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A Backyard Puma in Costa Rica: Translocation As a Successful Management Option

Victor H. Montalvo1*, Carolina Sáenz-Bolaños1, Isabel Hagnauer2, Juan C. Cruz-Díaz3, Pablo Vásquez4, Shirley Ramírez4, Todd K. Fuller5, and Eduardo Carrillo1

1Instituto Internacional en Conservación y Manejo de Vida Silvestre, Universidad Nacional, Apdo. 1350-3000, Heredia, Costa Rica. 2ZooAve, Fundación Restauración de la Naturaleza. Apdo. 1327- 4050, Alajuela, Costa Rica. 3Namá Conservation, Heredia 40101, Costa Rica. 4Ministerio de Ambiente y Energía, Departamento de Vida Silvestre, Apdo. 10127-1000, San José, Costa Rica. 5Department of Environmental Conservation, University of Massachusetts, Amherst, Massachusetts 01003, USA. *Corresponding author.

Urban Naturalist Notes, No. 1 (2022)

Abstract
In Costa Rica, most Pumas inhabit forested and rural areas, but during the last decade Puma sightings have increased in urban areas, sometimes resulting in distress to both humans and Pumas. Here we assess the success of translocating an adult male Puma 28 km from the suburbs of San Jose, Costa Rica to Braulio Carrillo National Park and monitored for 87 days post-release via satellite telemetry (n = 445 locations). The Puma used an area of 135 km2 and movements indicated “settling” behavior. This Puma used mostly areas of forest relatively nearer to streams and in proximity to protected areas and did not return to urban areas. Our translocation effort seemed to be an effective means of reducing conflict or potential conflict between Pumas and urban residents, however every translocation should be analyzed individually.

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Volume 9, 2022 Urban Naturalist Notes No. 1 A Backyard Puma in Costa Rica: Translocation As a Successful Management Option Victor H. Montalvo, Carolina Sáenz-Bolaños, Isabel Hagnauer, Juan C. Cruz-Díaz, Pablo Vásquez, Shirley Ramírez, Todd K. Fuller, and Eduardo Carrillo Urban Naturalist The Urban Naturalist (ISSN # 2328-8965) is published by the Eagle Hill Institute, PO Box 9, 59 Eagle Hill Road, Steuben, ME 04680- 0009. Phone 207-546-2821 Ext. 4, FAX 207-546-3042. E-mail: office@eaglehill.us. Webpage: http://www.eaglehill.us/urna. Copyright © 2022, all rights reserved. Published on an article by article basis. Special issue proposals are welcome. The Urban Naturalist is an open access journal. Authors: Submission guidelines are available at http://www.eaglehill.us/urna. Co-published journals: The Northeastern Naturalist, Southeastern Naturalist, Caribbean Naturalist, and Eastern Paleontologist, each with a separate Board of Editors. The Eagle Hill Institute is a tax exempt 501(c)(3) nonprofit corporation of the State of Maine (Federal ID # 010379899). Board of Editors Hal Brundage, Environmental Research and Consulting, Inc, Lewes, DE, USA Sabina Caula, Universidad de Carabobo, Naguanagua, Venezuela Sylvio Codella, Kean University, Union New Jersey, USA Julie Craves, University of Michigan-Dearborn, Dearborn, MI, USA Ana Faggi, Universidad de Flores/CONICET, Buenos Aires, Argentina Leonie Fischer, Technical University of Berlin, Berlin, Germany Chad Johnson, Arizona State University, Glendale, AZ, USA Sonja Knapp, Helmholtz Centre for Environmental Research–UFZ, Halle (Saale), Germany David Krauss, City University of New York, New York, NY, USA Joerg-Henner Lotze, Eagle Hill Institute, Steuben, ME. Publisher Kristi MacDonald, Hudsonia, Bard College, Annandale-on- Hudson, NY, USA Tibor Magura, University of Debrecen, Debrecen, Hungary Brooke Maslo, Rutgers University, New Brunswick, NJ, USA Mike McKinney, University of Tennessee, Knoxville, TN, USA. Journal Editor Desirée Narango, University of Massachusetts, Amherst, MA, USA Zoltán Németh, Department of Evolutionary Zoology and Human Biology, University of Debrecen, Debrecen, Hungary Joseph Rachlin, Lehman