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WILDLIFE MONITORING REPORT FOR CORCOVADO NATIONAL
PARK, COSTA RICA - 2020
TECHNICAL REPORT – AÑO DE LA PANDEMIA
Erik R Olson1*, Victoria Chevalier1, Joshua Kolasch1, Destiney Elder-Hall1, Alejandro Azofeifa2,
Eloy Olmos2, Danny Herrera2, Stephanie Mory Villaseñor2, Guido Saborío-R3
1 Northland College, 1411 Ellis Avenue, Ashland, WI 54806
2 Área de Conservación Osa, Sistema Nacional de Áreas de Conservación
3 Programa Nacional de Monitoreo Ecológico, Unidad de Seguimiento Estrategia Nacional de
Biodiversidad, Sistema Nacional de Áreas de Conservación
* corresponding author - Northland College, 1411 Ellis Avenue, Ashland, WI 54806, (1)715/682-1235,
eolson@northland.edu
Recommended citation: Olson, E.R., Chevalier, V., Kolasch, J., Elder-Hall, D., Azofeifa, A.,
Olmos, E., Herrera, D., Mory Villaseñor, S., Saborío-R, G. 2021. Wildlife monitoring report for
Corcovado National Park, Costa Rica - 2020. Technical Report. Pp 32. Northland College,
Wisconsin, USA. DOI: 10.13140/RG.2.2.29594.36800
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This research is the result of the JaguarOsa Project, a collaborative wildlife research and conservation
project. JaguarOsa is a partnership between Erik R Olson’s wildlife research lab at Northland College in
Wisconsin, USA and the Área de Conservación Osa of Sistema Nacional de Áreas de Conservación of
Costa Rica. This collaborative work aims to provide information to support and aid in the conservation of
wildlife, especially jaguars, white-lipped peccaries, Baird’s tapirs, primates, and pumas within the
protected areas (e.g., Corcovado National Park, Piedras Blancas National Park) of the Osa peninsula of
Costa Rica.
Esta investigación es el resultado del proyecto JaguarOsa, un proyecto colaborativo de investigación y
consersación de vida silvestre. JaguarOsa es una colaboración entre el laboratorio de investigación en
vida silves de Erik Olson, en Northland College, Wiconsin, EEUU, y el Área de Conservación Osa del
Sistema Nacional de Áreas de Conservación de Costa Rica. Esta colaboración busca proveer información
que apoye y sume en la conservación de la vida silvestre, especialmente para el jaguar, chancho de monte,
danta, pumas y primates dentro de las áreas protegidas (por ejemplo, el Parque Nacional Corcovado y el
Parque Nacional Piedras Blancas) en la Península de Osa.
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This research is dedicated to the memory of Parker J Matzinger.
He was a burst of light.
Parker J Matzinger
March 14, 1994 - January 26, 2017
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Resumen
Las cámaras trampa fueron instaladas entre el 28 de enero y el 15 de agosto del 2020. Se
instalaron y mantuvieron 18 estaciones, y en cinco de esas estaciones las cámaras fueron robadas
o destrozadas. Cada estación tuvo un promedio de 122 noches/trampa por cámara. Detectamos
un mínimo de 54 especies en 1 694 noches/trampa (incluyendo aves, roedores, murciélagos y
lagartijas). El muestreo del 2020 fue iniciado durante una pandemia global. Esto provocó retos
para el monitoreo y la conservación de la vida silvestre de Corcovado sin precedentes. Además,
nos presentó una oportunidad para explorar algunas de las ramificaciones de este cambio tan
drástico en el eco-turismo y actividades ilegales. Se detectaron jaguares en 9 eventos en cámaras
trampa diferentes durante el periodo de muestreo. Detectamos un mínimo de 5 individuos de
jaguar Panthera onca y un jaguar no identificado a nivel de individuo en el 46% de los sitios con
cámaras trampa. Al menos uno de los jaguares es nuevo en el área de estudio, elevando el
número mínimo de jaguares observados en Corcovado desde el 2015 a 10 individuos (8 machos
y 2 hembras, incluyendo un macho relativamente joven). Nuestros resultados indican que aunque
los jaguares podrían haber disminuido levemente desde el 2003, la abundancia de jaguares
parece estar aumentando más recientemente desde al menos el 2013. Interesantemente, Macho
Uno, un macho que ha sido detectado en Corcovado cada año de nuestro estudio desde el 2015 al
2019, y que fue detectado por otro estudio en el 2008, no fue detectado durante el muestreo del
2020 (pero fue detectado recientemente en el muestreo del 2021). Detectamos 4 de los jaguares
machos detectados previamente durante el muestreo del 2020, además de un nuevo macho
(Coco). La falta de detección de hembras no es algo raro, considerando que los jaguares hembras
tienden a evitar los sistemas de senderos y típicamente tiene menores tasas de detección en
monitoreos con cámaras trampa. Nuestros datos sugieren que hay esperanza para los jaguares del
Parque Nacional Corcovado. Detectamos 167 eventos independientes de chanchos de monte
Tayassu pecari en 92% de las estaciones. Nuestros datos indican que la población de chanchos
de monte ha cambiado durante el tiempo y ha podido aumentar en los años recientes. Los pumas
Puma concolor, fueron documentados en 36 eventos independientes en 62% de los sitios y la
abundancia de pumas parece estar estable en el tiempo, pero potencialmente disminuyendo en los
últimos años. Los zainos Pecari tajacu fueron detectados en 46 eventos independientes en 54%
de los sitios y la abundancia de zainos parece haber aumentado desde el 2003, pero disminuido
en los últimos años. Documentamos 109 eventos independientes de danta Tapirus bairdii en las
cámaras trampa, en 85% de los sitios. Durante el tiempo de este proyecto se han detectado un
