https://cpw.cvlcollections.org/files/original/995013c365a462e083b3aae34e8cde75.pdf dc0e0c295eb68f50f17334c040ffb952 PDF Text Text The research in this publication was partially or fully funded by Colorado Parks and Wildlife. Dan Prenzlow, Director, Colorado Parks and Wildlife • Parks and Wildlife Commission: Marvin McDaniel, Chair • Carrie Besnette Hauser, Vice-Chair Marie Haskett, Secretary • Taishya Adams • Betsy Blecha • Charles Garcia • Dallas May • Duke Phillips, IV • Luke B. Schafer • James Jay Tutchton • Eden Vardy �Received: 24 April 2019 | Revised: 15 July 2019 | Accepted: 19 July 2019 DOI: 10.1002/ece3.5559 ORIGINAL RESEARCH Human–Cougar interactions in the wildland–urban interface of Colorado's front range Mathew W. Alldredge1 1 | Frances E. Buderman2 | Kevin A. Blecha3 Colorado Parks and Wildlife, Fort Collins, CO, USA Abstract 2 As human populations continue to expand across the world, the need to understand Colorado State University, Fort Collins, CO, USA 3 Colorado Parks and Wildlife, Gunnison, CO, USA Correspondence Mathew W. Alldredge, Colorado Parks and Wildlife, 317 W. Prospect Rd., Fort Collins, CO 80526, USA. Email: mat.alldredge@state.co.us Funding information CPW game cash; Federal Aid and manage wildlife populations within the wildland–urban interface is becoming commonplace. This is especially true for large carnivores as these species are not always tolerated by the public and can pose a risk to human safety. Unfortunately, information on wildlife species within the wildland–urban interface is sparse, and knowledge from wildland ecosystems does not always translate well to human‐domi‐ nated systems. Across western North America, cougars (Puma concolor) are routinely utilizing wildland–urban habitats while human use of these areas for homes and recreation is increasing. From 2007 to 2015, we studied cougar resource selection, human–cougar interaction, and cougar conflict management within the wildland– urban landscape of the northern Front Range in Colorado, USA. Resource selec‐ tion of cougars within this landscape was typical of cougars in more remote settings but cougar interactions with humans tended to occur in locations cougars typically selected against, especially those in proximity to human structures. Within higher housing density areas, 83% of cougar use occurred at night, suggesting cougars gen‐ erally avoided human activity by partitioning time. Only 24% of monitored cougars were reported for some type of conflict behavior but 39% of cougars sampled during feeding site investigations of GPS collar data were found to consume domestic prey items. Aversive conditioning was difficult to implement and generally ineffective for altering cougar behaviors but was thought to potentially have long‐term benefits of reinforcing fear of humans in cougars within human‐dominated areas experiencing little cougar hunting pressure. Cougars are able to exploit wildland–urban landscapes effectively, and conflict is relatively uncommon compared with the proportion of cougar use. Individual characteristics and behaviors of cougars within these areas are highly varied; therefore, conflict management is unique to each situation and should target individual behaviors. The ability of individual cougars to learn to exploit these environments with minimal human–cougar interactions suggests that maintaining This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. © 2019 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd. Ecology and Evolution. 2019;9:10415–10431. � www.ecolevol.org | 10415 �10416 | ALLDREDGE et al. older age structures, especially females, and providing a matrix of habitats, including large connected open‐space areas, would be beneficial to cougars and effectively reduce the potential for conflict. KEYWORDS aversive conditioning, Colorado, conflict, Cougar, domestic predation, human interaction, livestock, predation, Puma concolor, residential development, wildland–urban interface 1 | I NTRO D U C TI O N Colorado protected cougars and began to actively manage for the sustainability of cougar populations in 1965, and by 1973, almost Human beings dominate the world, and their influences on ecosys‐ all of the western states were managing for cougar populations. tems and wildlife populations are well documented (Ellis, Goldewijk, Presumably, cougar populations began to increase and possibly re‐ Siebert, Lightman, & Ramankutty, 2010; Pickett et al., 2001; occupy historic ranges following this protection (Anderson, Lindzey, Theobald, 2005; Vitousek, Mooney, Lubchenco, & Melillo, 1997), but Knopff, Jalkotzy, & Boyce, 2010). Coinciding with this was the rapid the impact on wildlife communities in the wildland–urban interface expansion of human populations. In Colorado, much of this expan‐ is not well understood, especially with regard to carnivores (Kertson, sion occurred in mountainous regions that were prime cougar habi‐ Spencer, Marzluff, Hepinstall‐Cymerman, & Grue, 2011). Population tat. Expanding human and cougar populations throughout the West viability may be threatened by human development for some wild‐ have inevitably led to increasing cougar–human interactions in the life species (Hansen et al., 2005), but in other cases human develop‐ late 1900s and early 2000s (Beier, Riley, & Sauvajot, 2010), including ment may result in ecosystem alterations that are beneficial to other in Colorado (Halfpenny, Sanders, & McGrath, 1991). This significant species (Blecha, Boone, & Alldredge, 2018; Gehrt, Riley, & Cypher, increase in interaction during this time period led Colorado to initi‐ 2010). Cougars likely represent the nexus of these two forces, as the ate research to better understand this dynamic. wildland–urban interface represents a riskier environment where Knowledge of cougar space and resource use in the wildland–ur‐ survival is typically lower, but also represents a beneficial environ‐ ban interface is limited. Other research has documented decreased ment with alternate and consistent prey resources (Blecha et al., use of areas with increased human density (Beier & Barrett, 1993; 2018; Moss, Alldredge, & Pauli, 2016). Burdett et al., 2010; Dickson & Beier, 2002), but this does not ad‐ Carnivores are unique among wildlife as they tend to be highly dress how cougars use these areas. Kertson, Spencer, Marzluff, et al. adaptable and in general can exploit food resources within the wild‐ (2011) examined space use of cougars in Western Washington within land–urban interface, but they also generate considerable public at‐ a wildland–urban interface and found that space use and movement tention with regard to human safety (Gehrt et al., 2010). The ability rates within the urban areas were similar to those in wildland areas. of carnivores to exploit these wildland–urban interfaces has been They further suggest that the probability of human–cougar interac‐ documented for cougars (Burdett et al., 2010; Kertson, Spencer, & tions is maximized at a threshold residential density that modifies Grue, 2013), bobcats (Lynx rufus; Donovan et al., 2011), black bears available habitat but maintains enough wildland habitat to encour‐ (Ursus americanus; Baruch‐Mordo et al., 2014, Lewis et al., 2015), and age moderate levels of cougar use. In the Santa Cruz Mountains coyotes (Canis latrans; Gehrt, Anchor, & White, 2009, Poessel, Breck, of California, neighborhoods were found to be a deterrent to cou‐ & Gese, 2016). Unfortunately, as carnivores utilize the wildland–ur‐ gars for all behaviors or activities (Wilmers et al., 2013). Movement ban interface, and prey species increasingly inhabit urban areas, rates of cougars have also been shown to increase with increasing human–carnivore interactions will occur. human density (Buderman, Hooten, Alldredge, Hanks, & Ivan, 2018; Understanding cougar–human interactions requires an under‐ Wang, Smith, & Wilmers, 2017). Several studies have documented standing of the historical treatment of cougars throughout the his‐ an increase in use of smaller‐bodied prey, synanthropic wildlife, and tory of human settlement in North America. Cougars once occupied domestic pets in wildland–urban areas (Kertson, Spencer, Marzluff, the majority of North and South America, inhabiting most habitats et al., 2011), including two studies on the Front Range of Colorado from Northern British Columbia, south to Patagonia and from the (Blecha et al., 2018; Moss et al., 2016). Blecha et al. (2018) suggested Pacific to the Atlantic oceans (Rabinowitz, 2010). As North America that the utilization of higher housing density areas and, by exten‐ was settled, prime cougar habitat and prey were lost and persecu‐ sion, use of smaller‐bodied prey by cougars, is directly related to the tion of cougars began as livestock were killed and public percep‐ length of time since a cougar last fed. However, there is much more tions of cougars villainized. Throughout much of the 20th century, to be learned about cougar space use and movement patterns within a bounty was paid for cougars and they were killed wantonly across the wildland–urban interface. This includes the timing of use, where North America and completely removed from much of their historic human–cougar interactions occur relative to how cougars and hu‐ range (Rabinowitz, 2010). Even in the western states where cougars mans use the landscape, and management prescriptions to mitigate persisted, numbers were greatly reduced. these interactions. �| ALLDREDGE et al. 10417 In general, ungulates are the primary prey for cougars (Iriarte, human safety. Defining acceptable levels of human safety is difficult Franklin, Johnson, & Redford, 1990; Murphy & Ruth, 2010), except because people's perceptions are