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Ocelots (Leopardus pardalis) are a wide-ranging felid species, occurring from southern United States to northern Argentina. They occupy various habitats and are usually considered the most abundant wild cat species in the Neotropics. However, genetic studies that include free-ranging Mesoamerican ocelots are rare and generally based on small sample sizes. This is the first conservation genetics study on ocelots in Costa Rica and the second one in Mesoamerica that has conducted a genetic assessment of the species at a countrywide scale. We evaluated genetic diversity and population structure of ocelots using 15 microsatellite loci in 28 successfully genotyped individuals from throughout the country. We also compared genetic diversity of Costa Rican ocelots with that of jaguars (Panthera onca) and pumas (Puma concolor) in the country, and with ocelots in Belize. Genetic diversity of ocelots in Costa Rica was relatively high as measured by rarified allelic richness (AR = 5.50 ± 1.36) and expected heterozygosities (HE = 0.79 ± 0.08). We did not detect patterns of genetic substructure, suggesting high levels of gene flow throughout the country and no strong barriers to movement. As expected, genetic diversity of Costa Rican ocelots was higher than co-occurring jaguars and pumas. Additionally, levels of genetic diversity were slightly higher in Costa Rican ocelots when compared with their counterparts in Belize, confirming the south to north decrease in genetic diversity reported in other studies. Our study provides critical baseline information to understand the status of wild ocelot populations in Costa Rica. Future studies on ocelots and other threatened or keystone species should also integrate genetic monitoring and conservation genetics analysis to properly inform management decisions, guarantee their long-term survival, and improve the resilience of ecosystems.

Evaluating range-wide habitat use by a target species requires information on species occurrence over broad geographic regions, a process made difficult by species rarity, large spatiotemporal sampling domains, and imperfect detection. We address these challenges in an assessment of habitat use for jaguars (Panthera onca) outside protected areas in Central America. Occurrence records were acquired within 12 putative corridors using interviews with knowledgeable corridor residents. We developed a Bayesian hierarchical occupancy model to gain robust inference, allowing for heterogeneity introduced in the sampling process over space and time, using records of jaguar occurrence prone to false positives and false negatives. Probability of false detection of jaguars increased with the number of interviews conducted per unit (from 5.42% to 7.74% given

Large mammalian herbivores are experiencing population reductions and range declines. However, we lack regional knowledge of population status for many herbivores, particularly in developing countries. Addressing this knowledge gap is key to implementing tailored conservation strategies for species whose population declines are highly variable across their range. White-lipped peccaries (Tayassu pecari) are important ecosystem engineers in Neotropical forests and are highly sensitive to human disturbance. Despite maintaining a wide distributional range, white-lipped peccaries are experiencing substantial population declines in some portions of their range. We examined the regional distribution and population status of the species in Mesoamerica. We used a combination of techniques, including expert-based mapping and assessment of population status, and data-driven distribution modelling techniques to determine the status and range limits of white-lipped peccaries. Our analysis revealed declining and highly isolated populations of peccaries across Mesoamerica, with a range reduction of 87% from historic distribution and 63% from current IUCN range estimates for the region. White-lipped peccary distribution is affected by indices of human influence and forest cover, and more restricted than other sympatric large herbivores, with their largest populations confined to transboundary reserves. To conserve white-lipped peccaries in Mesoamerica, transboundary efforts will be needed that focus on both forest conservation and hunting management, increased cross-border coordination, and reconsideration of country and regional conservation priorities. Our methodology to detail regional white-lipped peccary status could be employed on other poorly-known large mammals. Universidad Nacional, Costa Rica El Colegio de la Frontera Sur, México Wildlife Conservation Society, United States Instituto Internacional en Conservación y Manejo de Vida Silvestre

Partial depigmentation (leucism) is rare in mammals. We report 17 camera trap records of a single female leucistic coyote (Canis latrans) in Costa Rica between 2014 and 2018. Resumen “Hembra de Canis latrans (Carnivora: Canidae) in Costa Rica”. La despigmentación parcial (leucismo) es poco común en mamíferos. Aquí reportamos un total de 17 registros fotográficos de una hembra leucística de coyote (Canis latrans) en Costa Rica entre 2014 y 2018.