total de 88 individuos de ocelote (52 detectados solo en una ocasión y 36 que han sido detectados
más de una vez) y al menos 24 individuos de tigrillos (1 tigrillo ha sido detectado más de una
vez, y el resto ha sido detectado solo una vez, los tigrillos fueron detectados por primera vez en
el 2017). Las siguientes especies parecen haber tenido un aumento en su abundancia y
distribución desde el 2003: pava Penelope pupurascens, cabro de monte Mazama americana,
armadillo Dasypus novemcinctus, tepezcuintle Cuniculus paca, tolomuco Eira barbara, y
manigordo Leopardus pardalis. Las siguientes especies parecen haber aumentado su abundancia
y distribución desde el 2003: el pavón Penelope pupurascens, el cabro de monte Mazama
americana, el armadillo de nueve bandas Dasypus novemcinctus, y el tepezcuintle Cuniculus
paca. Las siguientes especies parecen haber disminuido al principio pero luego aumentado
recientemente: el zorrillo hediendo Conepatus semistriatus, el yaguarundi Puma yaguarondi, and
la gallina de monte Tinamus major. Para el muestreo del 2021 quisimos aumentar el esfuerzo de
muestreo y la cobertura en el Parque instalando las cámaras en tres fases y estableciendo
estaciones nuevas. Sin embargo, retos asocidos con el transporte del equipo a Costa Rica desde
Estados Unidos, debido a la pandemia, retrasaron nuestros esfuerzos y los guardapaques en el
campo tuvieron que trabajar con menos cámaras de lo esperado. El robo o daño de las cámaras
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sigue siendo un problema dentro del Parque y representa un gran revés para los esfuerzos de
monitoreo. Como resultado de la pandemia global y las disminuciones asociadas en el
ecoturismo, documentamos una caída sustancial de humanos en el sistema de senderos (¡20
eventos en total!) y hubo un aumento aparente de actividades ilegales a lo largo del perímetro del
parque. El proyecto ha apoyado múltiples esfuerzos educativos y tiene dos publicaciones
científicas revisadas por pares, una tercera en revisión, una cuarta en preparación, seis informes
técnicos, contribuyó a dos estudios colaborativos del SINAC, contribuyó a dos estudios con
organizaciones conservacionistas locales y se presentó siete veces en conferencias profesionales.
El tráfico de drogas, la caza furtiva y otras actividades ilegales continúan amenazando la vida
silvestre de Corcovado y nuestra capacidad para monitorear dicha vida silvestre. En general, el
estado de la vida silvestre de Corcovado sigue siendo esperanzador y los esfuerzos científicos
para monitorear su vida silvestre son sólidos.
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Abstract
Camera traps were installed between 28 January and 15 August 2020. We installed and maintained 18
camera stations, at five of those stations, cameras were stolen or destroyed. Each camera station averaged
approximately 122 trap nights per camera. We detected a minimum of 54 species over 1,694 trap nights
(includes birds; excludes a number of unidentified birds, rodents, bats, and lizards). The 2020 survey was
initiated amidst a global pandemic. This posed unprecedented challenges to the monitoring and
conservation of wildlife within Corcovado. It also provided us an opportunity to explore some of the
ramifications of such a dramatic shift in eco-tourism and illegal activities. Jaguars were documented in 9
separate camera trap events during the survey period. We detected a minimum of 5 individual jaguars
Panthera onca and 1 unidentifiable jaguar at 46% of camera trap sites. At least one of the jaguars was
new to the study area, bringing the total number of jaguars observed in Corcovado since 2015 to a
minimum of 10 (eight males and two females; including one relatively young male). Our results indicate
that although jaguars may have declined slightly since 2003, jaguar abundance appears to be increasing
more recently since at least 2013. Notably, Macho Uno, a male jaguar, that has been detected in
Corcovado every year of our study (2015-2019), as well as, during a 2008 study, was not detected during
the 2020 survey (but has recently been detected in the 2021 survey). We detected four of the previously
detected male jaguars during 2020, as well as, one new male (Coco). The lack of detections of female
jaguars is not that surprising, considering female jaguars tend to avoid trail systems and typically have
lower detection rates for camera trap surveys. Our data suggests that there is hope for the jaguars of
Corcovado National Park. We detected 167 events of white-lipped peccaries Tayassu pecari from 92% of
the camera trap stations. Our data indicates that the white-lipped peccary population has varied over time
and may have increased in recent years. Pumas Puma concolor were documented in 36 events at 62% of
camera trap sites and puma abundance appears to be stable over time, but potentially decreasing in recent
years. Collared peccaries Pecari tajacu were documented in 46 events at 54% of camera trap sites and
collared peccary abundance appears to have increased since 2003 but decreased in recent years. We
documented 109 camera trap events of tapirs Tapirus bairdii at 85% of camera trap sites. Over the span
of this project, we have detected a total of 88 unique individual ocelots (52 captured only once
and 36 that have been recaptured at least once) and at least 24 individual margays (1 margay has
been recaptured, 23 margays were only detected only once; margays were first detected in 2017).