different when interactions do not in localities near housing development in which the cougar's di‐ directly affect them. Other difficulties associated with managing etary composition shifts to a higher reliance on smaller‐bodied prey cougar populations in areas with high levels of human interaction are (Kertson Spencer & Grue, 2011; Moss et al., 2016; Smith, Wang, & caused by the limited amount of information that is currently known Wilmers,2016). Many synanthropic wildlife species, such as raccoons about cougars in these exurban situations and responses of cougars (Procyon lotor), and domestic animals are found at higher densities to management prescriptions (CMGWG, 2005). in developed areas (Bateman & Fleming, 2012), which may explain Management plans generally require the removal of cougars that this switch to smaller‐bodied prey items, as they represent a con‐ represent a danger to human health and safety, but the appropriate sistent and abundant food resource. The use of alternate prey spe‐ management response to cougars that are overly familiar or habit‐ cies within these developed areas also includes the use of domestic uated to humans is unclear. Lethal control is losing public support animals, including hobby livestock, dogs (Canis familiarus), and cats (Reiter, Brunson, & Schmidt, 1999), so other options need to be ex‐ (Felis catus). The use of domestic animals by cougars has a high po‐ amined. Shivik and Martin (2000) and van Eeden et al. (2018) empha‐ tential to be a source of human–cougar conflict, especially livestock size the need to research and determine effective nonlethal control as this is usually detected by owners compared with pets that just techniques, or managers risk losing credibility with the public. go missing. In addition, the state of Colorado is statutorily respon‐ We define aversive conditioning as applying a negative stimu‐ sible for livestock damage from cougars ( 33‐3‐104) and therefore lus when an animal exhibits an undesirable behavior in an attempt must monetarily compensate owners when cougars take livestock. to modify or abolish that behavior. Aversive conditioning has been Understanding how cougars utilize domestic animals and whether attempted for a variety of species, including black bears, with mixed cougars are habituated to preying on domestics or if this is oppor‐ results (Beckman, Lackey, & Berger, 2004; Homstol, 2011; Leigh, tunistic feeding is important for understanding the wildland–urban 2007; Mazur, 2010; Rauer, Kaczensky, & Knauer, 2003). In these ex‐ dynamic for cougars. amples, specific behaviors are generally targeted, such as the use of Although cougar attacks on humans are rare (Apker, Updike, & trash or dumpsters by bears. McCarthy and Seavoy (1994) conclude Holdermann,2011; CMGWG, 2005), they have increased in recent that aversive conditioning for bears may be useful where single an‐ decades (Beier, 1991; Fithzugh, Kenyon, & Etling, 2003). There thropogenic food sources occur, but are questionable in urban areas have been 3 human fatalities from cougars and 16 nonfatal attacks where resources are widely distributed. in Colorado since 1990 (CPW 2018, Reported lion attacks on hu‐ There have been no studies confirming the effectiveness of aver‐ mans). Along with this has been a concomitant increase in other sive conditioning in cougars (CMGWG, 2005). Beier (1991) describes human–cougar interactions, which are likely due to habitat reduc‐ two unsuccessful attempts at aversive conditioning (one shot with tion, human encroachment, increased human recreational activi‐ rock salt, one treed and collared); however, one of these cougars ties, and possible increases in cougar populations (CMGWG, 2005). was already exhibiting aggressive behavior and the other was in Studies examining the intrinsic factors leading to cougar–human poor condition. McBride, Jansen, McBride, and Schulze (2005) used interactions have provided inconsistent results due to geography, hound capture and subsequent hound chases as a form of aversive methodology, and data quality. Cougar–human interaction may be conditioning on 4 Florida panthers with some degree of success. more likely in subadults (Aune, 1991; Mattson, 2007; Ruth, 1991) However, in Central Mexico, visual and sound deterrents were ef‐ or males (Tiechman, Cristescu, & Nielsen, 2013), especially when in‐ fectively used to scare felid predators away from livestock (Zarco‐ volving livestock depredations (Aune, 1991; Shaw, 1977; Suminski, Gonzalez & Monroy‐Vilchis, 2014). Aversive conditioning may be 1982; Torres, Mansfield, Foley, Lupo, & Brinkhaus, 1996). Analysis particularly difficult with cougars, as they are preying on naturally of conflict‐related mortalities in telemetered cougars has suspected occurring prey species in urban areas (deer, raccoons etc.), as well as higher likelihoods of conflict in male cougars (Thompson, Jenks, & on pets and livestock. In many cases, the undesirable behavior is not Fecske, 2014) or in subadult dispersing males and the oldest cougars associated with the prey species, but rather the location where they (Stoner, 2011). However, no significant age association was found are hunting and making kills. Therefore, associating a negative stim‐ when examining a large sample of necropsied cougar involved in pet ulus with an undesirable behavior (location) may be particularly dif‐ and livestock depredations (Torres et al., 1996). Other analyses have ficult or impossible for cougars compared with aversive conditioning suggested female cougars having a higher propensity to attack hu‐ on other species where there is a single source to condition against. mans (Mattson, 2007). The CMGWG (2005) concluded that a com‐ Large carnivores in the wildland–urban interface worldwide are bination of inexperience and unfamiliarity with their environment, often at risk because their requirements may conflict with those of as well as hunger, may cause young cougars to have more negative humans causing an urgent need for techniques to resolve and under‐ interactions with humans. stand conflicts between people and predators (Woodroffe, 2000). Wildlife managers throughout the western states, including To grow the knowledge base of how to potentially manage these those with Colorado Parks and Wildlife (CPW), are faced with deci‐ conflicts, we examined cougar spatio‐temporal landscape utilization sions about how to manage cougar populations and individual cou‐ patterns, prey utilization, and implementation of a potential conflict gars in order to maintain viable populations while also maintaining management techniques in an urban–wildland interface system in �10418 | the Colorado Front Range. Specifically, we model the habitat selec‐ ALLDREDGE et al. western edge approaching the continental divide. Housing density tion of a representative sample of GPS‐telemetered cougars and across the study area included wildland/rural (0–0.068 houses/ha, compare this to where sightings, conflicts, and harvest occurred as 69.7%), exurban (0.068–1.47 houses/ha), suburban (1.47–10 houses/ reported by agency personnel. We put the cougar landscape selec‐ ha, 2%), and urban (>10 houses/ha, 3%) (Blecha, 2015; Theobald, tion and conflict event patterns into context by summarizing the true 2005), creating a patchwork of habitats across the study, with the usage patterns and quantity of domestic prey species killed by the majority of urban areas along the eastern edge. Cities and counties sample of telemetered cougars on the same landscape. Along with throughout the area have also purchased and maintain large parcels these population‐wide data, we present case studies of individual of land as open space managed for human recreation and wildlife cougars with regard to conflict to address habitual versus opportu‐ populations. Naturally occurring prey species within this study area nistic behavioral responses. Finally, we address the feasibility and include elk (Cervus canadensis), mule deer (Odocoileus hemionus), cot‐ effectiveness of aversive conditioning techniques on cougars and tontail rabbit (Sylvilagus nuttallii), raccoon (Procyon lotor), and striped how this applies to future management of cougars within the urban– skunk (Mephitis mephitis). For detailed study area descriptions, see wildland interface. These results are likely applicable to many felid Moss et al. (2016) and Blecha et al. (2018). The majority of livestock species inhabiting such areas and may be broadly applicable to car‐ in the area was hobby livestock but will be referred generally as live‐ nivore species in urban settings. stock throughout. 2 | S T U DY A R E A This study was part of a long‐term cougar study (2007–2015) con‐ ducted in Colorado's northern Front Range including Boulder, 3 | M E TH O DS 3.1 | Cougar capture and incident data Cougars were captured from 2007 to 2015 (2014 and 15 to remove Jefferson, Gilpin, Clear Creek, and Larimer counties (Figure 1). This collars) using hounds, cage traps, foothold snares, and free‐darting, is a foothill–montane system covering an elevation gradient from and immobilized using medetomidine and ketamine hydrochloride. 