breeding season behavior called scraping, where they deposit urine and saliva at shared sites. Since both these fluids can contain CWD prions, scrape sites have the potential to serve as sentinel sites for environmental surveillance of CWD.
2. To examine this potential, we …

Biodistribution Analysis Of Peptide-Coated Magnetic Iron Nanoparticles: A Simple And Quantitative Method, Pavithra Natarajan, Katherine Horak, Jennifer Rowe, Sungmin Yoon, Joshua Lingo, John M. Tomich, Sherry D. Fleming

United States Department of Agriculture Wildlife Services: Staff Publications

Biodistribution tracks compounds or molecules of interest in vivo to understand a compound’s anticipated efficacy and safety. Nanoparticles deliver nucleic acid and drug payloads and enhance tumor permeability due to multiple properties such as high surface area to volume ratio, surface functionalization, and modifications. Studying the in vivo biodistribution of nanoparticles documents the effectiveness and safety of nanoparticles and facilitates a more application-driven approach for nanoparticle development that allows for more successful translation into clinical use. In this study, we present a relatively simple method to determine the biodistribution of magnetic iron nanoparticles in mice. In vitro, cells take up …

African Swine Fever At The Wildlife-Livestock Interface: Challenges For Management And Outbreak Response Within Invasive Wild Pigs In The United States, Vienna R. Brown, Ryan S. Miller, Kim M. Pepin, Keith M. Carlisle, Merril A, Cook, Cole F. Vanicek, Lindsey K. Holmstrom, Lisa T. Rochette, Timothy J. Smyser

United States Department of Agriculture Wildlife Services: Staff Publications

African swine fever (ASF) causes significant morbidity and mortality in both domestic and wild suids (Sus scrofa), and disease outbreaks convey profound economic costs to impacted industries due to death loss, the cost of culling exposed/infected animals as the primary disease control measure, and trade restrictions. The co-occurrence of domestic and wild suids significantly complicates ASF management given the potential for wild populations to serve as persistent sources for spillover. We describe the unique threat of African swine fever virus (ASFV) introduction to the United States from epidemiological and ecological perspectives with a specific focus on disease management at the …

Overview Of Global Wildlife–Livestock Interfaces: The Array Of Conflicts And Strategies To Mitigate Them, Kurt C. Vercauteren, Stewart Breck

United States Department of Agriculture Wildlife Services: Staff Publications

Implications

Wildlife–livestock conflict can be broadly categorized as either competition for resources, disease transmission between wildlife and domestic species, or predation.
Management strategies can either focus on management of the livestock or management of the wildlife. Dynamic integration of both these strategies likely provides the best solution in many scenarios.
Reducing conflict and maintaining ecologically appropriate and viable populations of wildlife and thriving human endeavors should be the ultimate goal.

Use Of Rhodamine B As A Biomarker In A Simulated Oral Vaccine Deployment Against Bovine Tuberculosis In White-Tailed Deer, David Dressel, Kurt Vercauteren, Michael Lavelle, Nathan P. Snow, Henry Campa Iii

United States Department of Agriculture Wildlife Services: Staff Publications

Introduction Free-ranging white-tailed deer (Odocoileus virginianus) in northeastern lower Michigan, United States are a self-sustaining reservoir for bovine tuberculosis (bTB). Farm mitigation practices, baiting bans, and antlerless deer harvests have been ineffective in eliminating bTB in white-tailed deer and risks to cattle. The apparent prevalence has remained relatively constant in deer, prompting interest among wildlife researchers, managers, and veterinarians for an effective means of vaccinating deer against bTB. The commonly used human vaccine for bTB, Bacillus Calmette Guerin (BCG), is the primary candidate with oral delivery being the logical means for vaccinating deer.