College, City University of New York, New York, NY, USA Travis Ryan, Center for Urban Ecology, Butler University, Indianapolis, IN, USA Michael Strohbach, Technische Universität Braunschweig, Institute of Geoecology, Braunschweig, Germany Katalin Szlavecz, Johns Hopkins University, Baltimore, MD, USA Advisory Board Myla Aronson, Rutgers University, New Brunswick, NJ, USA Mark McDonnell, Royal Botanic Gardens Victoria and University of Melbourne, Melbourne, Australia Charles Nilon, University of Missouri, Columbia, MO, USA Dagmar Haase, Helmholtz Centre for Environmental Research–UFZ, Leipzig, Germany Sarel Cilliers, North-West University, Potchefstroom, South Africa Maria Ignatieva, University of Western Australia, Perth, Western Australia, Australia ♦ The Urban Naturalist is a peer-reviewed and edited interdisciplinary natural history journal with a global focus on urban areas (ISSN 2328- 8965 [online]). ♦ The journal features research articles, notes, and research summaries on terrestrial, freshwater, and marine organisms and their habitats. ♦ It offers article-by-article online publication for prompt distribution to a global audience. ♦ It offers authors the option of publishing large files such as data tables, and audio and video clips as online supplemental files. ♦ Special issues - The Urban Naturalist welcomes proposals for special issues that are based on conference proceedings or on a series of invitational articles. Special issue editors can rely on the publisher’s years of experiences in efficiently handling most details relating to the publication of special issues. ♦ Indexing - The Urban Naturalist is a young journal whose indexing at this time is by way of author entries in Google Scholar and Researchgate. Its indexing coverage is expected to become comparable to that of the Institute's first 3 journals (Northeastern Naturalist, Southeastern Naturalist, and Journal of the North Atlantic). These 3 journals are included in full-text in BioOne.org and JSTOR.org and are indexed in Web of Science (clarivate.com) and EBSCO.com. ♦ The journal's staff is pleased to discuss ideas for manuscripts and to assist during all stages of manuscript preparation. The journal has a page charge to help defray a portion of the costs of publishing manuscripts. Instructions for Authors are available online on the journal’s website (http://www.eaglehill.us/urna). ♦ It is co-published with the Northeastern Naturalist, Southeastern Naturalist, Caribbean Naturalist, Eastern Paleontologist, Eastern Biologist, and Journal of the North Atlantic. ♦ It is available online in full-text version on the journal's website (http://www.eaglehill.us/urna). Arrangements for inclusion in other databases are being pursued. Cover Photograph: Local firefighters assisting with the translocation of an adult male Puma captured in the San Juan de Tibás suburbs of San Jose, Costa Rica. Photograph © I. Hagnauer. Urban Naturalist Notes V.H. Montalvo et. al. Vol. 9, 2022 N1:1–7 1 2021 A Backyard Puma in Costa Rica: Translocation As a Successful Management Option Victor H. Montalvo1*, Carolina Sáenz-Bolaños1, Isabel Hagnauer2, Juan C. Cruz- Díaz3, Pablo Vásquez4, Shirley Ramírez4, Todd K. Fuller5, and Eduardo Carrillo1 Abstract - In Costa Rica, most Pumas inhabit forested and rural areas, but during the last decade Puma sightings have increased in urban areas, sometimes resulting in distress to both humans and Pumas. Here we assess the success of translocating an adult male Puma 28 km from the suburbs of San Jose, Costa Rica to Braulio Carrillo National Park and monitored for 87 days post-release via satellite telemetry (n = 445 locations). The Puma used an area of 135 km2 and movements indicated “settling” behavior. This Puma used mostly areas of forest relatively nearer to streams and in proximity to protected areas and did not return to urban areas. Our translocation effort seemed to be an