1,590 m along the urban eastern edge to 3,170 m along the wildland Female cougars > 12 months old and male cougars > 24 months old F I G U R E 1 Study area used to investigate human–cougar interactions along the Front Range of Colorado �| ALLDREDGE et al. and >55 kg were collared with satellite GPS collars (Vectronics, GPS 10419 locations throughout the study area or home range where the ani‐ Plus Globalstar). Ear tag transmitters (ATS VHF, 28 g) or ARGOS mal may or may not have been observed. In the literature, this par‐ Satellite, 28 g) was used for cougars too small to be collared. Age was ticular application of the use‐availability framework has sometimes estimated using gum‐line recession or date of birth, animals were been referred to as using latent selection difference functions (LSD; weighed, morphometric measurements were taken, and blood and e.g., Erickson et al., 2014; Latham et al., 2011; Lendrum et al., 2018; tissue samples were collected for all individuals captured. All capture Roever et al., 2014). Typically, they are referred to as LSDs when and handling was done under approved capture and handling guide‐ the availability locations (represented by a 0 response in a logistic lines (ACUC 01‐2007 and 16‐2008). For the duration of the study, regression) represent something that can also be viewed as a used GPS collars were set to collect 7 to 8 locations per day (maintaining location (e.g., locations used by another species or during an alterna‐ a schedule at night of every 3 hr and reducing locations during the tive behavioral state), as opposed to a random selection of locations day when only collecting 7 per day). See Blecha and Alldredge (2015) where the animal was not observed. Given this sampling scheme, we for detailed description of capture, handling, and GPS collar settings. were able to assess the difference between where cougar incidents Cougar incident data between 2001 and 2014 within the north‐ did and did not occur, conditioned on cougar presence. ern Front Range of Colorado were collected and summarized by A habitat selection model was necessary to visualize the relative event type and location. When CPW is contacted about cougar risk of a cougar incidents occurring on the landscape. Because the sightings or conflicts, CPW personnel fill out and keep records on cougar incidents analysis relied on selecting random available lo‐ these events (known as “incident forms”). Data on these events, cations from used cougar locations, the cougar incident regression along with CPW's cougar harvest data, were summarized and used equation actually represents the relative risk of a cougar incident in analyses to determine site characteristics of where these events conditioned on cougar selection. Spatially interpolating this func‐ occurred. There were 629 conflict and sighting incidents reported tion assumes that cougar preference is uniform across the land‐ from 2001 to 2014. We will refer to a direct human encounter with scape. To account for this, we needed to multiply the relative risk of a cougar or property damage by a cougar as a conflict and a report a cougar incident (conditioned on cougar selection) by relative cou‐ that a cougar was seen in the area as a sighting. Collectively, we will gar habitat selection. We therefore fit a resource selection function refer to these as “cougar incidents.” (RSF) assuming an exponential form and estimated the coefficients using a Bayesian hierarchical logistic regression (Johnson, Nielsen, 3.2 | Timing of use and incident locations Merrill, McDonald, & Boyce, 2006). The used sample for the cou‐ gar habitat selection model were the 5,000 randomly selected GPS Understanding the timing of when cougars use areas with different points used as the available sample in the cougar incidents model, housing densities is important to understanding cougar behavior and and 25,000 available points were sampled uniformly across the potential management strategies. We examined four housing den‐ study area. Informal sensitivity tests were performed to determine sity classes aggregated at a 300 m scale; 0 houses per ha (wildland), that the number of used and available locations were appropriate. 0 to 1.47 (rural and exurban), 1.47 to 10 (suburban), and greater than Housing density level was used as a factor predictor, but suburban 10 houses per ha (urban). Hour of the GPS collar locations was dis‐ and urban were combined due to the small number of urban grid cretized into four bins: night (22:00–05:00), morning (05:00–09:00), cells. Continuous predictor variables included canopy presence, day (09:00–17:00), and evening (17:00–22:00) time periods. For each distance to canopy, heat loading (CHILI, Theobald, Harrison‐Atlas, individual and time period, we summed the number of GPS collar lo‐ Monahan, & Albano, 2015), elevation, distance to roads (Colorado cation observations in each housing density class. To standardize the Department of Transportation), distance to housing (Blecha, 2015), proportional use of the housing densities across time (because the and topographic wetness (Theobald, 2007), all of which were stan‐ time periods vary in duration), we then divided the resulting value by dardized to the mean and standard deviation across the study area. the total number of locations (for that individual) that fell into each We also used an additional interaction term between housing den‐ time bin class. sity and distance to housing. Due to high correlation among the To model the risk of cougar incidents, we employed a use‐avail‐ covariates with the interaction term, we orthogonalized (a mathe‐ ability framework fit using a logistic regression (Manly, McDonald, matical process that decorrelates sets of values) the covariates to fit Thomas, McDonald, & Erickson, 2007). In the use‐availablility frame‐ the model and then back‐transformed the coefficients to be compa‐ work, covariates from locations where cougar incidents occurred rable to the results from the cougar incident analyses. We present (used) are contrasted with random locations selected from an area the relative selection strength across the study area and the poste‐ considered available for cougar incidents (available). In this applica‐ rior mean and 95% credible intervals for the regression coefficients tion, available locations were restricted to GPS locations of collared (e.g., the log relative selection strength of a given covariate, such cougars using a latent selection difference function given that avail‐ that negative values indicate avoidance and positive values indicate ability locations can also be viewed as a used location during an al‐ preference; Avgar, Lele, Keim, & Boyce, 2017; Lele, Merrill, Keim, ternative behavioral state (e.g., Erickson, Found‐Jackson, & Boyce, & Boyce, 2013). Our primary reason for fitting a habitat selection 2014; Latham, Latham, & Boyce, 2011; Lendrum et al., 2018; Roever, model was to help visualize the conditional relationship between Beyer, Chase, & Aarde, 2014) as opposed to a random selection of cougar incidents and cougar use; therefore, we did not fit age or �10420 | ALLDREDGE et al. sex‐specific habitat selection models (for most conflicts, sex and project personnel and classified by feeding event presence, prey age of the cougar were unknown). species, and whether the species was domestic, for 56 cougars. In the cougar incident models, the conflict and sighting locations Sampling strata were based on each unique cougar and month mon‐ were considered the used sample, and a randomly selected subset of itored. See Blecha and Alldredge (2015) for detailed sampling and 5,000 cougar locations, regardless of the individual, was the avail‐ field methods. At the population level, we calculated an annual per‐ able sample. Because the available sample in this analysis is related capita average proportion and kill rate of domestic prey in the diet to what we define as the used sample in the habitat selection model by accounting for the unequal sampling probability of clusters across described in the above paragraph, the computational load of using monthly strata. Given that the number of kills cougar make, and all 233,348 locations as the used sample (necessitating a minimum therefore the corresponding number of clusters available to be sam‐ of 1,166,740 available locations given the conventional guidance on pled, is variable throughout the year (Knopff, Knopff, & Boyce, 2010, habitat selection analysis) was computationally infeasible. We used CPW unpublished data), we accounted for this unequal sampling the same model specification for the cougar incident models as for probability across our monthly strata (R package: SURVEY). Sample habitat selection, but without the interaction between housing den‐ coverage probability by month was calculated using the expected sity and distance to housing. Without the interaction term, orthogo‐ per‐capita feeding rate (f) multiplied by the number of unique com‐ nalization of covariates was not necessary. binations (k) of cougar and year monitored in each month, divided by For all models, we estimated coefficients using a Bayesian hier‐ the total kills sampled (n) per month. Confidence interval was cal‐ archical logistic regression, which was fit in R (R Core Team, 2017) culated based on techniques for small expected proportions (Korn using a Gibbs sampler with adaptive tuning. Adaptive tuning oc‐ & Graubard, 1998). On an individual level, the total feeding events curred during the first 20,000 iterations; the final tuning coefficient investigated, proportion of feeding events determined to be domes‐ was then used for a subsequent 20,000 iterations, with the first tic, and the measure of effort expended (the number of months an 2,000 iterations being discarded. Throughout we will use the ter‐ individual cougar's clusters were investigated) were summarized for minology relative selection strength or relative risk when discussing each collared cougar. coefficient estimates, as noted by Lele et al. (2013) and Avgar et al. Finally, a generalized additive model (binomial error family) was (2017). We present both the estimated coefficients (e.g., the log rel‐ used to test for variables associated with feeding on domestic prey ative risk of a given covariate) and a modified spatial description of over wild prey, given a cougar feeding event (domestic prey coded as the relative risk of a cougar incident on the landscape. To present the 1 and wild prey coded as 0). Candidate models included all combina‐ latter, we used the inverse logit of the estimated cougar incident re‐ tion subsets of Julian calendar day number (1–365 days: cubic cyclic gression equations, excluding the intercept, to visually describe the spline term), cougar age, cougar sex (male as baseline), and housing relative risk of a cougar incident and constrain the values between density (400 m buffer). Cougar age (by month) was determined dy‐ zero and one, and then multiplied each surface by a surface repre‐ namically, as some individuals were monitored over several years. senting relative cougar habitat selection. Nonlinear or interaction responses were considered in housing den‐ sity, cougar age, and cougar sex with simple smooth splines or sim‐ 3.3 | Sampled collared cougar case histories and domestic prey In order to understand cougar incident behavior, we summarized the number of collared cougars that engaged in conflict behavior ple quadratic terms. Models were evaluated via log likelihood, AICcc, and AICc weight. 