The following species appear to have increased in abundance and distribution since 2003: crested guan
Penelope pupurascens, red brocket deer Mazama americana, nine-banded armadillo Dasypus
novemcinctus, and agouti paca Cuniculus paca. The following species appear to have decreased at first
and then increased more recently: striped hog-nosed skunk Conepatus semistriatus, jaguarundi Puma
yaguarondi, and the great tinamou Tinamus major. For 2021 survey efforts, we attempted to increase our
coverage of the park by installing cameras in three phases and establishing multiple new camera stations.
However, challenges associated with transporting equipment to Costa Rica from the US, due to the
pandemic, slowed our efforts and park rangers in the field worked with fewer cameras than expected.
Theft or damage of cameras continues to be a problem within the park and represents a major setback for
monitoring efforts. As a result of the global pandemic and the associated declines in eco-tourism, we
documented a substantial drop in humans on trail systems (20 events total!) and there was an apparent
increase of illegal activities along the perimeter of park. The project has supported multiple education
efforts and has two peer reviewed scientific publications, a third in review, a fourth in preparation, six
technical reports, contributed to two SINAC collaborative studies, contributed to two studies with local
conservation organizations, and presented seven times at professional conferences. Drug running,
poaching, and other illegal activities continue to threaten the wildlife of Corcovado and our ability to
monitor said wildlife. Overall, the status of Corcovado’s wildlife remains hopeful and the scientific
efforts to monitor it’s wildlife remains strong.
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Figure Captions
Figure 1. Camera station locations within Corcovado National Park for the 2020 (black circles
and X’s), 2019 (blue), 2018 (magenta), and 2017 (gray; gridlines represent 4X4km camera trap
network grid). Land cover classification data courtesy of NASA DEVELOP and Osa
Conservation
Figura 1. Ubicación de las estaciones de cámaras trampa en el Parque Nacional Corcovado para
el 2020 (negro y X’s), 2019 (azul), 2018 (magenta), y 2017 (gris; líneas grises representan la
cuadrícula de 4x4 km). La clasificación de la cobertura de la tierra es cortesía de NASA
DEVELOPMNET y Conservación Osa
Figure 2. Images of the nine jaguars (Panthera onca) a) Macho Uno (m) (2008, 2015, 2016,
2017, 2018, 2019), b) Dia (f) (2016), c) Vivi (f) (2015, 2017), d) Espejo (m) (2016, 2018, 2019,
2020), e) Calvin (m) (2017, 2019, 2020), f) Don Álvaro (m) (2019), g) El Trotomundo (m)
(2016, 2017, 2018, 2019, 2020), h) Champeon (2019), and i) El Hijo (2019, 2020) detected
during the 2015-2020 camera trap surveys in Corcovado National Park. These pictures exclude
one male (Coco, 2020) and multiple undeterminable events of jaguars
Figura 2. Imágenes de los nueves jaguares (Panthera onca) a) Macho Uno (m) (2008, 2015,
2016, 2017, 2018, 2019), b) Dia (f) (2016), c) Vivi (f) (2015, 2017), d) Espejo (m) (2016, 2018,
2019, 2020), e) Calvin (m) (2017, 2019, 2020), f) Don Álvaro (m) (2019), g) El Trotomundo (m)
(2016, 2017, 2018, 2019, 2020), h) Champeon (2019), y i) El Hijo (2019, 2020) detectados
durante el 2015 al 2020 mediante el monitoreo con cámaras trampa en el Parque Nacional
Corcovado. Estas imágenes exluyen un nuevo macho (Coco, 2020)
Figure 3. Macho Uno, a male jaguar (Panthera onca), caught on camera in 2008 (mistakenly
identified as an adult female, though likely a juvenile male at the time; Carazo 2009), 2015
(Olson et al. 2016), 2016 (Olson et al. 2017), 2017 (Olson et al. 2018), 2018 (Olson et al. 2020a),
and 2019 (Olson et al. 2020b) in Corcovado National Park. Olson et al. (2019) recently published
a paper regarding Macho Uno in the journal Cat News (issue 69 spring, pp. 4-6) suggesting that
the resiliency of one of the oldest wild jaguars ever recorded is a sign of hope for the jaguars of
Corcovado National Park. Macho Uno was not detected via this survey in 2020
Figura 3. Macho uno, un jaguar macho (Panthera onca), capturado en cámara en 2008
(identificado erróneamente como una hembra adulta, que era un juvenil macho en esa época,
Carazo 2009), 2015 (Olson et al. 2016), 2016 (Olson et al. 2017), 2017 (Olson et al. 2018), 2018
(Olson et al. 2020), y 2019 en el Parque Nacional Corcovado. Olson et al. (2019) recientemente
publicaron un artículo sobre Macho Uno en la revista científica Cat News (issue 69 spring, pp. 4-
6) sugiriendo que la permanencia de uno de los jaguares más viejos jamás reportados es una seña
de esperanza para los jaguares del Parque Nacional Corcovado
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Figure 4. Jaguar (blue) and white-lipped peccary (orange) abundance (tracks/km) from Carrillo
(2012) and Azofeifa et al. (2021). R2 values are associated with dashed lines representing
estimates of a best fit line for the data. Data indicates that jaguars may have declined in
abundance (tracks/km) from 1990’s through early 2010’s but are now increasing. Data also
indicates that white-lipped peccary abundance is highly variable over time but may be increasing
in recent years
Figura 4. Abundancia (huellas/km) de jaguar (azul) y chancho de monte (anaranjado) de Carrillo
(2012) y Azofeifa et al. (2021). Los valores de R2 están asociados con líneas punteadas
representando la estimación de la línea de mejor ajuste de los datos. Datos indican que la
abundancia de huellas de jaguar puede haber disminuido desde 1990 al principio del 2010, pero
están aumentando ahora. Los datos además indican que la abundancia de chanchos de monte es
muy variable en el tiempo, pero están aumentando ahora
Figure 5. Abundance (RAI) from camera trap data (left; Carazo 2009; Olson et al. 2016, 2017,
2018, 2020, and this report; RAI=Events/1000TN) for jaguar (light blue), white-lipped peccary