Connectivity of natural areas through biological corridors is essential for ecosystem resilience and biodiversity conservation. However, robust assessments of biodiversity in corridor areas are often hindered by logistical constraints and the statistical challenges of modeling data from multiple species. Herein, we used a hierarchical community occupancy model in a Bayesian framework to evaluate the status of medium and large-sized mammals in a critical link of the Mesoamerican Biological Corridor (MBC) in Costa Rica. We used camera traps deployed from 2013–2017 to detect 18 medium (1–15 kg) and 6 large (>15 kg) mammal species in a portion of two Jaguar Conservation Units (JCUs) and the Corridor linking them. Camera traps operated for 16,904 trap nights across 209 stations, covering an area of 880 km2. Forest cover was the most important driver of medium and large-sized mammal habitat use, with forest specialists such as jaguars (Panthera onca) and pumas (Puma concolor) strongly associated with high forest cover, while habitat generalists such as coyotes (Canis latrans) and raccoons (Procyon lotor) were associated with low forest cover. Medium and large-sized mammal species richness was lower in the Corridor area (x¯= 9.78±1.84) than in the portions evaluated of the two JCUs (x¯= 11.50±1.52). Puma and jaguar habitat use probabilities were strongly correlated with large prey species richness (jaguar,r= 0.59, pr= 0.72, pr= 0.36, p = 0.003; puma,r= 0.23, p = 0.064). Low estimated jaguar habitat use probability in one JCU (Central Volcanic Cordillera:x¯= 0.15±0.11) suggests that this is not the jaguar stronghold previously assumed. In addition, the western half of the Corridor has low richness of large mammals, making it necessary to take urgent actions to secure habitat connectivity for mammal populations.

Poca información ha sido generada sobre la biodiversidad del Refugio Nacional de Vida Silvestre Barra del Colorado, Costa Rica. En el presente estudio determinamos la abundancia relativa y el patrón de actividad de ciertas especies de mamíferos terrestres. Durante el periodo 2014 se empleamos diez cámaras trampa, las cuales fueron distribuidas dentro del refugio. Después de un total de 1 611 noches de muestreo, se identificaron 15 especies de mamíferos distribuidos en siete órdenes y 11 familias. Las especies más abundantes fueron Dasyprocta punctata, Leopardus pardalis, Tayassu pecari,Mazama temama, Pecari tajacu y Tapirus bairdii, mientras que Tamandua mexicana reportó la menor abundancia. Se estimaron los patrones de actividad para ocho especies, los cuales fueron similares a lo previamente descrito en la literatura. Estudios adicionales son necesarios con el fin de incrementar nuestro conocimiento sobre la biodiversidad del refugio, información que será esencial para el adecuado manejo del área.

El intercambio ecológico y la viabilidad de la biodiversidad de Costa Rica se ven amenazados por ladegradación y la fragmentación del bosque. Esta investigación pretende proponer una herramienta paraidentificar el posible impacto de las actividades pecuarias sobre áreas de conservación del Caribe. Para ello, serealizó la extracción de ADN de las heces de 14 jaguares (Panthera onca) y 13 pumas (Puma concolor) de vidalibre mediante el kit Fujifilm, Fuji. Estas fueron identificadas previamente por individuo y especie, mediantemétodos moleculares. Luego, se cuantificaron 16 genes de resistencias a los antimicrobianos (ARGs) (tetA,tetB, tetC, tetK, tetM, tetQ, tetS, tetW y tetY, catI, catII, sulI, sulII, qnrS, vanA y mecA) mediante qPCR. Lacaracterización de los sitios de muestreo se hizo mediante QGIS y ArGIS, al emplear datos geoespacialesde la región y el censo de SENASA 2014. De los 14 ARGs hallados, 9.3 % tenían una resistencia baja, 16 %intermedia y 13.9 % alta. Los genes tetQ (85.2 %) y tetY (70.3 %) fueron los más frecuentes, mientras que tetS(11.1 %) fue el menos usual. Las sulfonamidas (sulI y sulII) (70.3 % cada uno), fenicoles (catII y catI) (18.5 %y 52 %, respectivamente) y quinolonas (qnrS) (11 %) presentaron valores representativos. No se identificarongenes de vancomicina (vanA) ni meticilina (mecA). Los jaguares tuvieron más número de ARGs (37 %). ElParque Nacional Braulio Carrillo fue el área donde se encontraron las más altas concentraciones de tet, suly cat. El gen qnrS se identificó, principalmente, fuera de áreas protegidas. De acuerdo con el análisis integralde los datos, el impacto pecuario podría influir sobre la presencia y concentración de ARGs.