3.4 | Aversive conditioning during the study and the types of behaviors by individual. Sample We also assessed the efficacy of aversive conditioning on cougars size is small and the types of conflict are varied, so detailed analy‐ in urban areas by describing cougar responses to several different ses were not warranted. We also believe that individual variation types of aversive conditioning. Aversive conditioning treatment was among cougars is important and would be masked by generalities applied to an individual GPS collared cougar in an attempt to alter its about habituation or the lack of it. Therefore, we present all case behavior in the future. The primary behavior that we focused on was histories on a time line showing the type of conflict and when it use of undesirable locations, such as urban neighborhoods and areas occurred for each individual in an attempt to demonstrate patterns near schools. Research from this study has already demonstrated of conflict among individuals. This summary includes only reported that cougars are generally using these areas to acquire food (Blecha conflicts and does not include domestic kills documented during et al., 2018), so aversive conditioning was typically applied to situa‐ feeding site investigations (see below) that were not reported by tions where a cougar had made a kill in an urban area. A similar situ‐ owners as conflict. ation would be a cougar that had sought cover (thick brush or under Cougar feeding patterns on domestic prey were derived and a porch) in urban areas during the day. A secondary situation for summarized based on the field investigation of GPS location clusters aversive conditioning is a cougar that had killed a domestic animal (including single point clusters representing small prey) produced (livestock or pet). In Colorado, a livestock owner can request that a by GPS collared cougars (Anderson & Lindzey, 2006). A stratified cougar be killed if it has killed livestock, so aversive conditioning in random sample of potential feeding sites were ground‐truthed by these situations was uncommon. �| ALLDREDGE et al. Aversive conditioning was not done in more remote areas or lower housing density exurban areas where cougars were preying on naturally occurring prey items, even if this occurred near hous‐ ing or if the cougar was just seen and reported. These situations were viewed as natural behaviors for cougars and would not nor‐ 10421 TA B L E 1 Timing of use within habitat density classifications relative to daily activity periods: night (22:00–05:00 hr), morning (05:00–09:00 hr), day (09:00–17:00 hr), and evening (17:00–22:00 hr) mally elicit a response from wildlife managers. All attempts of aver‐ Housing density (houses/ha) Night Morning Day Evening sive conditioning were done for situations and reports that would 0 0.22 0.27 0.28 0.24 normally result in wildlife managers hazing, trapping, or removing 0–1.47 0.40 0.17 0.14 0.29 a cougar. All cougars that were aversively conditioned were either 1.47–10 0.62 0.06 0.05 0.27 previously collared or were collared as part of the treatment so that >10 0.83 0.05 0.02 0.11 responses could be assessed. Initially aversive conditioning treatments involved shooting the offending individual with 2 to 3 bean bag rounds fired from a 12‐ Note: Proportion use was standardized to account for differing GPS location schedules within time periods. gauge shotgun at a distance > 10 m up to 30 m to avoid injury to the animal. Releases were always set so that the treatment could be con‐ selection strength declined as a function of distance to housing; sistently done by trained personnel. In many situations, the cougar however, this effect describes the baseline cougar response to hous‐ was in an undesirable location, so capture and relocation were also ing (in rural areas; β = −1.33; Figure 2). In exurban areas, there was an necessary. In these cases, the cougar was free‐range darted or cap‐ overall positive effect of distance to housing on selection (β = 14.63, tured in a cage trap, relocated to a nearby open space 4–20 km away, for an overall effect of 13.33); however, in suburban and urban areas, and shot with bean bag rounds on release. Relocation distance was cougars demonstrated strong selection with decreasing distance intentionally kept small in order to assess the effects of the aversive to housing (β = −52.51, for an overall effect of −53.84; Figure 2). conditioning treatment on behavior rather than the effect of reloca‐ Cougars showed a preference for areas further from roads (β = 1.64; tion. In the latter part of the study (2011–2014), based on responses Figure 2). The presence of canopy cover increased cougar preference to aversive conditioning, we assessed the response of cougars to (β = 0.15); however, cougars demonstrated avoidance of areas with removing their kills from undesirable locations. In these cases, GPS increasing distance to canopy (β = −1.90), heat loading (β = −0.15), data were used to find kills in these areas, and then, kills were re‐ elevation (β = −1.29), and topographic wetness (β = −0.46; Figure 2). moved as soon as they were detected. Conditional on where conflict is more likely to occur based on The types of human–cougar interactions were varied, most situ‐ cougar selection models, the presence of canopy cover (β = −0.30) ations were unique, and sample sizes were small so we summarized and increasing distance to roads (β = −1.45) decreased the relative these interactions into general categories including undesirable loca‐ risk of a reported conflict event (Figure 3). Increasing distance to tion (areas of high housing density, inside city limits, near school), un‐ canopy (β = 1.36), elevation (β = 0.84), and topographic wetness desirable kill location, and domestic predation. Aversive treatments (β = 0.45) increased the relative risk of a reported conflict event were also unique to each situation; therefore, we present these data (Figure 3). Being in an exurban (β = 1.67) or suburban/urban (β = 2.76) and cougar responses as summaries because the variability in each area also increased the relative risk of a reported conflict event, with situation did not warrant statistical analysis. the relative risk being higher in suburban/urban areas (Figure 3). The relative risk of a sighting event is also implicitly conditioned 4 | R E S U LT S 4.1 | Timing of use and incident locations on where human activities occur; however, this component was unobserved and unmodeled, and therefore, care should be taken when interpreting the effects of covariates on relative risk of a cou‐ gar sighting. Increasing elevation (β = 0.46) and topographic wet‐ We examined the timing of when cougars used various housing den‐ ness (β = 0.24) had a positive effect on the relative risk of a cougar sities and where incidents occurred relative to how cougars used the sighting, given cougar selection (Figure 4). Similar to conflict events, landscape for 76 (43 females, 33 males) independent age cougars being in an exurban (β = 1.18) or suburban/urban area (β = 2.60) also with GPS collars. Of these cougars, 6 females (14.0%) and 11 males increased the relative risk of a sighting event. Increasing distance (33.3%) never used areas with housing densities over 1.47 houses/ to road (β = −4.61) and increasing distance to housing (−6.38) de‐ ha (suburban/urban) and 32 females (74.4%) and 28 males (84.8%) creased the relative risk of a sighting event (Figure 4). never used areas with more than 10 houses/ha (urban). Cougar use Spatially, the relative risk of a sighting event was similar to the of higher housing density areas was predominantly at night (Table 1). relative risk of a conflict event, however less heterogeneous across Eighty‐three percent of cougar use in areas exceeding 10 houses/ha the landscape, which is likely due to the nonsignificant effects of and 62% of cougar use in areas between 1.47 and 10 houses/ha oc‐ canopy, distance to canopy, heat loading, and the small positive ef‐ curred between 22:00 and 05:00 hr. Cougars selected exurban over fects of elevation and topographic wetness (Figure 4). Cougar pre‐ rural habitat (β = 9.45) and strongly avoided suburban/urban habi‐ dation on domestic animals generally occurred in areas that cougars tat relative to rural habitat (β = −34.29; Figure 2). Cougar's relative were selecting for and within the higher housing density areas along �10422 | ALLDREDGE et al. F I G U R E 2 Relative probability of