Materials and methods We developed …

Identifying Priority Mitigation Areas For Human–Osprey Conflict, Natasha K. Murphy, Melanie R. Boudreau, Brian S. Dorr, Kate Slankard, Scott A. Rush

United States Department of Agriculture Wildlife Services: Staff Publications

Raptor nests on human‐built structures represent a significant source of conflict, as they can result in bird mortality, fires, or power outages due to falling nest materials or animals connecting with energized conductors. Power companies typically try to mitigate these conflicts to avoid service disruptions. Performing mitigation measures across all potentially problematic power infrastructure is generally not a practical solution given logistical, time, or budgetary constraints. Therefore, there is a need for quantitative, landscapescale tools to identify conflict risk and prioritize mitigation. We examined the influence of habitat and transmission infrastructure distribution on the potential risk of osprey (Pandion …

Estimating Landscape Spread Of A Low-Prevalence Disease Using Multiple Surveillance Methods, Abigail Feuka, Aakash Pandey, Melinda Cosgrove, Megan Moriarty, Anthony Duffiney, Henry Campa Iii, Kurt C. Vercuteren, Kimberly M. Pepin

United States Department of Agriculture Wildlife Services: Staff Publications

1. Diseases occurring at low prevalence in free-ranging wildlife can be difficult to study and predict at a landscape scale. This problem is compounded by imperfect pathogen detection.

2. We used data from multiple surveillance methods to determine factors influencing spread of a low-prevalence disease, bovine tuberculosis (bTB), among white-tailed deer (Odocoileus virginianus) in a bTB endemic area over a 10-year period. We fit dynamic occupancy models to predict spatial spread of bTB through a county in Michigan, USA, within the bTB endemic area, while accounting for imperfect detection and a suite of ecological processes potentially driving bTB …

Efficacy Of An Inflatable Deterrent For Reducing New World Vulture Human‑Wildlife Conflict, Bryan M. Kluever, Betsy A. Evans, Noah M. Osterhoudt, Eric A. Tillman

United States Department of Agriculture Wildlife Services: Staff Publications

Increasing urbanization coupled with spatial expansion and numerical increase of New World vulture populations has engendered a rise in human-vulture conflict, creating a need for effective tools to mitigate vulture-related damage. Visual frightening devices that mimic the presence of human or other predators can be employed in human-vulture conflict scenarios to increase perceived risk by the pest species, thereby eliciting an antipredator behavioral response, such as fleeing. One visual frightening device, inflatable scarecrows, recently proved effective at reducing passerine attendance at feral swine feeders, but their effectiveness when directed at other species and conflict scenarios has varied. Our primary objective …

Using Prescribed Fire And Biosolids Applications As Grassland Management Tools: Do Wildlife Respond?, Brian E. Washburn, Michael J. Begier

United States Department of Agriculture Wildlife Services: Staff Publications

Prescribed burning is a management tool commonly used in forested ecosystems in the southeastern United States, but the influence of this method on grassland vegetation and wildlife in this geographic region is unknown. During 2009–2015, we conducted a study to determine if the application of prescribed burning and/or long-term biosolid applications alter plant communities and/or wildlife use of grassland areas at Marine Corps Air Station Cherry Point, Havelock, North Carolina. We monitored vegetation growth, measured plant community composition, and documented wildlife activity in four study plots for 3 years after the implementation of annual winter prescribed burns. Prescribed burning reduced …

Effects Of Pregnancy Prevention On Brucella Abortus Shedding In American Bison (Bison Bison), Pauline Nol, Rebecca Frey, Morgan Wehtje, Jack C. Rhyan, Patrick Ryan Clarke, Matthew Mccollum, Christine Quance, Douglas C. Eckery, Suelee Robbe-Austerman

United States Department of Agriculture Wildlife Services: Staff Publications

Products of parturition are the predominant source of Brucella abortus for transmission in bison (Bison bison). Our objective was to assess whether preventing pregnancy in Brucella-seropositive bison reduced B. abortus shedding. Brucella-seropositive and -seronegative bison from Yellowstone National Park, Wyoming, USA were used in a replicated experiment. Each of two replicates (rep1, rep2) included a group of seropositive females treated with a single dose of gonadotropin-releasing hormone-based immunocontraceptive (Treatment rep1, n = 15; Treatment rep2, n = 20) and an untreated group (Control rep1, n = 14; Control rep2, …