Little is known about the effects of jaguars on the population of marine turtles nesting in Tortuguero National Park, Costa Rica. This study assessed jaguar predation impact on three species of marine turtles (Chelonia mydas, Dermochelys coriáceaand Eretmochelys imbricata)that nest in Tortuguero beach. Jaguar predation data was obtained by using two methodologies, literature review (historical records prior the year 2005) and weekly surveys along the 29 km stretch of beach during the period 2005-2013. Our results indicated that jaguar predation has increased from one marine turtle in 1981 to 198 in 2013. Jaguars consumed annually an average of 120 (SD= 45) and 2 (SD= 3) green turtles and leatherbacks in Tortuguero beach, respectively. Based on our results we concluded that jaguars do not represent a threat to the population of green turtles that nest in Tortuguero beach, and it is not the main cause for population decline for leatherbacks and hawksbills. Future research should focus on continuing to monitor this predator-prey relationship as well as the factors that influence it so the proper management decisions can be taken.

Due largely to their success in undisturbed areas, the conservation of large carnivores is tied to protected areas. However, because of their large area requirements, large carnivores – and their conservation – will also always be linked to areas outside of protected areas. Retaliatory killing of jaguars near reserves and in corridors between reserves threatens to severely decrease jaguar populations in protected areas and their corridors, thus decreasing the ecological viability of protected areas for the species. Resolving this conflict will allow jaguars to utilize the human-dominated landscape, provide opportunities for corridor conservation, and enhance the effectiveness of protected areas. We review methods to reduce depredation of livestock and prevent lethal control of jaguars. Approaches to mitigate conflict include insurance schemes, management of wild prey, and above all, improving livestock husbandry and management. Improvements that are recommended for specific problem farms and ranches might include electric fencing, night enclosures, designs for newborn holding pens, the use of guard animals, and partial herd immersion of creole cattle races or water buffalo. We describe multiple testing scenarios and results from throughout Latin America for reducing livestock depredation across protected areas and agricultural landscapes.

Broad scale population estimates of declining species are desired for conservation efforts. However, for many secretive species including large carnivores, such estimates are often difficult. Based on published density estimates obtained through camera trapping, presence/absence data, and globally available predictive variables derived from satellite imagery, we modelled density and occurrence of a large carnivore, the jaguar, across the species’ entire range. We then combined these models in a hierarchical framework to estimate the total population. Our models indicate that potential jaguar density is best predicted by measures of primary productivity, with the highest densities in the most productive tropical habitats and a clear declining gradient with distance from the equator. Jaguar distribution, in contrast, is determined by the combined effects of human impacts and environmental factors: probability of jaguar occurrence increased with forest cover, mean temperature, and annual precipitation and declined with increases in human foot print index and human density. Probability of occurrence was also significantly higher for protected areas than outside of them. We estimated the world’s jaguar population at 173,000 (95% CI: 138,000–208,000) individuals, mostly concentrated in the Amazon Basin; elsewhere, populations tend to be small and fragmented. The high number of jaguars results from the large total area still occupied (almost 9 million km2) and low human densities (< 1 person/km2) coinciding with high primary productivity in the core area of jaguar range. Our results show the importance of protected areas for jaguar persistence. We conclude that combining modelling of density and distribution can reveal ecological patterns and processes at global scales, can provide robust estimates for use in species assessments, and can guide broad-scale conservation actions.

The purpose of protected areas is to safeguard natural habitats and their biodiversity. However, few protected areas are large enough for the long-term conservation of resident species, making the landscapes surrounding protected areas equally important for species conservation. This is especially true for wide-ranging carnivores such as the jaguar (Panthera onca), for which conservation strategies stress the maintenance of range-wide habitat connectivity via protected areas and dispersal corridors. We used 587 interviews with local people in 441 36-km2 sampling units to assess the presence of jaguar and six prey species in a critical corridor in Central America, linking Bosawas Biosphere Reserve and Wawashan Nature Reserve in Nicaragua. We analyzed these data using single-season false-positive detection models in a site occupancy framework. We found that agricultural encroachment, particularly pervasive within a protected area, was the main factor limiting the presence of jaguar and three large-bodied prey species: white-lipped peccary (Tayassu pecari), collared peccary (Pecari tajacu), and red brocket deer (Mazama americana). We suggest that improperly-managed protected areas can suffer the loss of apex predators and their prey, thereby jeopardizing range-wide conservation strategies for threatened species.