cougar use within the study area, given the covariates and their effect sizes, and the log relative selection strength of a given covariate, such that negative values indicate avoidance and positive values indicate preference. Black circles on the map indicate cougar harvest locations, while gray symbols indicate reported cougar predation on domestic animals and livestock. Suburban/Urban Housing Dist. Int. and Exurban Housing Dist. Int. represent the additional effect of distance to housing when an individual was in a suburban/urban area or exurban area, respectively (e.g., the interaction term between housing density and distance to housing) F I G U R E 3 Relative risk of a reported cougar conflict event given the relative probability of cougar use within the study area, and the log relative risk strength of a given covariate. Negative values indicate decreased risk, and positive values indicate increased risk. Black circles on the map indicate cougar harvest locations the eastern edge of the study area (Figure 2). Some predation on collared because of conflict behavior (3 livestock predation and 2 livestock and dogs did occur in lower housing density areas in the because of location) (Figure 5). Of the females, 9 were initially cap‐ western and central portions of the study area. Although some cou‐ tured and collared because of conflict behavior (3 livestock preda‐ gar harvest occurred in the higher housing density areas along the tion, 1 domestic pet predation, and 5 because of location). A total of eastern edge of the study area, the majority was in lower housing 24 conflict events were reported for male cougars: 14 for livestock density areas in the western and central portions (Figure 2). In gen‐ predation, 2 for predation on dogs, and 8 for using an undesirable eral, cougar harvest did not occur in areas where cougars tended to location. AM13 was the only male that showed repeated livestock be seen or in conflict with humans (Figure 3). predation over a short time period, killing a llama (Lama glama) and 2 small horses over a 5‐month period but then going for several years 4.2 | Cougar case histories and domestic prey before killing livestock again. AM14 never killed livestock until he was 6 years old and then was euthanized for killing multiple llamas in Of 52 males and 50 females that were captured and monitored dur‐ one area over a month. A total of 26 conflict events were reported ing the study, 11 males and 13 females were reported by the pub‐ for female cougars: 7 for livestock predation, 2 for predation on lic for conflict behavior. Of the males, 5 were initially captured and dogs, and 17 for using an undesirable location. �| ALLDREDGE et al. 10423 F I G U R E 4 Relative risk of a cougar sighting event, given the relative probability of cougar use, within the study area, and the log relative risk strength of a given covariate, such that negative values indicate decreased risk and positive values indicate increased risk. Black circles on the map indicate cougar harvest locations F I G U R E 5 Timeline for monitored cougars involved in at least one human conflict event in the Front Range of Colorado from 2007 to 2016. Letters indicate a capture event (C), a mortality event (H: hunting, N: natural, E: euthanized), or collar removal (R). Symbols indicate a conflict event, which may have overlapped with a capture or mortality event. Values at the start of each row indicate age at first capture Domestic prey did not appear in kill sites for 34 of 56 cougars sampled in kill site investigations. Of all of the GPS clusters that were of domestic prey found at cougar feeding sites ranged from 0.01 in January to 0.10 in May (Table 3). investigated, 1,625 were verified as kill sites, and of those, 68 con‐ Conditional on a feeding event, domestic prey item pres‐ tained domestic prey items: 43 cat, 17 dog, 2 alpaca, 2 goat, 1 sheep, ence was most parsimoniously described as a function of housing 1 llama, 1 fowl, and 1 unidentifiable pet (based on the presence of a density with a quadratic (β HD = 1.019 (SE = 0.246), β HD2 = −0.128 collar). Domestic species constituted 3.87% (95% C.I.: 2.87%–5.0%) (SE = 0.042)), cougar age (βage = −0.762 (SE = 0.179)), cougar sex of items preyed upon annually. Interindividual variation (by cougar) (βmale = 0.479 (SE = 0.138)), and an interaction between sex and age of domestic predation was present based on a nonuniform distribu‐ (βmale,age = 0.387 (SE = 0.134)), holding 77.6% of the AICc weight. tion in domestic prey proportion when considering each cougar as a Julian calendar day appeared in the second‐ranked model holding single sample. Domestic prey proportion by individual ranged from 11.1% AICc weight. The influence of housing density on domestic 0 to 0.75 (Avg. 0.16, Table 2). However, the high proportion (0.75) of prey killing probability was strongest at ~5 houses per ha (Figure 6). domestic prey was for a single individual that was only monitored This influence of housing density appears to diminish with >5 houses for two months in the study (4 total kill sites documented). The in‐ per ha, but this declining response may be an artifact of the small dividual with the next highest proportion of 0.357 was monitored sample size of feeding events in these highest housing densities for 15 months with 28 total kill sites documented. The proportion (1.7% of all 1625 feeding sites). Cougar age and sex were interacting �10424 | Cougar ID ALLDREDGE et al. Months monitored Events investigated Proportion domestic AF58 2 4 0.75 AF56 15 28 0.36 AF37 3 3 0.33 AM46 3 4 0.25 AM70 10 16 0.25 AM74 16 34 0.21 AF24 3 5 0.20 AM606 7 17 0.18 AM21 2 6 0.17 AF32 9 20 0.15 AF88 4 7 0.14 AF40 36 73 0.10 AM44 30 52 0.08 AF73 21 43 0.07 AF52 30 68 0.06 AF57 25 54 0.06 AF86 9 19 0.05 AM14 29 42 0.05 AF69 24 64 0.05 AM84 13 25 0.04 AF50 16 37 0.03 AF59 22 40 0.03 TA B L E 2 Proportion of domestic prey found at cougar kill sites in relation to the number of kill sites investigated and the number of months kill sites were investigated, reported by cougar Note: Kill sites were investigated for 56 cougars but domestic prey was found at kill sites for only these 22 individuals. TA B L E 3 Observed monthly proportion of domestic prey found at kill sites (pd), per‐capita feeding rate based on all prey items (fm), number of cougars monitored each month across years (km), and the number of ground‐truthed clusters confirmed as kill sites for all prey items (nm) pd fm 4.3 | Aversive conditioning From 2007 to 2011, 7 female and 6 male cougars were aversively conditioned from 1 to 3 times each for a total of 17 and 8 aversive conditioning treatments on females and males, respectively. Aversive km nm conditioning treatments for being in undesirable location were con‐ January 0.01 6.09 70 152 ducted 14 and 3 times for females and males, respectively, and all of February 0.06 5.76 62 115 these treatments included shooting the offending individual with 1 to March 0.04 5.68 65 139 2 beanbag rounds. A naturally occurring food source was also involved April 0.05 6.72 68 149 May 0.10 6.39 64 134 June 0.04 11.31 66 138 July 0.03 12.51 63 173 August 0.05 9.91 65 144 September 0.01 10.78 61 127 October 0.06 10.43 58 111 tial conflict (neighborhoods within city limits) after treatments. Only 1 November 0.02 7.92 64 124 male continued to use these areas after treatment; however, 2 males December 0.03 8.01 58 119 were later aversively conditioned for killing livestock. in 16 of these cases. Because of the location, 15 aversive treatments for undesirable location also included relocation to nearby open space. Relocation distances between 4 and 9 km (n = 13) resulted in the cou‐ gars return within 1 to 2 days post‐treatment. Relocation distances of 16 and 19 km for the remaining 2 cougars resulted in a return to the individual's home range within 6 days, but no immediate return to the location of conflict. All females continued to use locations of poten‐ The remaining 8 aversive conditioning treatments involved cou‐ variables revealing that age of the individual did not affect the like‐ gars killing livestock (n = 5) or domestic dogs (n = 3). In four cases, lihood of domestic prey at a kill for males, but subadult female cou‐ aversive conditioning was done on site and did not involve capture or gars were more likely than all other age–sex types to prey upon relocation, while the remaining 4 were relocated. All relocated cou‐ domestic animals (Figure 7). gars returned to the capture location within 2 days, but immediately �| ALLDREDGE et al. F I G U R E 6 Model predicted domestic prey probability and 95% confidence intervals, given a feeding event, as a response to housing density for male and female cougars 10425 F I G U R E 7 Model predicted domestic prey probability, given a feeding event, as a response to cougar age and sex as interacting variables. Housing density was held at 1.47 houses per ha, which is the transition point from exurban to suburban housing density left the area, presumably due to prey carcass removal. Two cougars never killed livestock again post‐treatment and 2 were euthanized for the data on such individuals are limited to age and sex. This study is killing livestock within 6 months following treatment. The remaining unique in that it was designed to examine human–cougar interac‐ cougars opportunistically killed livestock for the rest of their lives tion, so cougars were not lethally removed following conflict, and (4 to 6 additional years each making 1 to 2 total livestock kills) and instead allowed cougars to be monitored for behavioral patterns. were eventually euthanized for livestock depredation (e.g., AM13, Figure 5). Our findings on cougar space use