Effects Of The Immunocontraceptive Gonacon On Pregnancy In Brucella-Seropositive American Bison (Bison Bison), Rebecca K. Frey, Morgan E. Wehtje, Pauline Nol, Patrick R. Clarke, Jack C. Rhyan, Matthew P. Mccollum, Lowell A. Miller, Douglas C. Eckery

United States Department of Agriculture Wildlife Services: Staff Publications

The purpose of this study was to determine if the number of pregnancies in naturally infected Brucella abortus-positive bison (Bison bison) cows would be reduced over a period of 5 yr after one treatment with 3000 mg gonadotropin-releasing hormone immunocontraceptive (GonaCon) compared to a similar group of naturally infected B. abortus-positive bison cows not treated with GonaCon. In each of the 5 yr, GonaCon-treated cows produced fewer offspring in relation to number of cows than the nontreated cows. Fisher’s Exact test comparing offspring produced during the first reproductive season showed a significant difference between the …

Antipredator Blood-Squirting Defense In Horned Lizards (Phrynosoma): Chemical Isolation Of Plasma Component(S), Pogonomyrmex Ant Dietary Origin, And Evolution, Wade C. Sherbrooke, Bruce A. Kimball

United States Department of Agriculture Wildlife Services: Staff Publications

North American horned lizards of the genus Phrynosoma are largely myrmecophagous and squirt systemic blood from circumocular sinuses during encounters with certain mammalian predators. The predators react with distinct revulsion. Using bioassay-guided fractionation of blood plasma, we endeavored to identify the source and characteristics of the active chemical compound(s) responsible for this negative oral response. Our results of coyote bioassays and a series of novel mouse bioassay experiments were largely concordant. The active compound(s) was likely plasma-borne with a molecular weight (mw) of between 800 and 1,600. To identify the source of the active compound(s), we noted bioassay responses …

Internal And External Factors Affect Pair Mate Proximity In Captive Coyotes, Canis Latrans, Alexandra Turano, Stacey P. Brummer, Jeffrey T. Schultz, Julie Young, Sara M. Freeman

United States Department of Agriculture Wildlife Services: Staff Publications

No abstract provided.

Toxoplasma Gondii Survey In Waterfowl And Gulls From Eight Usa States, Sawsan Ammar, Tania J. Dawant, Janetta Kelly, Graham Hickling, Justin Brown, Kyle R. Van Why, Rebecca Poulson, Chunlei Su, Richard W. Gerhold Jr.

United States Department of Agriculture Wildlife Services: Staff Publications

Sera from 391 waterbirds from eight USA states were tested for Toxoplasma gondii antibodies using the modified agglutination test. Fifteen different waterbird species (26.6%; n = 104) were seropositive. Of the adults, 25.4% (n = 52) showed a significantly higher T. gondii seroprevalence compared with juveniles (13.4%; n = 17); however, sex was not a significant factor.

Increased Bat Activity At Urban Water Sources: Implications For Cross-Species Transmission Of Bat Rabies To Mesocarnivores, Lias A. Hastings, Carol L. Chambers, David L. Bergman, Tad C. Theimer

United States Department of Agriculture Wildlife Services: Staff Publications

We examined the potential for urban water sources, specifically golf course ponds, to act as centers for rabies transmission from bats to mesocarnivores in the arid southwestern United States where surface water is often limited. Because residential housing can act as den and roost sites for both mesocarnivores and bats, we also examined the effect of housing density around water sources on activity. Using ultrasonic acoustic recorders to assess bat activity and camera traps to estimate mesocarnivore activity, we compared 14 pairs of wet and dry locations over two years by surveying twice during the summer, once prior to summer …