effective means of reducing conflict or potential conflict between Pumas and urban residents, however every translocation should be analyzed individually. Introduction. In Costa Rica, Puma (Puma concolor Linnaeus) populations still regularly occur in protected and rural areas (Carrillo et al. 1999, Wainwright 2007), but Puma sightings in urban areas also have increased in the last decade. Their occurrence in urban areas is of consequence to both Pumas and humans, because conflict could arise if a Puma preys on domestic pets or otherwise frequents backyards and scares residents (Fontúrbel and Simonetti 2011, Linnell et al. 1997). Non-lethal methods are highly recommended to deal with humancarnivore conflict in such areas (Priatna et al. 2012, Weise et al. 2015), and thus translocation of “problem” animals has become an option for management and conservation (Fontúrbel and Simonetti 2011, Linnell et al. 1997). But despite a wealth of recent research on and management of Pumas across their range (e.g., Hornocker and Negri 2010), only a few studies have followed up the behavior and fate of translocated Pumas after release (Adania et al. 2017, Belden and Hagedorn 1993, Ross and Jalkotzy 1995, Ruth et al. 1998), and much less is known about Puma movements in Central America than other regions (Laundre and Hernandez 2010). Here we report on the movements and habitat use patterns of an adult male Puma translocated from an urban area to the nearest protected area and monitored for 3 months and reflect on translocation as an effective conflict-mitigation strategy. Methods. This tale began on 9 March 2019 in the San Juan de Tibás suburb area about 4 km north of downtown San José, the capital city of Costa Rica (pop. = 2.1 million in metropolitan area; https://www.nationsonline.org/oneworld/costa_rica.htm) (Fig. 1; 9°57'38N, 84°04'47W). Elevation in Tibás ranges from 900 m to 1150 m, with an average temperature of 23°C and annual precipitation of 290 cm (Instituto Meteorológico Nacional, San José, Costa Rica; https://www.imn.ac.cr/en/mapa). The capture site was adjacent to a riparian corridor of the Río Virilla that extends east towards its headwaters near Irazu Volcano and south of Braulio Carrillo National Park (BCNP; 600–1,500 m elevation) where the Puma was subsequently translocated. The 476-km2 BCNP was es- 1Instituto Internacional en Conservación y Manejo de Vida Silvestre, Universidad Nacional, Apdo. 1350-3000, Heredia, Costa Rica. 2ZooAve, Fundación Restauración de la Naturaleza. Apdo. 1327- 4050, Alajuela, Costa Rica. 3Namá Conservation, Heredia 40101, Costa Rica. 4Ministerio de Ambiente y Energía, Departamento de Vida Silvestre, Apdo. 10127-1000, San José, Costa Rica. 5Department of Environmental Conservation, University of Massachusetts, Amherst, Massachusetts 01003, USA. *Corresponding author: victor.montalvo.guadamuz@una.cr. Associate Editor: Michael Strohbach, Technische Universität Braunschweig, Institute of Geoecology. Vol. 9, 2022 URBAN NATURALIST NOTES N1:1–7 Urban Naturalist Notes V.H. Montalvo et. al. Vol. 9, 2022 N1:1–7 2 tablished in 1978 and is located in the central mountain range of Costa Rica; more than 90% of the park is covered in primary forest. A Puma that was apparently unable to escape from a walled backyard of neighborhood house prompted a “911” emergency call from a local resident, resulting in a response from personnel of the Tibás fire and police departments who were subsequently joined by a veterinarian, by personnel from the Ministry of Environment and Energy, and by researchers from the National University. We immobilized the Puma using a dart projectile (Dan-Inject, Kolding, Denmark; https://www.dan-inject.com) filled with a combination of 5 mg/kg of ketamine (Bremer Pharma GmbH, Warburg, Germany) and 2mg/kg xylazine (Procin Equus 10%, Pisa Agropecuaria). The 39-kg adult (indicated by tooth wear) male was fitted with a GPS