(orange), tapir (dark blue), collared peccary (yellow), and puma (grey) (no data for years 2004-
2007 and 2009-2014). Frequency of occurrence from camera trap data (right; Olson et al. 2016,
2017, 2018, 2020, and this report) for jaguars (light blue) and white-lipped peccaries (orange)
with linear trend lines fit to the data. Note that camera trap station locations have changed
somewhat over time due to camera placement, camera malfunction, or theft – this especially
influences tapir detections
Figura 5. Abundancia relativa en base a los datos de cámaras trampa (izquierda; Carazo 2009;
Olson et al. 2016, 2017, 2018, 2020, y este reporte; RAI=Eventos/1000NT) para el jaguar
(celeste), chancho de monte (anaranjado), danta (azul oscuro), zaino (amarillo), y puma (gris)
(sin datos para 2004-2007 y 2009-2014). Frequncia de ocurrencia en base a los datos de cámaras
trampa (derecha; Olson et al. 2016, 2017, 2018, 2020, y este reporte) para el jaguar (celeste) y
chancho de monte (anaranjado) con líneas punteadas uniendo los puntos. Tenga en cuenta que
las ubicaciones de las estaciones de cámaras trampa han cambiado algo con el tiempo debido a la
ubicación de la cámara, el mal funcionamiento de la cámara o el robo; esto influye especialmente
en las detecciones de tapires
Figure 6. Jaguar distribution within 4X4 km grid cells as determined by camera trap or track
survey data collected in Corcovado National Park, Costa Rica from 2013-2020 (areas without
any survey effort are not shaded any color). Camera trap locations for 2020 are shown for
reference (15 of 23 (65%) grid cells have evidence of jaguars)
Figura 6. Distribución del jaguar dentro de la cuadrícula de 4x4 km, determinada mediante las
cámaras trampa y el monitoreo de rastros, en el Parque Nacional Corcovado, Costa Rica del
2013 al 2020 (áreas sin ningún esfuerzo de muestreo no están coloreadas). La ubicación de las
cámaras trampa para el 2020 son mostradas como referencia (15 de 23 (65%) celdas de
cuadrícula tienen evidencia del jaguar
Figure 7. White-lipped peccary distribution within 4X4 grid cells as determined by camera trap
or track survey data collected in Corcovado National Park, Costa Rica from 2013-2020 (areas
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without any survey effort are not shaded any color). Camera trap locations for 2020 are shown
for reference
Figura 7. Distribución del chancho de monte dentro de la cuadrícula de 4x4 km, determinada
mediante las cámaras trampa y el monitoreo de rastros, en el Parque Nacional Corcovado, Costa
Rica del 2013 al 2020 (áreas sin ningún esfuerzo de muestreo no están coloreadas). La ubicación
de las cámaras trampa para el 2020 son mostradas como referencia
Table Captions
Table 1. Number of events and relative abundance (RAI; RAI=E (E/1000TN) of wildlife species
detected on camera trap in Corcovado National Park, Costa Rica and relative trend in RAI
(decreasing=↓, increasing=↑, stable=↔, ?=indicates greater uncertainty) from 2003 to 2020
Cuadro 1. Número de eventos y abundancia relativa (RAI; RAI=E (E/1000TN) de las especies de vida
silvestre detectadas en cámaras trampa en el Parque Nacional Corcovado, Costa Rica y la tendencia
relativa en RAI (disminuyendo=↓, aumentando=↑, estable=↔, ?=indica mayor incertidumbre) desde el
2003 al 2020
Table 2. Frequency of occurrence (FOC; FOC=camera stations present/total number of camera
stations) of wildlife species detected on camera trap in Corcovado National Park, Costa Rica and
relative trend in FOC (decreasing=↓, increasing=↑, stable=↔, ?=uncertainty) from 2003 to 2020
Cuadro 2. Frecuencia de ocurrencia (FOC, FOC = estaciones presente/ número total de estaciones) de las
especies de vida silvestre detectadas en cámaras trampa en el Parque Nacional Corcovado, Costa Rica y la
tendencia relativa en el FOC (disminuyendo=↓, aumentando=↑, estable=↔, ?=incierto) desde el 2003 y
hasta el 2020
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Background
This long-term monitoring effort was designed to monitor trends in wildlife diversity and
abundance within Corcovado National Park. This study was initiated in 2015 (Olson et al. 2016)
and expanded in 2016 (Olson et al. 2017). Special focus was paid to ensure the project was
designed to assess the status of three endangered species, jaguar Panthera onca, white-lipped
peccaries Tayassu pecari, and Baird’s tapir Tapirus bairdii within Corcovado National Park. In
2015 and 2016, we surveyed the park using camera traps spaced approximately 1.6 to 3.4 km
apart. However, in 2017 we implemented camera trapping efforts according to a 4X4 km grid
overlaid across the entire the Osa Peninsula to support national wildlife monitoring. This
transition resulted in the elimination of a few camera trap stations within the bajura area of the
park (i.e., areas near beaches and lowlands around the Corcovado Lagoon) and allowed us to
expand our monitoring efforts into the more mountainous regions of the park (Figure 1). Despite
these changes, we retained most of our historical camera trapping stations. In 2020, we installed
cameras according to the same protocols implemented in 2017-2019. In 2020, our survey
encompassed around ~37% of Corcovado National Park (Figure 1). We utilized tourist and
research trails for camera placement. The study area was ~160km2 in 2020 (2015 ~150 km2;
2016 ~170 km2; 2017 ~240 km2; 2018 ~170km2; 2019 ~170km2). Historically, our surveys run
February through June – spanning a minimum of three months. However, in 2019 we decided to
retain cameras in the field all year to better understand the seasonal variation in wildlife
abundance and activity. In 2020, we returned to our traditional monitoring window, with cameras
active from the end of January to mid-August. Our monitoring effort represents some of the most
intensive (spatially and temporally) wildlife monitoring done in the park to date (i.e., longest
consecutive time maintaining cameras throughout the greatest portion of the park).