Experimental Assessment Of Laser Scarecrows For Reducing Avian Damage To Sweet Corn, Sean T. Manz, Kathryn E. Sieving, Rebecca N. Brown, Page E. Klug, Bryan M. Kluever

United States Department of Agriculture Wildlife Services: Staff Publications

Background

Birds damage crops, costing millions of dollars annually, and growers utilize a variety of lethal and nonlethal deterrents in an attempt to reduce crop damage by birds. We experimentally tested laser scarecrows for their effectiveness at reducing sweet corn (Zea mays) damage. We presented 18 captive flocks of free-flying European starlings (Sturnus vulgaris) with fresh sweet corn ears distributed on two plots where laser and control treatments were alternated each day and allowed each flock to forage over 5 days. In 16 trials, fresh sweet corn ears were mounted on wooden sticks distributed from 0 …

Unraveling Parasite Identification In Invasive And Endangered Squirrel Species: Potential For Parasite Spillover, Deandra Jones, Patricia S. Stock, Emilie Lefoulon, Maria V. Mazzamuto, W. Eugene Hall, David L. Bergman, John L. Kaprowski

United States Department of Agriculture Wildlife Services: Staff Publications

Invasive species can impact ecosystem health by introducing parasites during their establishment in new areas. However, the dynamics of parasite loads between invasive and native species remain poorly understood. In the 1940s, Abert’s squirrels (Sciurus aberti) were introduced to the Pinaleño Mountains in southeastern Arizona where they now co-occur with endemic endangered Mt. Graham red squirrels (Tamiasciurus fremonti grahamensis). The study aimed to identify and quantify parasites in both Abert’s and Mt. Graham red squirrels, with a focus on investigating possible parasite spillover between these two species and examining how host-related factors affect parasite presence. PCR …

Coyotes In The Great Basin Desert Do Not Exhibit A Spatial Response Following The Removal Of Anthropogenic Water Sources, Nadine A. Pershyn, Eric M. Gese, Erica F. Stuber, Bryan M. Kluever

United States Department of Agriculture Wildlife Services: Staff Publications

Coyote (Canis latrans) range expansion into desert ecosystems has highlighted the role of anthropogenic water sources in arid ecosystems. Despite hypotheses that additional water facilitated this expansion, previous studies reported that coyotes did not exhibit a spatial or dietary response to removal of anthropogenic water. We used GPS data to examine if coyotes responded to water removal at a finer spatial scale than previously investigated. Our integrated step selection analysis did not find evidence that coyotes adjusted their distance to water following water removal. Vegetation was an important factor in habitat selection of coyotes, with riparian and agricultural …

Frequency And Vehicle Damage Costs Of Wild Pig‑Vehicle Collisions In The United States, 2015–2022, Sophie C. Mckee, Jeremiah L. Psiropoulos, John J. Mayer

United States Department of Agriculture Wildlife Services: Staff Publications

The increasing frequency of wildlife-vehicle collisions (WVCs) is a globally recognized problem that can severely impact both the animals and humans involved in these accidents. Introduced wild pigs (Sus scrofa) have been present in various parts of the USA for centuries, but no study on wild pig-vehicle collisions (WPVCs) has been conducted on a national scale for the USA Using data provided by State Farm, we derived the number of vehicle collisions with wild pigs by state in the USA over the 2015–2022 period and the associated vehicle damage costs. In 2022, we estimated that a total of …

An Estimate Of The Marginal Annual Economic Contribution Of Wild-Pig Hunting In Texas, Sophie C. Mckee, Keith M. Carlisle, John Tomeček, Stephanie A. Shwiff, Rachael Connally, Maureen Frank, Mike Bodenchuk, Nicole Didero

United States Department of Agriculture Wildlife Services: Staff Publications

Context

Wild-pig (Sus scrofa) hunting is a culturally significant recreational and commercial activity in Texas, but the overall impacts are contested. Except for one 1980s study, there has been no research to formally quantify any economic benefits of wild-pig hunting in Texas, which is an important component for weighing the costs and benefits of wild-pig presence in the state.