collar (Lotek Engineering, Newmarket, ON, Canada; http://www.lotek.com) programmed to record the Puma’s position every 2 hours. Three hours after immobilization, the Puma was released in Braulio Carrillo National Park, 28 km from the capture site and east of the main highway that crosses the mountain range to the Pacific lowlands (Fig 1.; 10°09'42N, 83°56'14W). Handling and capture protocols followed guidelines of the American Society of Mammologists (Sikes et al. 2011). Figure 1. Monthly locations and modified minimum polygon delineation of monthly ranges of a male Puma translocated from the suburbs of San Jose to Braulio Carrillo National Park in central Costa Rica on 9 March 2019. Urban Naturalist Notes V.H. Montalvo et. al. Vol. 9, 2022 N1:1–7 3 We used the location data to assess whether or not the Puma settled into a home range away from urban areas. We estimated the size of the area traversed by the Puma each month by drawing modified minimum polygons (Harvey and Barbour 1965) after plotting and connected the sequential locations of the Puma, and assessed temporal changes in movement rates for each month by summarizing the calculated distances between first locations on each pair of consecutive days. To test patterns of habitat use, we generated 500 random locations within the area of a larger 95% autocorrelated kernel range (Signer and Balkenhol 2015; estimated for the entire monitoring period) to contrast (using a c2-test) expected (random) land used frequencies with observed Puma locations. Available locations were randomly generated across six land use categories: 1) Forest - primary and secondary forest (62% availability), 2) Human infrastructure - roads and buildings (6%), 3) Non-forest - pasture, hayfield, shrubland (17%), 4) Forest plantation (1%), 5) Agriculture - crops, coffee plantations (1%), and 6) Other - cloud-covered (no data) and un classified (13%). We also calculated for each location the distance to unpaved roads, urban infrastructure, forest cover, protected areas, and rivers/streams (i.e., water). The spatial layers were based on a vegetation coverage produced by Sistema Nacional de Areas de Conservación (SINAC; http://www.sinac.go.cr/EN-US/Pages/default.aspx) and the Costa Rica Digital Atlas (Ortiz- Malavasi 2014). We fit a set of Generalized lineal Models (GLM), assuming binomial distribution (Di Blanco et al. 2015), for the previous distance variables to additively test the intensity of selection related to habitat attributes. The empirical support of candidate models was evaluated using the Akaike Information Criterion (AIC) (Burnham and Anderson 2002), and throughout the model comparison we determined the most plausible models using the highest Akaike weight (⍵ ; range = 0 to 1; Anderson 2007). Results and Discussion. In total, 445 locations (42% of potential) were registered during 87 continuous days of post-translocation monitoring (9 March–4 June 2019), and average of ~5 locations per day. Though the Puma’s satellite telemetry collar was programed for a 2-year life, we lost its signal after 87 days and were unable to identify whether the collar failed, or the Puma was killed and the collar destroyed. Within an overall 3-month range of 135 km2, monthly range size decreased from 92 to 32 km2 (Table 1, Fig. 1). As the Puma’s range size decreased, the average distance moved on consecutive days (1.8 to 2.0 km/day) and the median distance (1.3 to 1.9 km/day) stayed about the same (Table 1). The core area of where it seemed to settle was about 21 km from its capture location, and 13 km from its release site (Fig. 1). Habitat use analysis indicated that the Puma was not located in human infrastructure (6% available), agriculture (1%), or forest plantation (1%) land use categories, and used forest (74% use) more, and non-forest habitats less, than expected (7% use; c2 = 28.13, P < 0.0001). Although the Puma explored areas closer