Methods
We used non-invasive monitoring techniques (i.e., camera traps) to monitor the wildlife of
Corcovado National Park (Long et al. 2008). We identified thirty camera stations within a 4X4
km grid to install cameras (Figure 1). We attempted to maintain even spacing between cameras
and only one camera per grid cell. However, we used historical camera station locations within
grid cells where possible to maintain consistency over time (Harmsen et al. 2017). We placed
cameras, locally, in areas with the highest probability of capturing wildlife (i.e., along trails, near
river crossings, mud pits, ridgelines) and programmed camera traps to record the time, date,
temperature, and moon phase for each photo. We set cameras to take bursts of three photographs
with less than one second between each photograph and a no refractory period (i.e., rapid-fire)
between events (Apps & McNutt 2018). To avoid pseudo-replication, we defined an event as any
photo-series of a species and considered subsequent photos of the same species within 30
minutes of a previous photo to be the same event (O’Brien et al., 2003; Naing et al. 2015; Allen
et al. 2019). We calculated the relative abundance index (RAI) as: RAI = (E/TN) * 1000, where E
is the number of events and TN is the total number of trap nights (Allen et al. 2019) We used RAI
because it is considered an accurate index of abundance (Parsons et al. 2017; Palmer et al. 2018)
or site use (Sollmann 2018; though see Stewart et al. 2018). For individual identification of
jaguars, E.R. Olson identified individuals based on spot patterns and other physical features, and
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determined sex based on the presence of male genitalia (Harmsen et al. 2017). For individual
identification of ocelots, V. Chevalier and E.R. Olson identified individuals using rosette spot
patterns using program Hotspotter and expert validation.
Figure 1. Camera station locations within Corcovado National Park for the 2020 (black circles
and X’s), 2019 (blue), 2018 (magenta), and 2017 (gray; gridlines represent 4X4km camera trap
network grid). Land cover classification data courtesy of NASA DEVELOP and Osa
Conservation.
Results & Discussion
Eighteen camera trap stations were installed and maintained between 28 January and 15 August
2020. Cameras were destroyed or stolen at five of the camera stations, resulting in 13 functional
camera stations. Of the functioning camera stations, there was an average of 122 trap nights per
station. We detected a minimum of 54 species over 1,694 trap nights (includes birds; excludes a
number of unidentified birds, rodents, bats, and lizards), including a minimum of 5 jaguars Panthera
onca and at least one new jaguar for Corcovado.
Jaguars in 2020
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Jaguars were documented in 9 separate camera trap events during the survey period (Table 1).
One jaguar could not be identified to the individual-level. The 2020 study identified a minimum
of 5 individual jaguars (Figure 2) and one unidentifiable jaguar at 6 camera trap stations during
the survey period (46% of camera trap stations). We were able to identify 5 jaguars resulting in 5
males. Four of the previously detected males were re-detected during the 2020 survey effort (El
Trotamundo, Espejo, Calvin, El Hijo). One new male, Coco, was detected in a grid cell with no
jaguar events to date. Macho Uno was not detected in 2020, but was previously detected in every
year of our study since 2015 and prior study in 2008 (Carazo 2009; Olson et al. 2016; Olson et
al. 2017; Olson et al. 2018; Olson et al. 2019a; Olson et al. 2019b; Olson et al. 2020a; Olson et
al. 2020b; Figures 2 and 3). El Trotamundo, was detected in 2016, 2017, 2018, and 2019 (Olson
et al. 2017; Olson et al. 2018; Olson et al. 2020a; Olson et al. 2020b; Macho2 in 2016; Figure 2).
Espejo was detected previously in 2018 and 2019 and just outside the park in 2016 (Olson et al.
2020a; Olson et al. 2020b; Figure 2). Calvin was detected previously in 2017 and 2019 (Olson et
al. 2018; Olson et al. 2020b; Figure 2). El Hijo was detected in 2019 for the first time (Olson et
al. 2020b; Figure 2). In 2020, El Hijo was observed three times, El Trotamundo observed twice,
Espejo observed once, Calvin observed once, and Coco observed once. However, there was one
event of an unidentifiable individual and 3 events of the same individual that could be any one of
two individuals (due to incomplete jaguar profiles) or a new individual.
The lack of detection of females in 2020 is not surprising. Females tend to avoid trail systems
and use the landscape differently than do males. In fact, most camera trap studies tend to exhibit
male biased detection rates (Harmsen et al. 2017). Thus, it is expected that females are present
within Corcovado National Park but have lower detection rates. Additionally, Dia was detected
in an area where we have not been able to install camera traps since 2016 due to safety concerns
– thus, our ability to detect Dia may be in part due to our inability to survey that area.