Corridors are critical elements in the long-term conservation of wide-ranging species like the jaguar ( Panthera onca ). Jaguar corridors across the range of the species were initially identified using a GIS-based least-cost corridor model. However, due to inherent errors in remotely sensed data and model uncertainties, these corridors warrant field verification before conservation efforts can begin. We developed a novel corridor assessment protocol based on interview data and site occupancy modeling. We divided our pilot study area, in southeastern Nicaragua, into 71, 6 × 6 km sampling units and conducted 160 structured interviews with local residents. Interviews were designed to collect data on jaguar and seven prey species so that detection/non-detection matrices could be constructed for each sampling unit. Jaguars were reportedly detected in 57% of the sampling units and had a detection probability of 28%. With the exception of white-lipped peccary, prey species were reportedly detected in 82–100% of the sampling units. Though the use of interview data may violate some assumptions of the occupancy modeling approach for determining ‘proportion of area occupied’, we countered these shortcomings through study design and interpreting the occupancy parameter, psi, as ‘probability of habitat used’. Probability of habitat use was modeled for each target species using single state or multistate models. A combination of the estimated probabilities of habitat use for jaguar and prey was selected to identify the final jaguar corridor. This protocol provides an efficient field methodology for identifying corridors for easily-identifiable species, across large study areas comprised of unprotected, private lands.

The jaguar Panthera onca is threatened throughout its range and categorized as Near Threatened on the IUCN Red List. To inform conservation of the jaguar population in Corcovado National Park, Costa Rica, population size was estimated using data from a 3-month camera trap study. Individuals were identified from their coat patterns. The resulting density estimate of 6.98 ± SD 2.36 individuals per 100 km2 was lower than expected. The sex ratio was 1.33 males per female, and the minimum home ranges of two males were 25.64 and 6.57 km2. Hunting pressure on jaguar and white-lipped peccaries Tayassu pecari, the jaguar's main prey in the Park, may be responsible for the low jaguar density as space does not seem to be a limiting factor. The numbers of females may have been underestimated because of sampling bias and therefore the sex ratio obtained in this and similar studies must be interpreted cautiously. Better protection of the corridor that connects the Park with other protected areas is essential to guarantee long-term survival of the jaguar in Costa Rica.

Little is known about the effects of jaguars on the population of marine turtles nesting in Tortuguero National Park, Costa Rica. This study assessed jaguar predation impact on three species of marine turtles (Chelonia mydas, Dermochelys coriacea and Eretmochelys imbricata) that nest in Tortuguero beach. Jaguar predation data was obtained by using two methodologies, literature review (historical records prior the year 2005) and weekly surveys along the 29 km stretch of beach during the period 2005-2013. Our results indicated that jaguar predation has increased from one marine turtle in 1981 to 198 in 2013. Jaguars consumed annually an average of 120 (SD= 45) and 2 (SD= 3) green turtles and leatherbacks in Tortuguero beach, respectively. Based on our results we concluded that jaguars do not represent a threat to the population of green turtles that nest in Tortuguero beach, and it is not the main cause for population decline for leatherbacks and hawksbills. Future research should focus on continuing to monitor this predator-prey relationship as well as the factors that influence it so the proper management decisions can be taken. Existe poco conocimiento sobre el impacto que tienen los jaguares sobre las tortugas marinas que anidan en el Parque Nacional Tortuguero, Costa Rica. Este estudio evaluó el impacto de la depredación de jaguar sobre tres especies de tortugas marinas (Chelonia mydas, Dermochelys coriácea y Eretmochelys imbricata) que anidan en Tortuguero. Los reportes de depredación fueron obtenidos empleando dos metodologías, revisión literaria (eventos registrados antes del 2005) y monitoreos semanales a lo largo de la playa (durante el periodo 2005-2013). La depredación del jaguar se ha incrementado de una tortuga en 1981 a 198 tortugas en 2013. Asimismo, los jaguares consumieron anualmente un promedio de 120 (SD= 45) y 2 (SD= 3) tortugas verdes y tortugas baula en Tortuguero, respectivamente. Nuestros resultados indican que los jaguares no representan una amenaza para la población de tortugas verdes que anida en Tortuguero, y no son la causa principal de la disminución poblacional de la tortuga baula y carey. No obstante, se recomienda continuar con el monitoreo con el fin de entender cómo esta interacción depredador-presa evolucionará en el futuro.