within the wildland–urban in‐ terface are similar to those of Kertson et al. (2013), Maletzke et al. During 2011–2014, prey remains were removed from cougar (2017), Stoner (2011), and Wang et al. (2017), which are the only caches (n = 12) in undesirable locations and no aversive conditioning other studies that examined cougar populations that interacted was done. In all cases, cougars returned to investigate the area. In 2 within urban areas but also had access to large wildland habitats. cases, the cougar did not leave the area and killed another small prey Common assumptions are that increasing cougar complaints are as‐ item (raccoons) in the vicinity of the original kill. In the remaining sociated with increasing subadult and younger age classes (Lambert 10 cases, the cougar left the area and made its next kill away from et al., 2006; Robinson, Wielgus, Cooley, & Cooley, 2008) and that human‐developed areas (in open space) but did continue to use de‐ use of exurban habitats is limited to subadults and transients. The re‐ veloped areas. sults presented here refute these assumptions, demonstrating that all age classes, especially females, utilize these areas. Although all age classes are using these human‐dominated areas, it is females and 5 | D I S CU S S I O N younger age classes that are more likely to be involved with domes‐ tic animal predation conflicts. Kertson et al. (2013) showed a similar We presented data from a long‐term study of cougars utilizing the pattern of use among all demographic classes utilizing residential urban Front Range of Colorado describing how cougars use urban areas. Stoner (2011) suggested that it was the maternal females and areas, the timing of use, predation on domestic animals, and the inefficient hunters (i.e., very young dispersing animals or senescent potential for aversive conditioning to change cougar behaviors. females) that exploited these riskier urban–wildland habitats to cap‐ Although reports of cougars using urban areas are becoming more italize on food resources. common, few studies have examined these dynamics. Our results are Areas with high densities of humans are often thought to be species‐specific, but are likely broadly applicable to large obligate low quality habitat for cougars because of increased cougar mor‐ carnivore species that utilize the wildland–urban interface and are tality due to roadkill, lethal removal following depredation involving bound to interact with humans. Little is known about cougar conflict pets or livestock, and policies favoring cougar removal to maintain and depredation on domestic prey because of the difficulty in col‐ human safety (CMGWG, 2005). While these sources of mortality lecting data on repeat depredation. Conflict and depredation histori‐ occurred during our study, our data suggest that cougar removals cally have resulted in lethal removal of the offending individual, so related to depredation and human safety were uncommon relative �10426 | to the amount of time cougars used these areas and the number of ALLDREDGE et al. selection (i.e., if we compare incident locations to where cougars pets taken by cougars, primarily because cougars went undetected. are), the factors driving these occurrences are different from areas Cougar population density in this area was estimated at 4.1 indepen‐ that cougars are selecting for. Cougar kills of domestic animals show dent cougars per 100 km2, one of the highest reported cougar den‐ a similar pattern, especially for domestic cats (Figure 2), which are sities in the literature (Alldredge, Blecha, & Lewis, 2019), indicating generally closely associated with housing. Some livestock and large that cougars are likely doing very well in these urban habitats. Similar dog conflicts occurred in areas away from housing that are more typ‐ patterns of high carnivore density and human conflict in urban areas ically selected by cougars, likely because of overlap with preferred have been documented for leopard (Panthera pardus) and striped habitats between cougars and free‐ranging livestock. However, this hyena (Hyaena hayena) (Athreya, Odden, Linnell, Krishnaswamy, & does suggest that animal husbandry practices, especially for dogs, Karanth, 2013), suggesting the adaptability of carnivores may gen‐ could help reduce conflict (such as not letting dogs roam freely, es‐ erally allow them to exploit urban environments despite human con‐ pecially when cougars are active, and covering the top of kennels). flict and increased risk of mortality. Given the amount of private property that occurred within our Our data suggest that the majority (79%) of cougars within the study area, cougar harvest was limited. Examining harvest locations wildland–urban interface are avoiding areas with higher housing (Figures 2 and 3) shows that most harvest occurred in areas where density. Of the cougars that use higher housing density areas, they sightings and conflicts were less common. This may indicate that generally use these areas at night and leave before morning. Based cougar hunting as currently practiced might have limited success as on this, it would appear that cougars view these areas as risky en‐ a management strategy to reduce human–cougar interactions, likely vironments and are avoiding them during periods of higher human due to large amounts of private property limiting common methods activity. Blecha et al. (2018) found that cougars using urban areas of harvest. However, cougars have large home ranges so harvest were doing so for food acquisition. This suggests that the poten‐ that occurs outside the wildland–urban interface may include ani‐ tial food resource within higher housing density areas is worth the mals that also utilize urban areas. Robinson et al. (2008) point out risk, which may be especially true for females with kittens as their that cougar harvest to reduce population size within small areas is energetic demands increase. Females with kittens near urban areas generally ineffective because of high immigration rates as there was tended to den kittens outside of these areas and make nightly for‐ a temporal difference between cougar use and human activity. ays into neighborhoods in search of prey. Cougars in urban Western Being in an undesirable location was the primary type of conflict Washington showed similar landscape use patterns of exploiting po‐ for female cougars, especially older individuals. All but 2 conflicts tential prey resources within urban settings while minimizing the po‐ associated with undesirable location were reported from a cougar tential for interactions with people (Kertson, Spencer & Grue, 2011; kill of a naturally occurring prey item, demonstrating that the major‐ Kertson, Spencer, Marzluff, et al., 2011). Shifting activity patterns to ity of these conflicts involved cougars using higher housing density nocturnal periods in exurban areas have also been reported for other areas to acquire prey. Moss et al. (2016) in our study area as well as large carnivores, such as black bears exploiting urban food resources Kertson, Spencer, and Grue (2011) and Robins, Kertson, Faulkner, (Lyons, 2005). and Wirsing (2019) all documented a significant use of alternative Cougar resource selection showed consistent patterns of habitat small‐bodied prey in urban areas compared with studies in wildland use, with the exception of avoidance of the lowest density hous‐ areas that documented ungulates as the primary prey for cougars. ing (wildland/rural) relative to exurban habitat. Because wildland Undesirable location conflicts occurred across all age classes, but and rural habitat are combined, it may be that these areas provide may appear slightly higher for younger cougars (Figure 4), likely rep‐ fewer prey resources, especially during the winter, compared with resenting transient individuals. The potential undesirable location exurban habitat. It is possible that this may be an artifact of lower conflict was probably higher for older age females because they reg‐ sampling effort in the western portion of the study area as capture ularly used higher housing density areas (based on GPS locations), efforts focused more on the exurban eastern edge, but a concur‐ but older females were rarely seen or reported by the public. rent study estimating cougar density in this study area suggested Conflict associated with livestock depredation was more com‐ a similar distribution of cougars (Alldredge et al., 2019). Regardless, mon for male cougars and appeared to be opportunistic, although, the avoidance of housing was still predictable showing cougar avoid‐ on some occasions, an individual would kill multiple livestock over ance of humans when in more rural and open areas, and avoidance a short time period. Some cougars were euthanized following live‐ of suburban and urban areas. Space use patterns of cougars in this stock depredation, so it is difficult to fully assess whether cougars landscape demonstrate the highly adaptable behaviors found in were habituated to preying on livestock. Situations where collared many large carnivores as habitat generalists, including black bears cougars could not be recaptured following livestock depredation (Baruch‐Mordo et al., 2014; Lewis et al., 2015), coyotes (Gehrt et were our only opportunities to gather information on the repeat al., 2009; Poessel et al., 2016), and other felids (Burdett et al., 2010; behavior or frequency of livestock kills. Information from cougars Donovan et al., 2011). The effect of covariates on the relative risk of cougar incidents that killed livestock that could not be recaptured revealed an op‐ portunistic use of livestock based on long time intervals between was often the opposite