Aims and methods

To fill that research gap, we surveyed a sample of resident and non-resident 2018–2019 hunting-license holders in Texas about various topics related to their wild-pig hunting activities, including expenditures

Key results

On the basis of the …

Rapid Point-Of-Care Testing For Detection Of Antibodies To Toxoplasma Gondii In Black Vultures And Ring-Billed Gulls From Pennsylvania, Alexa Rosypal Von Dohlen, Sydney Randall, Jordyan Grays, Mark A. Dugo, Brian J. Hunt, Justin Brown, Kyle Van Why, Sawsan Ammar, Richard Gerhold

United States Department of Agriculture Wildlife Services: Staff Publications

Toxoplasma gondii is a zoonotic protozoan parasite that infects most warm-blooded animals, including birds. Scavenging birds are epidemiologically important hosts because they can serve as indicators of environmental T. gondii levels. A rapid point-of-care (POC) test that detects antibodies to T. gondii in humans is commercially available. In this research, we assessed the ability of the human POC test to detect anti–T. gondii antibodies in 106 black vultures (Coragyps atratus) and 23 ring-billed gulls (Larus delawarensis) from Pennsylvania, USA. Serum samples were tested with the POC test and compared to the modified …

What Is Known, Unknown, And Needed To Be Known About Damage Caused By Wild Pigs, Kurt C. Vercauteren, Kim M. Pepin, Seth M. Cook, Sophie Mckee, Abigail Pagels, Keely J. Kohen, Ingrid A. Messner, Michael P. Glow, Nathan P. Snow

United States Department of Agriculture Wildlife Services: Staff Publications

Damage assessments provide evidence for initiating and evaluating management programs that protect natural resources and human livelihoods against invasive species. Wild pigs (Sus scrofa) cause extensive damage in their native and non-native ranges, but the extent of current damage and efficacy of management of the damage (i.e., population control of wild pigs, exclusion fencing, etc.) remains poorly described. We conducted a systematic review of physical damage caused by wild pigs to summarize what is known and identify knowledge gaps for damage assessment. Wild pig damage assessments have been overwhelmingly qualitative (84% of studies) and measured differently across studies, …

Use Of Nest Boxes By European Starlings (Sturnus Vulgaris): Effects Of Perceived Nest Predation Risk, Bradley F. Blackwell, Morgan B. Pfeiffer, Joshua L. Hoblet, Bruce N. Buckingham

United States Department of Agriculture Wildlife Services: Staff Publications

The European starling (Sturnus vulgaris) has expanded beyond its native Eurasian range, exploiting both natural cavities and human structures for nesting. We hypothesized that starling exposure to enhanced risk of nest predation at nest boxes (surrogates for nest sites in other structures), due to predator access, would negatively affect occupancy (establishment of a nest and ≥ 1 egg) and subsequent nest success. We also hypothesized that starlings would show no distinction in occupancy between nest boxes relative to the presence/absence of old nest material (i.e., material from the previous season), conditions that can contribute to nest predation risk. …

Discrimination Among Similarly Colored Goose Species In Federal Harvest Surveys, Joshua L. Dooley, Paul F. Doherty Jr., David L. Otis, Gary C. White, Daniel R. Taylor, Doreen L. Griffin, Syephen C. Chandler, Stephanie M. Catino, Kathy K. Fleming, Robert V. Raftovich, Antoinette J. Piaggio

United States Department of Agriculture Wildlife Services: Staff Publications

Each year in the United States, fall‐winter (sport) harvests of goose species are estimated from federal surveys coordinated by the United States Fish and Wildlife Service, including the Migratory Bird Harvest Survey to estimate total goose harvest and the Parts Collection Survey (PCS) to estimate the species and age composition. For the PCS, randomly selected hunters collect tail and wing feathers of each goose shot during the hunting season, and then biologists determine the age class and species of each sample at organized events (Wingbees) in each of the 4 flyways (Pacific, Central, Mississippi, and Atlantic). For similarly colored goose …