ResumenLa Iniciativa del Corredor del Jaguar pretende mantener la conectividad de esta especie. En Costa Rica, esta iniciativa inició en el Subcorredor Biológico Barbilla-Destierro. En este Subcorredor, se ha identificado la pavimentación de un camino de lastre (asociado a las obras de mitigación social del Proyecto Hidroeléctrico Reventazón), como una potencial barrera para la conectividad del jaguar (Panthera onca) y otras especies. Este estudio sugiere un método para identificar los cruces de fauna sobre el camino. Se tomaron datos de junio del 2011 a enero del 2012. Se compararon los registros de avistamientos y atropellos entre una sección de lastre y una de pavimento del mismo camino. Para identificar los pasos de fauna se sobrepusieron distintas fuentes de información: recorridos, entrevistas y conectividad estructural. Al comparar los caminos se encontró que en el pavimento circulan más vehículos, hay más casas, hay más atropellos y se registran menos especies y avistamientos de fauna. Se identificaron tres sitios de cruces de fauna y cinco sectores. Se registró el cruce del camino por jaguar y caucel (Leopardus wiedii) y el uso de una alcantarilla por mapaches (Procyonlotor). Esta metodología puede ser implementada por proyectos viales para identificar los sitios de cruce de fauna y brindar recomendaciones sobre las medidas ambientales de mitigación a implementar asociadas a mejoras o construcción de caminos. AbstractThe Jaguar Corridor Initiative aims to maintain the connectivity of this species. In Costa Rica, this initiative started on the Barbilla-Destierro Biological Subcorridor. The future paving of a road as an offset measure of Reventazon Hydroelectric Project was identified as a potential barrier for the connectivity of the jaguar (Panthera onca) and other animals. In this study we suggest a method that can be used to identify animal crossing points on a gravel road. Data was collected from June 2011 to January 2012. We made a comparison between gravel and paved sections of the same road. To identify animal crossing sites and sectors, information from surveys, interviews and structural connectivity was overlapped. When comparing the gravel and paved sections, the paved section had more cars using it, more houses at its edges, more wildlife roadkills and fewer sightings and species. Three wildlife crossings sites and five sectors were identified. Jaguar and margay (Leopardus wiedii) were registered crossing the road, and raccoons (Procyon lotor) frequently used a culvert as an underpass. This methodology may be used as a practical tool that can be implemented by government and enterprises to identify wildlife crossing sites and to recommend mitigation actions that would reduce the impact caused by the creation and improvement of roads in the region.