of the effect of the covariate on cougar se‐ these events and did not support habituation to preying on livestock. lection. These results demonstrate that if we condition on cougar Cougars likely encountered livestock frequently but infrequently �| ALLDREDGE et al. 10427 preyed on livestock suggesting that cougars are selecting against as the individual returned to their cache, generally in high‐quality livestock, which can be seen with older age classes that infrequently cougar habitat. In these situations, the offending cougar had already killed livestock. Torres et al. (1996) also found male cougars preyed received a reward (food) for the behavior that we were attempting on livestock significantly more than females in California. to condition against. Other data presented here would also suggest Assessing depredation of pets (cats and dogs) from conflict that cougars likely utilize these undesirable locations regularly for reports would suggest that these events were rare. Data from acquiring resources or kill domestic pets and go undetected in both California from 1972 to 1995 also suggested pet depredation was situations. In ideal circumstances, aversive conditioning would occur relatively uncommon compared with predation on livestock based at the point in time the undesired behavior is initiated (immediately on permits issued in response to a complaint (Torres et al., 1996). as they enter the poor location or right as the attack on livestock is However, pet depredation by cougars based on kill site investiga‐ initiated). With current GPS technology and real‐time data, it is con‐ tions in our study suggests that this is more common than indicated ceivable that aversive conditioning could be applied as cougars enter from conflict reports. This is likely because the small body mass of these areas or approach livestock, but this would require a huge ef‐ pets allow prey to be easily cached or moved from the property by fort and provides no reasonable long‐term management applications cougars and thus rarely discovered by pet owners. For the Front as all cougars would need to have GPS collars. Range of Colorado, on average, we estimated that 4% of a cougar's Although not conclusive, our data suggest that cougars were annual prey (individual kills) was domestic species, primarily small not habituated to these undesired behaviors that result in conflict pets. In general, pet depredation appeared to be opportunistic, but events, but were using available resources opportunistically or for two cougars regularly killed pets. We documented, from sampled foraging opportunities as others were limited. Certainly some cou‐ GPS feeding sites, a 4‐year‐old male that killed 7 dogs over a 14‐ gars were removed after repeated livestock depredations, suggest‐ month interval and a 2‐year‐old female that killed 8 cats and 2 dogs ing habituation. However, cougars that could not be removed after over a 14‐month interval. These two instances represent minimums repeated livestock depredation showed a tendency to go back to because these numbers only represent kill sites that were sampled. naturally occurring prey and utilized livestock only opportunistically. Housing density was the best predictor variable associated Similarly, some cougars used higher housing density areas with some with cougars feeding on domestic animals given a sampled feed‐ regularity but this may have been driven by food resources. In this ing event which was expected because of the strong association area, cougars have been shown to have higher avoidance of human between domestic animals and houses. There was some evidence populated areas following feeding events and decreased avoidance that calendar day was a factor influencing cougar use of domestic as time increases since their last feeding event (Blecha et al., 2018). animals, as the proportion of domestic prey found in feeding sites Most of the higher housing density areas within our study (within increased slightly during May (Table 3). Cougars during this study cities) had stable and consistent naturally occurring prey including also increased use of higher housing density areas during May, co‐ deer, raccoons, and rabbits (Blecha, 2015). These factors only in‐ inciding with an increased use of smaller nonungulate prey (Blecha crease the difficulty of applying aversive conditioning techniques et al., 2018). Housing density is also a good proxy for the spatial within the wildland–urban interface. availability of small domestic prey, especially outdoor and feral Removal of cached kills from undesirable locations proved some‐ cats (Blecha, 2015), the primary domestic prey item found in this what effective as a means of getting a cougar to leave an urban area, study. Domestic prey items were more likely to be present at the especially when more prey was not immediately available in the area. feeding sites of female cougars, with lower use of domestic pets as Cougars tended to leave the area and hunt elsewhere except when females age. Overall, to reduce conflicts associated with cougars raccoons were immediately available. It is possible that cache re‐ killing pets, we recommend managing for an older age class cou‐ moval could have long‐term effects on cougars, as these areas would gar population. Other studies only provided cougar demographic be inefficient for cougars to acquire needed resources, although it relationships based on conflicts reported to agencies (Aune, 1991; is doubtful that managers could find and remove enough carcasses Tiechman et al., 2013; Torres et al., 1996) or the conflict‐related (i.e., those cached by unmarked cougars) to have the desired effect. mortality events of collared cougars (Stoner, 2011; Thompson et Removal of livestock kills could have similar effects and cause cou‐ al., 2014). Future studies should investigate real versus perceived gars to leave the area and search for prey elsewhere but only if re‐ domestic predation frequency using GPS collar sampling methods maining livestock are unavailable. of feeding sites. Although aversive conditioning appears to be relatively ineffec‐ Aversive conditioning within this study was generally ineffective tive, there may be hidden benefits that make the effort worthwhile. for altering cougar use of urban areas or other undesirable locations, In all of our efforts to aversively condition cougars, it is likely that which is likely a result of how treatments were applied because of we reinforced the idea to cougars that humans present a risk and logistical constraints and cougar behavior. In general, aversive treat‐ should be avoided. In many exurban areas of the Front Range, it is ments on cougars in undesirable locations were done on cougars re‐ becoming more and more common for people to report cougars lay‐ turning to their kill of a naturally occurring prey item they made the ing near roads or houses in broad daylight, seemingly unconcerned previous night in a populated area. Similarly, aversive conditioning of about humans passing by. In some cases, people can even stop to cougars following depredation of domestic animals was conducted take pictures of such cougars. This is in contrast to remote areas �10428 | ALLDREDGE et al. of Colorado where people rarely see cougars. This apparent bold people can take to help prevent or limit conflict and promote coex‐ behavior or habituation to human activity may not be desirable and istence in the future. First, it is important to educate the public on could lead to increased conflicts. Similar phenomena of increased cougar behavior and habits in these areas so that people understand boldness have been documented for other carnivore species in what cougars are doing and what they can do to reduce future con‐ urban settings, such as brown bears (Ursus arctos; Fernandez‐Gil, flict. Maintaining large (>2,000 ha) open space or natural areas with 2014) and coyotes (Baker & Timm, 1998; Timm, Baker, Bennett, ample prey and cover for cougars within the urban habitat matrix & Coolahan, 2004), including coyotes in Colorado (Breck et al., will provide areas for cougars with limited human activity. We also 2019). It has been postulated that historically intense human per‐ recommend educating the public about living with cougars, includ‐ secution of some species selected against bolder individuals and ing protecting livestock and pets from cougars and manipulating that there has been a recent release from this selective pressure urban habitats to limit attractions for prey species and limiting hid‐ in human‐dominated landscapes that reduce hunting opportu‐ ing cover. People are emotionally attached to hobby livestock and nities, thus allowing bolder, more aggressive individuals to thrive pets, and cougar interactions with these animals generally result in in these riskier urban environments (Martinez‐Abrain, Jimenez, & a negative outcome for the cougar even though it is engaging in a Oro, 2018). There is some question whether such differences in natural activity and in many situations is in quality cougar habitat. behavioral traits are a result of phenotypic plasticity or of intrin‐ Therefore, better animal husbandry will help limit cougar conflict sic differences (Miranda, Schielzeth, Sonntag, & Partecke, 2013). and negative attitudes toward coexisting with cougars (CMGWG, Regardless, carnivore species do appear to be bolder and more ag‐ 2005). As a final step, we would recommend habitat manipulations gressive in urban settings and these phenomena correlate well with within higher housing density areas or areas where cougar presence increasing cougar sightings and conflict that has been observed in is not desirable, including green belts within these