Population Growth Rates Of Pronghorn: Influence Of Temporally And Spatially Explicit Conditions, Density Dependence, And Scale, Eric M. Gese, Cole A. Bleke, Shane B. Roberts, Paul Atwood, Patricia A. Terletzky

United States Department of Agriculture Wildlife Services: Staff Publications

Density-independent and density-dependent population regulation has long been a subject of investigation. We examined density-dependent and density-independent factors on growth rates of pronghorn (Antilocapra americana (Ord, 1815)) using a retrospective analysis of population survey data. Across Idaho, we found that as the proportion of the subpopulation harvested the previous year increased, growth rates increased. Similarly, as fawn recruitment increased, growth rates increased. We also found when the growth rate in the previous year increased, the growth rate the subsequent year decreased. When subpopulations were examined independently, we found that in a low-elevation desert subpopulation, growth rates were influenced by …

Natural Selection On Feralization Genes Contributed To The Invasive Spread Of Wild Pigs Throughout The United States, Niek W. G. Barmentlo, Patrick G. Meirmans, William H. Stiver, Joseph G. Yarkovich, Blake E. Mccann, Antoinette J. Piaggio, Dominic Wright, Timothy J. Smyser, Mirte Bosse

United States Department of Agriculture Wildlife Services: Staff Publications

Despite a long presence in the contiguous United States (US), the distribution of invasive wild pigs (Sus scrofa × domesticus) has expanded rapidly since the 1980s, suggesting a more recent evolutionary shift towards greater invasiveness. Contemporary populations of wild pigs represent exoferal hybrid descendants of domestic pigs and European wild boar, with such hybridization expected to enrich genetic diversity and increase the adaptive potential of populations. Our objective was to characterize how genetic enrichment through hybridization increases the invasiveness of populations by identifying signals of selection and the ancestral origins of selected loci. Our study focused on invasive …

Downed Wood Removal Effects On Survival And Site Fidelity Of Woodrats In A California Oak Woodland, Timothy J. Smyser, Michael A. Hardy, Amy J. Davis, William L. Preston, William D. Tietje

United States Department of Agriculture Wildlife Services: Staff Publications

Owing to the increasing intensity and frequency of wildfires in the western United States, the removal of woody debris (downed dead wood and snags) from fire‐prone landscapes is being evaluated for wildfire mitigation. Consequently, the study of the ecological value of coarse woody debris to sustain dependent species has become of foremost importance. From 2004–2009, we used a before‐after control‐impact (BACI) study design to assess the effects of downed wood removal on a population of big‐eared woodrats (Neotoma macrotis) in an oak woodland (Quercus spp.) in coastal‐central California, USA. Using Pollock's robust design mark‐recapture analyses (with 12 …

Effects Of Lethal Management On Gray Wolf Pack Persistence And Reproduction In Wisconsin, Usa, Alejandra Zubiria Perez, Kenneth F. Kellner, David M. Macfarland, Jennifer L. Price Tack, David B. Ruid, Glenn E. Stauffer, Jerold L. Belant

United States Department of Agriculture Wildlife Services: Staff Publications

Direct human-caused mortality accounts for about half of all large mammal mortality in North America. For social species like gray wolves (Canis lupus), the death of pack members can disrupt pack structure and cause pack dissolution, and mortality of breeding adults or wolves during reproduction and pup-rearing can decrease pup recruitment. We estimated minimum and maximum probability of wolf pack persistence in Wisconsin, USA, during biological years (15 April–14 April) 2011–2019 and evaluated the influence of pack size and legal harvest mortality on pack persistence during 2012–2014. Harvests comprised 75–161 mortalities within 194 monitored packs during 2012–2014, with …