In this study, El Trotomundo was observed at Las Latas and OWA. He was observed at Las
Latas in 2017 (Olson et al. 2018), at Bajura Rio Corcovado in 2018 and 2019 (Olson et al.
2020a; Olson et al. 2020b), and along the Playa Sirena in 2016 (Unknown2 in 2016; Olson et al.
2017).
Espejo, another large male, was observed at Ollas. Espejo has been observed at Las Latas, Bajura
Llorito, Ollas, and Bajura Rio Corcovado in 2019 (Olson et al. 2020b), and was observed at Las
Latas, Bajura Llorito, Sector Ticho, and Bajura Rio Corcovado in 2018 (Olson et al. 2020a).
Espejo appears to be covering a relatively large area of the park.
We did not generate density estimates for our study. Density estimates from camera trap surveys
have come under increased scrutiny recently (Maffei et al. 2011; Tobler & Powell 2013), and
while they are still valuable, experts have suggested reporting minimum known alive by trap
nights and study area. We plan on eventually examining changes in jaguar occupancy and
detection probability over time within our study area and hopefully to estimate density of jaguars
using methods similar to Harmsen et al.’s (2017) long-term study.
Between 2015 and 2020, we have detected a minimum of 10 individual jaguars; 8 males (Macho
Uno, Calvin, El Trotomundo, Espejo, Champion, El Hijo, Don Álvaro, Coco), 2 females (Vivi,
Page 13 of 32
Dia), and 7 unknowns. However, it is possible that some of the unknown jaguars are also one of
the known jaguars. Additionally, it is important to note that our survey area does not cover all of
Corcovado and that Corcovado could contain both resident and dispersing jaguars. It is also
important to note that we do not have complete profiles for Don Álvaro and Coco yet. Thus, we
may be able to retroactively assign identity to some unknown observations of jaguars once we
confirm a good left profile image for these individuals.
Based on our monitoring history of jaguars within the park, it appears as though the following
male jaguars are more likely residents of Corcovado: Macho Uno, Espejo, El Trotomundo,
Calvin, and El Hijo. The other individuals have only more recently been detected and therefore
status assignment is going to be less reliable. The two female jaguars detected (Dia, Vivi) are
most likely also resident jaguars of Corcovado, especially Vivi, but low detection probability of
females makes status assignment less certain.
It should be noted that two separate jaguars were captured on camera trap in 2016 in the Golfo
Dulce Forest Reserve, just outside of Corcovado National Park by MINAE SINAC-ACOSA
cameras. One of the jaguars captured is Espejo, indicating that Espejo has been using the area
around the park since at least 2016. The other jaguar appears to be a different individual from
those that we have captured in Corcovado National Park between 2015 and 2020 but was
recaptured in the Gulfo Dulce Forest Reserve in 2019 (Bone-Guzmán and Chacón-Madrigal
2020), suggesting that it is a resident of the forest reserve adjacent to the park. A male jaguar was
also captured on camera in Piro at Osa Conservation by Osa Conservation (A. Whitworth, pers.
comm.) and that jaguar does not a match with any of the jaguars detected thus far.
Status of the Jaguar
Salom-Perez et al. (2007) documented a minimum of 4 individual jaguars (3 males and 1 female)
during the course of their study (363 TN), but observed a total of 7 jaguars after they included
other opportunistic camera trap observations thereafter (4 males and 3 females). They
documented 4 jaguars within their estimated 86 km2 study area located in the bajura east of the
lagoon. They calculated a density of 6.98 ± SD 2.36 per 100 km2 for their study area, which
covered 20% of the park.
Carazo (2009) documented a minimum of 5 individual jaguars (3 males, 2 females, and 1
unknown) during the course of their study (668 TN). They documented 4 jaguars within their
roughly 40 km2 study area located in the bajura east of the lagoon. Their study area covered
roughly 10% of the park.
In 2015, Olson et al. (2016) documented a minimum of 2 individual jaguars (1 male and 1
female) during the course of their study (592 TN) but observed a total of 3 jaguars after they
included other opportunistic observations thereafter (1 male, 1 female, and 1 unkown). They
documented 2 jaguars within their estimated 153 km2 study area covering the extent of the
bajura. Their study area covered ~33% of the park.
Page 14 of 32
Table 1. Number of events and relative abundance (RAI; RAI=E (E/1000TN) of wildlife species detected on camera trap in Corcovado National
Park, Costa and relative trend in RAI (decreasing=↓, increasing=↑, stable=↔, ?=indicates greater uncertainty) from 2003 to 2020
Scientific Name
2003*
RAI
2008*
RAI
2015*
RAI
2016*
RAI
2017*
RAI
2018*
RAI
2019*
RAI
2020*
RAI
Trend
Ateles geoffroyi
**
**
1 (1.7)
___
___
___
1(0.3)
2 (1.2)
___
Conepatus semistriatus
9 (4.0)
3 (4.5)
___
___
1 (2.4)
6 (6.8)
17 (5.6)
24 (14.2)
↓↑
Cuniculus paca
40 (17.8)
17 (25.4)
6 (10.1)
14 (17.7)
25 (59.7)
59 (66.9)
314 (95.3)
137 (80.9)
↑
Crax rubra
277 (123.4)
59 (88.3)
95 (160.5)
110 (138.7)
38 (90.7)
162 (183.7)
304 (92.2)
202 (119.2)
↔
Dasyprocta punctata
514 (229.1)
151 (226.0)
221 (373.3)
168 (211.9)
354 (844.9)
455 (546.2)1
1914 (580.7)
762 (449.8)
↔
Dasypus novemcinctus
1 (0.4)
**
___
1 (1.3)
3 (7.2)
20 (22.7)
45 (13.7)
12 (7.1)
↑
Didelphis marsupialis
16 (7.1)
24 (35.9)
15 (25.3)
18 (22.7)
7 (16.7)
60 (68.0)
214 (64.9)
47 (27.7)
***
Eira barbara
20 (8.9)
5 (7.5)
5 (8.5)
3 (3.8)
4 (9.5)
12 (13.6)
34 (10.3)
12 (7.1)
↔
Leopardus pardalis
75 (33.4)
37 (55.4)
33 (55.7)
27 (34.0)
12 (28.6)
18 (20.4)
95 (28.8)
29 (17.1)
↑↓?