to San Jose in April, it seemed to retreat to and settle in more remote areas by May (Fig. 1). The most plausible and statistically supported distance-related habitat model included all five variables (Table 2; ⍵=0.99). Distance to each showed a negative relationship with Puma presence of this single male individual; e.g., as proximity to a variable (e.g., unpaved roads) increased, so did Puma presence (Table 3). Though all variables assessed show statistical significance, distance to forest was the most influential v ariable, followed by distance to water; an additional correlation test showed none between forest and water (t = 0.163, d.f. = 443, P = 0.870, Cor = 0.008). Experts state that there are no truly urban Puma populations, only populations abutting urban areas into which dispersing juveniles and the occasional adult will enter (Beier et al. 2010). This seems the case here with an adult male following a riverine corridor from the mountains and ending up in a backyard. Urban Naturalist Notes V.H. Montalvo et. al. Vol. 9, 2022 N1:1–7 4 Recent translocation mitigation efforts for large felids, though often politically expedient, have had mixed success (Hoogesteijn et al. 2020, Morapedi et al. 2021, Priatna et al. 2012, Weise et al. 2015), and translocation, in general, is a rarely used conservation technique for Pumas, sometimes considered ineffectual (Sweanor and Logan 2010). Notably, it has been used successfully for augmenting the population of Florida panthers (Benson et al. 2011, Hostetler et al. 2013, Johnson et al. 2010), and thus has been suggested as a future measure to enhance Puma populations isolated by urbanization effects (Vickers et al. 2015). However, translocations are not much used for mitigating livestock conflicts (Guerisoli et al. 2020), or especially for removing potential interactions in urban settings. Still, public attitudes indicate that translocations might be the preferred management options as Puma-human interactions increase (Manfredo et al. 1998, Thornton and Quinn 2009). Previous translocations of Pumas seem to suggest that Pumas will settle fairly rapidly and behave much like non-translocated animals (Belden and Hagedorn 1993, Ross and Jalkotzy 1995, Ruth et al. 1998, Adania et al. 2017). Notably, our report of the translocation of a single male Puma in Costa Rica is the first done in order to mitigate its occurrence in an urban area. We moved it 28 km from the capture site assuming that, due to the riparian connectivity from the capture site (Río Virilla) to a national park, the Puma came from there. Male Pumas dispersing Table 1. Monthly range size (modified minimum polygon) and daily movements of a translocated male Puma in central Costa Rica. Month Range Daily Distance (km) Moveda Size (km2) N Mean Median Range N March 92 104 1.92 1.31 0.01-7.30 21 April 49 126 1.80 1.28 0.03-5.59 30 May 32 215 1.99 1.89 0.01-6.36 31 aDistance between first locations on each date on consecutive day s. Table 2. Ranked candidate models of habitat selection for a male Puma translocated central Costa Rica (AIC = Akaike Information Criterion; Δ AIC = Akaike Information Criterion differences from the top model; ⍵ = Likelihood empirical weight). Modela AIC Δ AIC ⍵ Use ~ DR+ DU +DF + DPA + DW 1129.5 0 0.99 Use ~ DR+ DU +DF + DPA 1140.5 11.1 0.001 Use ~ DR+ DU +DF 1164.3 34.8 <0.001 Use ~ DF 1188.5 59.0 <0.001 Use ~ DR+ DU 1219.4 89.9 <0.001 Use ~ DPA 1220.2 90.8 <0.001 Use ~ DU 1227 97.5 <0.001 Use ~ DW 1227.2 97.8 <0.001 Use ~ DR 1231.9 102.4 <0.001 Use ~ Intercept 1237.2 107.7 <0.002 aDR = Distance to unpaved Roads; DU = Distance to Urban infrastructure; DF = Distance to Forest cover; DPA = Distance to Protected Areas; DW = Distance to Water. Urban Naturalist Notes V.H. Montalvo et. al. Vol. 9, 2022 N1:1–7 5 from their natal ranges typically move 100 km and more, and when encountering urban areas often move along wildland-urban