Habitat characteristics and human activities are known to play a major role in the occupancy of jaguars Panthera onca across their range, however the key variables influencing jaguar distribution in Tortuguero National Park, Costa Rica, have yet to be identified. This study evaluated jaguar occupancy in Tortuguero National Park and the surrounding area. Jaguar detection/non-detection data was collected using digital camera traps distributed within the boundaries of the protected area. Local community members were also interviewed to determine jaguar occurrence in the Park’s buffer zone. Occupancy models were then applied to identify the habitat characteristics that may better explain jaguar distribution across the study area. From June 2012 to June 2013, a total of 4 339 camera trap days were used to identify 18 individual jaguars inside the protected area; 17 of these jaguars were exclusively detected within the coastal habitat, whilst the remaining individual was detected solely within the interior of the Park. Interviewees reported 61 occasions of jaguar presence inside the buffer zone, between 1995 and 2013, with 80% of these described by the communities of Lomas de Sierpe, Barra de Parismina and La Aurora. These communities also reported the highest levels of livestock predation by jaguars (85% of attacks). In the study area, jaguar occurrence was positively correlated with the seasonal presence of nesting green turtles Chelonia mydas, and negatively correlated with distance to the Park boundary. Our findings suggested that the current occupancy of the jaguar in the study area may be a response to: 1) the vast availability of prey (marine turtles) on Tortuguero beach, 2) the decline of its primary prey species as a result of illegal hunting inside the Park, and 3) the increase in anthropogenic pressures in the Park boundaries. Rev. Biol. Trop. 62 (4): 1449-1458. Epub 2014 December 01. Las características del hábitat y las actividades humanas juegan un papel importante en la presencia del jaguar Panthera onca en toda su área de distribución, sin embargo, las variables clave que influyen en la distribución del jaguar en el Parque Nacional Tortuguero, Costa Rica, aún no se han identificado. Por lo tanto se evaluó la presencia del jaguar Panthera onca en este parque nacional y su área de amortiguamiento. Se recolectaron datos de detección/no detección del jaguar mediante cámaras trampa ubicadas dentro del parque, y se realizaron encuestas en las comunidades del área de amortiguamiento. Posteriormente, se emplearon modelos de ocupación para identificar los atributos del hábitat que mejor explicaban la presencia del felino en el área. Se identificaron 18 jaguares dentro del parque, de los cuales 17 estuvieron exclusivamente en el hábitat costero. En el área de amortiguamiento, las comunidades con una mayor presencia del felino (Lomas de Sierpe, Barra de Parismina y La Aurora) coincidieron con las zonas más conflictivas, en relación a la cacería y la depredación de jaguar sobre el ganado. La probabilidad de ocupación del jaguar se incrementa a medida que aumenta la presencia de la tortuga verde Chelonia mydas, y disminuye conforme la distancia al límite del parque se incrementa. Nuestros resultados indican que la actual presencia del jaguar se debe a: 1) la alta disponibilidad de presas (tortugas marinas) en el hábitat costero, 2) la disminución de las principales especies presa como resultado de la cacería y 3) al incremento de las actividades humanas en el área de amortiguamiento del parque.

Olfactory communication among mammals remains poorly studied yet may be key to understanding their ecology. This is particularly true for mammalian carnivores, which rely extensively on scent marking for communication. Previous research suggests that carnivore latrines play a large role in both intra- and interspecific communication. Despite the apparent complexity of mammal use of latrines, little work has examined behavior patterns of species that visit latrines. We used motion-triggered video cameras to study use and behavior of mammals at ocelot ( Leopardus pardalis ) latrines in Costa Rica. We documented temporal patterns of use by the focal species (ocelots), diversity of mammalian species using latrines, and behaviors that occurred at these sites. Ocelots showed peaks in visitation every 60–67 days and a shorter gap between intersexual versus intrasexual visits, supporting the idea that ocelot latrines are used to communicate information about reproductive status. Fourteen terrestrial mammal species visited the latrines, and these species engaged in a variety of behaviors, including mark investigation, scent marking, and acceptance of scent marks. The complexity and frequency of behaviors by nonfocal species suggest that latrines may play as important a role in communication for these other species as they do for ocelots. La comunicacion olfativa entre mamiferos ha sido poco estudiada, sin embargo puede ser la clave para el entendimiento de su ecologia. Esto es particularmente cierto para mamiferos carnivoros quienes dependen extensivamente de el marcaje de rastros olfativos para su comunicacion. Estudios previos siguieren que las letrinas juegan un rol importante en la comunicacion intra- e inter-especifica. A pesar de la aparente complejidad del uso de letrinas por parte de mamiferos, pocos trabajos han examinado los patrones de comportamiento de las especies que las visitan. Utilizamos camaras de videos sensibles al movimiento para estudiar el uso y comportamiento de mamiferos en letrinas de ocelotes ( Leopardus pardalis ) en Costa Rica. Documentamos los patrones de uso temporal por parte de especies focales (ocelotes), la diversidad de especies de mamiferos y los comportamientos que ocurrieron en estos lugares. Los resultados indican que el uso por parte de los ocelotes muestran picos de visitas cada 60 a 67 dias y un intervalo mas corto para visitas inter-sexuales que para visitas intra-sexuales. Esto apoya la idea de que las letrinas de los ocelotes son utilizadas para comunicar estatus reproductivos. Un total de catorce mamiferos terrestres visitaron las letrinas y estas especies mostraron una variedad de comportamientos, incluyendo investigacion, marcaje y aceptacion de rastros olfativos. La complejidad y frecuencia de comportamientos por especies no focales, sugieren que las letrinas pueden jugar un rol importante en la comunicacion de otras especies como lo es para los ocelotes.