Leopardus wiedii
2 (0.9)
4 (6.0)
___
___
___
5 (5.7)
16 (4.9)
14 (8.3)
↓↑?
Mazama americana
36 (16.0)
8 (12.0)
13 (22.0)
12 (15.1)
82 (195.7)
37 (42.0)
46 (14.0)
76 (44.9)
↑
Nasua narica
30 (13.4)
7 (10.5)
6 (10.1)
10 (12.6)
4 (9.6)
29 (32.9)
76 (23.1)
79 (46.6)
↑?
Panthera onca
24 (10.7)
5 (7.5)
3 (5.1)
6 (7.6)
8 (19.1)
12 (13.6)
39 (11.8)
9 (5.3)
↔?
Pecari tajacu
29 (12.9)
8 (12.0)
11 (18.6)
25 (31.5)
27 (64.4)
33 (37.4)
168 (51.0)
46 (27.2)
↑
Penelope purpurascens
1 (0.4)
4 (6.0)
___
___
1 (2.4)
6 (6.8)
8 (2.4)
9 (5.3)
↓↑
Procyon spp.
3 (1.3)
**
15 (25.3)
4 (5.0)
3 (7.2)
9 (10.2)
18 (5.5)
5 (3.0)
?
Puma concolor
58 (25.8)
15 (22.5)
31 (52.4)
23 (29.0)
9 (21.5)
29 (32.8)
54 (16.4)
36 (21.3)
↔
Puma yaguarondi
4 (1.8)
2 (3.0)
___
___
___
___
7 (2.1)
5 (3.0)
↓↑
Sciurus spp.
**
**
___
___
1 (2.4)
8 (9.1)
59 (18.0)
27 (15.9)
↑?
Tamandua mexicana
7 (3.1)
**
___
1 (1.3)
___
2 (2.3)
16 (4.9)
9 (5.3)
↑?
Tapirus bairdii
111 (49.5)
47 (70.4)
122 (206.1)
102 (128.6)
39 (93.1)
43 (48.3) 1
124 (37.6)
109 (64.3)
↔
Tayassu pecari
113 (50.4)
55 (82.3)
94 (158.8)
41 (51.7)
31 (74.0)
107 (121.3)
264 (80.1)
167 (98.6)
↔↑
Tinamus major
151 (67.3)
16 (24.0)
6 (10.1)
2 (2.5)
4 (9.5)
68 (77.1)
201 (61.0)
68 (40.1)
↓↑
*2003 (2244 TN, 13 camera stations) & 2008 (668 TN, 28 camera stations) data from Carazo 2009, data adjusted to Events/1000TN; 2015 data from Olson et al.
2016 (592 TN, 21 camera stations); 2016 from Olson et al. 2017 (793 TN, 21 camera stations); 2017 from Olson et al. 2018 (419 TN, 17 camera stations); 2018
from Olson et al. 2020a (882 TN, 17 camera stations1); 2019 from Olson et al. 2020b (3296 TN, 14 camera stations); 2020 from this report (1964 TN, 13 camera
stations); **Unreported; *** this data likely represents more than one species of possum; 1 RAI calculated based on data from an additional camera station in
2018 (camera at site was only functioning for 8 days; 890 TN total).
Page 15 of 32
Table 2. Frequency of occurrence (FOC; FOC=camera stations present/total number of camera stations)
of wildlife species detected on camera trap in Corcovado National Park, Costa and relative trend in FOC
(decreasing=↓, increasing=↑, stable=↔, ?=uncertainty) from 2015 to 2020
Scientific Name
2015*
FOC (#)
2016*
FOC (#)
2017*
FOC (#)
2018*
FOC (#)
2019*
FOC (#)
2020*
FOC (#)
Trend
Ateles geoffroyi
5% (1)
___
___
___
7% (1)
15% (2)
?
Conepatus
semistriatus
___
___
7% (1)
11% (2)
50% (7)
38% (5)
↑
Cuniculus paca
24% (5)
24% (5)
38% (6)
47% (8)
86% (12)
62% (8)
↑
Crax rubra
76% (16)
71% (15)
50% (8)
94% (16)
100%
(14)
100%
(13)
↔
Dasyprocta
punctata
91% (19)
71% (15)
75% (12)
100%
(17)1
100%
(14)
100%
(13)
↔
Dasypus
novemcinctus
___
5% (1)
13% (2)
24% (4)
57% (8)
38% (5)
↑
Didelphis
marsupialis
29% (6)
19% (6)
19% (3)
53% (9)
86% (12)
62% (8)
↑**
Eira barbara
14% (3)
19% (2)
19% (3)
35% (6)
79% (11)
54% (7)
↑
Leopardus
pardalis
62% (13)
29% (6)
50% (8)
65% (11)
100%
(14)
77% (10)
↔
Leopardus wiedii
___
___
___
18% (3)
50% (7)
38% (5)
↑?