edges, make frequent forays in peninsulas of habitat that intrude into urban areas, and may settle in habitats at the urban edge (Beier et al. 2010). During the 87 days we monitored the Puma, we documented that it moved from the release location, explored areas closer to San Jose, then seemed to settle back in more remote areas. The overall area it used during 3 months (135 km2) was similar to annual home range sizes of resident Pumas near urban areas elsewhere (Dickson and Beier 2002, Laundré and Loxterman 2007). Also, daily movements of our male Puma (1.8–2.0 km) were similar to male Pumas monitored in Mexico (Nuñez and Miller 2019). Habitat use analysis showed the importance of distance to forest cover and streams to Puma occurrence, followed by distance to protected areas, similar to the pattern for Pumas across their distribution in the wild (Guerisoli et al. 2019). Use of developed areas by Pumas substantially increased their risk of mortality (Moss et al. 2016), and thus fear of humans may be a major factor affecting Puma movements (Nickel et al. 2021, Smith et al. 2019). Previous post-translocation information on large carnivores indicates that translocated individuals have reduced survival and reproduction (Linnell et al. 1997), since moving throughout unknown areas would increase loss of human fear, stress, vehicle collision, and starvation (Boast et al. 2018, Ruth et al. 1998, Seddon et al. 2012), and sometimes a propensity to return to original capture sites (Ruth et al. 1998). Common factors associated with successful large felid translocations suggests that Pumas should be moved to areas of: 1) low density of intrainterspecific competitors, 2) low human density, 3) high prey availability, and 4) large size (Belden and Hagedorn 1993, Boast et al. 2018, Hoogesteijn et al. 2020, Priatna et al. 2012); for Pumas, animals that are not yet of breeding age or established a home range may be more successful, as well (Sweanor and Logan 2010). Although, there are protocols and translocation manuals available for some species (e.g., Ceballos 2018), each potential translocation should be assessed individually to allow translocated individuals to successfully established a home range (Linnell et al. 1997, Ruth et al. 1998). In addition, it may be useful in the future to help reduce conflicts with Pumas in urban areas by increasing the awareness of Pumas through targeted education, facilitating of stakeholder involvement, developing of proactive Puma management strategies, and exploring adaptive management (Thornton and Quinn 2009). Acknowledgements We thank the Tibás fire and police departments for their valuable help during the Puma translocation. We also thank Fabricio Carbonell from the Área de Conservación Cordillera Volcánica, the two anonymous reviewers for helping to improve this manuscript and the Project Survival Cat Haven for donating the satellite collar. Table 3. Coefficients of explanatory variables of the top model for a resource selection function of a translocated male Puma in central Costa Rica. Coefficientsa ß SE Z-value P value Intercept 1.645 0.215 7.645 < 0.00001 DR -0.003 <0.001 -3.550 < 0.00001 DU -0.001 <0.001 -4.884 < 0.00001 DF -0.229 0.058 -3.961 < 0.00001 DPA -0.007 0.001 -5.022 < 0.00001 DW -0.220 0.006 -3.593 <0.00001 aDR = Distance to unpaved Roads; DU = Distance to Urban infrastructure; DF = Distance to Forest cover; DPA = Distance to Protected Areas; DW = Distance to Water. Urban Naturalist Notes V.H. Montalvo et. al. Vol. 9, 2022 N1:1–7 6 Data Availability Statement The data that support the findings of this study are available on request from the corresponding author. Literature Cited Adania, C., W.D. De Carvalho, L.M. Rosalino, J. De Cassio, and P. Crawshaw. 2017. First soft release of a relocated Puma in South America. Mammal Research 62:121–128. Anderson, D.R. 2007. 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