areas. Habitat the Front Range of Colorado and other urban settings in the West. manipulations should be directed toward limiting hiding cover for Given this trend observed across carnivore species, it seems that cougars and, more importantly, limiting habitats that are directly aversive conditioning of cougars in higher housing density areas beneficial for prey species. Our data presented here and from Moss could provide indirect benefits in continuing to instill the fear of et al. (2016) and Blecha et al. (2018) strongly suggest that cougars humans in animals occupying such areas. It may even be beneficial are utilizing these areas to take advantage of alternate prey species, to actively haze cougars that are seen in these settings, that appear and limiting prey in these areas should limit the motivation of cou‐ to be overly comfortable around human activity, or that exhibit lit‐ gars to use these risky habitats. tle or no fear of humans. Cougar characteristics and circumstances surrounding conflict situations were highly variable and did not show consistent patterns. AC K N OW L E D G M E N T S In some situations, an individual would repeat conflict behaviors over This project was funded by Colorado Division of Wildlife Federal a short time period but then stop exhibiting these behaviors and uti‐ Aid in Wildlife Restoration Project W‐153‐R and Colorado Division lize the wildland–urban interface for long periods of time without of Wildlife game cash funds. We appreciate all of the field efforts conflict. In other situations, individual cougars were removed after of the technicians on this study. We also appreciate the numerous repeat conflict behaviors, resolving issues within certain areas. land owners, including county and city properties that allowed us Individual variation in cougar utilization of human‐dominated land‐ access. The efforts of those that reviewed this manuscript were scapes and behavior is not unique to this study (Aune, 1991; Kertson greatly appreciated. et al., 2013; Riley & Aune, 1997; Robins et al., 2019; Sweanor, Logan, Bauer, Millsap, & Boyce, 2008). Given this variation, it would appear that cougar conflict management should be focused on individual C O N FL I C T O F I N T E R E S T cougars for each unique set of circumstances. Our data also support The authors declare that they have no conflict of interest. the conclusions of Kertson et al. (2013) suggesting that human–cou‐ gar interactions are a function of individual behavior (learned and innate) and circumstances and that managing for and maintaining an older age structure of cougars in the wildland–urban setting would be beneficial. The coincidence of high cougar density, and extensive use of higher housing density areas and rapid human expansion along the Colorado Front Range, has created a situation where the potential AU T H O R C O N T R I B U T I O N S M.W.A. designed the study, collected field data, analyzed data, and wrote the manuscript. F.E.B. analyzed data and wrote the manu‐ script. K.A.B. collected field data, analyzed data, and wrote the man‐ uscript. All authors gave final approval for publication. for human–cougar interactions and conflicts is high. However, the realized level of this interaction is relatively low compared with the potential, suggesting that cougars can coexist with people reason‐ ably well. Kertson et al. (2013) came to the same conclusion in an urban setting in western Washington. However, there are steps that E T H I C A L A P P R OVA L This research complies with the laws of the country in which it was performed. Capture and handling of animals was approved by the �| ALLDREDGE et al. institutional animal care and use committee (CPW ACUC ACUC 01‐2007 and 16‐2008). DATA AVA I L A B I L I T Y S TAT E M E N T The raw data supporting this research are openly available from the Dryad data archive https​://doi.org/10.5061/dryad.bt3ng2j. ORCID Mathew W. Alldredge https://orcid.org/0000-0002-7133-9621 REFERENCES Alldredge, M. W., Blecha, T., & Lewis, J. (2019). Less invasive monitor‐ ing of cougars in Colorado's Front Range: An evaluation and review. Wildlife Society Bulletin, 43, 223-230. Anderson, C. R., & Lindzey, F. G. (2006). Estimating cougar predation rates from GPS location clusters. Journal of Wildlife Management, 67, 307–316. Anderson, C. R. J., Lindzey, F., Knopff, K. 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Animal Conservation, 17, 371–378. https​:// doi.org/10.1111/acv.12104​ How to cite this article: Alldredge MW, Buderman FE, Blecha KA. Human–Cougar interactions in the wildland–urban interface of Colorado's front range. Ecol Evol. 2019;9:10415– 10431. https​://doi.org/10.1002/ece3.5559 � Dublin Core The Dublin Core metadata element set is common to all Omeka records, including items, files, and collections. For more information see, http://dublincore.org/documents/dces/. Title A name given to the resource Journal Articles Description An account of the resource CPW peer-reviewed journal publications Text A resource consisting primarily of words for reading. Examples include books, letters, dissertations, poems, newspapers, articles, archives of mailing lists. Note that facsimiles or images of texts are still of the genre Text. Dublin Core The Dublin Core metadata element set is common to all Omeka records, including items, files, and collections. 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Title A name given to the resource Human-Cougar interactions in the wildland-urban interface of Colorado's front range Rights Information about rights held in and over the resource <a href="http://rightsstatements.org/vocab/InC-NC/1.0/" target="_blank" rel="noreferrer noopener">In Copyright - Non-Commercial Use Permitted</a> <a href="https://creativecommons.org/licenses/by/4.0/" target="_blank" rel="noreferrer noopener">Attribution 4.0 International (CC BY 4.0)</a> Date Created Date of creation of the resource. 2019-08-20 Subject The topic of the resource Aversive conditioning Colorado Conflict Cougar Domestic predation Human interaction Livestock, predation <em>Puma concolor</em> Residential development Wildland–urban interface Description An account of the resource <span>As human populations continue to expand across the world, the need to understand and manage wildlife populations within the wildland</span><b>–</b><span>urban interface is becoming commonplace. This is especially true for large carnivores as these species are not always tolerated by the public and can pose a risk to human safety. Unfortunately, information on wildlife species within the wildland</span><b>–</b><span>urban interface is sparse, and knowledge from wildland ecosystems does not always translate well to human-dominated systems. Across western North America, cougars (</span><i>Puma concolor</i><span>) are routinely utilizing wildland</span><b>–</b><span>urban habitats while human use of these areas for homes and recreation is increasing. From 2007 to 2015, we studied cougar resource selection, human–cougar interaction, and cougar conflict management within the wildland</span><b>–</b><span>urban landscape of the northern Front Range in Colorado, USA. Resource selection of cougars within this landscape was typical of cougars in more remote settings but cougar interactions with humans tended to occur in locations cougars typically selected against, especially those in proximity to human structures. Within higher housing density areas, 83% of cougar use occurred at night, suggesting cougars generally avoided human activity by partitioning time. Only 24% of monitored cougars were reported for some type of conflict behavior but 39% of cougars sampled during feeding site investigations of GPS collar data were found to consume domestic prey items. Aversive conditioning was difficult to implement and generally ineffective for altering cougar behaviors but was thought to potentially have long-term benefits of reinforcing fear of humans in cougars within human-dominated areas experiencing little cougar hunting pressure. Cougars are able to exploit wildland</span><b>–</b><span>urban landscapes effectively, and conflict is relatively uncommon compared with the proportion of cougar use. Individual characteristics and behaviors of cougars within these areas are highly varied; therefore, conflict management is unique to each situation and should target individual behaviors. The ability of individual cougars to learn to exploit these environments with minimal human–cougar interactions suggests that maintaining older age structures, especially females, and providing a matrix of habitats, including large connected open-space areas, would be beneficial to cougars and effectively reduce the potential for conflict.</span> Creator An entity primarily responsible for making the resource Alldredge, Mathew W. Buderman, Frances E. Blecha, Kevin A. Language A language of the resource English Is Part Of A related resource in which the described resource is physically or logically included. Ecology and Evolution Format The file format, physical medium, or dimensions of the resource application/pdf Bibliographic Citation A bibliographic reference for the resource. Recommended practice is to include sufficient bibliographic detail to identify the resource as unambiguously as possible. <p>Alldredge, M. W., F. E. Buderman, and K. A. Blecha. 2019. Human–Cougar interactions in the wildland–urban interface of Colorado's front range. Ecology and Evolution 9:10415–10431. <a href="https://doi.org/10.1002/ece3.5559" target="_blank" rel="noreferrer noopener">https://doi.org/10.1002/ece3.5559</a></p> Extent The size or duration of the resource. 17 pages Type The nature or genre of the resource Article