Mazama
americana
29% (6)
29% (6)
38% (6)
47% (8)
71% (10)
62% (8)
↑
Nasua narica
19% (4)
19% (4)
25% (4)
65% (11)
100%
(14)
69% (9)
↑
Panthera onca
10% (2)
19% (4)
31% (5)
24% (4)
79% (11)
46% (6)
↑
Pecari tajacu
29% (6)
52% (11)
31% (5)
59% (10)
86% (12)
54% (7)
↔
Penelope
purpurascens
___
___
6% (1)
12% (2)
43% (6)
38% (5)
↑
Procyon spp.
24% (5)
14% (3)
6% (1)
18% (3)
57% (8)
8% (1)
↔
Puma concolor
52% (11)
52% (11)
38% (6)
47% (8)
79% (11)
62% (8)
↔
Puma yaguarondi
___
___
___
___
29% (4)
31% (4)
↑
Sciurus spp.
___
___
6% (1)
24% (4)
57% (8)
23% (3)
↑
Tamandua
mexicana
___
5% (1)
___
6% (1)
50% (7)
46% (6)
↑
Tapirus bairdii
86% (18)
62% (13)
50% (8)
61% (11) 1
93% (13)
85% (11)
↔
Tayassu pecari
52% (11)
43% (9)
31% (5)
65% (11)
86% (12)
92% (12)
↑?
Tinamus major
19% (4)
5% (1)
13% (2)
59% (10)
100%
(14)
62% (8)
↑
*2003 (2244 TN, 13 camera stations) & 2008 (668 TN, 28 camera stations) data from Carazo 2009, data adjusted to
Events/1000TN; 2015 data from Olson et al. 2016 (592 TN, 21 camera stations); 2016 from Olson et al. 2017 (793
TN, 21 camera stations); 2017 from Olson et al. 2018 (419 TN, 17 camera stations); 2018 from Olson et al. 2020a
(882 TN, 17 camera stations1); 2019 from Olson et al. 2020b (3296 TN, 14 camera stations); 2020 from this report
(1964 TN, 13 camera stations); **Unreported; *** this data likely represents more than one species of possum; 1
RAI calculated based on data from an additional camera station in 2018 (camera at site was only functioning for 8
days; 890 TN total).
Page 16 of 32
In 2016, Olson et al. (2017) examined jaguar abundance within Corcovado National park using a
two-stage camera trap grid in 2016. Olson et al. (2017) documented a minimum of 3 individual
jaguars (2 male and 1 female) during the course of their study (793 TN). They documented 3
jaguars within their estimated ~170 km2 study area covering the extent of the bajura. Their study
area covered ~40% of the park.
In 2017, Olson et al. (2018) examined jaguar abundance within Corcovado National park using a
two-stage camera trap grid in 2017. The area covered by that study was expanded, but camera
trap density was reduced, in part due to a large number of camera trap stations that were
compromised for a variety of reasons. Olson et al. (2018) documented a minimum of 5
individual jaguars (3 male, 1 female, 1 unknown) during the course of their study (419 TN).
They documented 5 jaguars within their estimated ~240 km2 study area covering the extent of a
wide range of habitat types within the park (but with significantly reduced camera trap station
density). Their study area covered ~56% of the park.
In 2018, Olson et al. (2020a) examined jaguar abundance within Corcovado National park using
a two-stage camera trap grid in 2018. There were fewer camera trap thefts or damages due to the
use of security boxes and more dedicated wildlife monitoring field staff – resulting in a survey
effort (trap nights) 2.1 times that of 2017. Olson et al. (2020a) documented a minimum of 3
individual jaguars (3 males, 1 unknown) during the course of their study (882 TN). The study
area was like that of the 2016 survey effort (~170 km2). They documented a minimum of 3
jaguars within their estimated ~170 km2 study area covering the extent of a wide range of habitat
types within the park. Their study area covered ~40% of the park.
With 3,296 trap nights, the 2019 monitoring effort represented the most intensive camera trap survey
effort ever implemented in Corcovado National Park to date (roughly 5.5 times greater effort than most
previous studies). Olson et al. (2020b) documented a minimum of 7 individual jaguars (7 male, 1
unknown) during the course of this study (3,296 TN). Their estimated study area covered ~170
km2 representing ~40% of the park and a wide range of habitat types (Olson et al. 2020b). This
study represents the most intensive spatial and temporal coverage for a camera trap study in the
park to date.
For 2020, we detected a minimum of 5 individual jaguars. The increased temporal coverage for
some cameras likely led to a relatively high survey effort for 2020 (1,694 TN for 13 stations).
The study area was like that of 2018 and 2019, with a study area of ~160 km2 representing ~37%
of the park – though theft or damage of 5 camera stations resulted in poorer coverage of some
areas of the study area.
Assessment - Our research indicates that trends in jaguar abundance are not as clear as previously
thought. It is likely that jaguar abundance has declined in Corcovado since 1990, however, the
extent of that decline is not clear. Jaguar abundance appears to have fluctuated since 2003 but
remained relatively stable, if not increased. There are some indications that jaguar abundance
may have increased since 2013 (Azofeifa et al. 2019). It is important to acknowledge that this
data comes from a variety of sources and that track survey data can be a questionable approach
for assessing trends in population abundance. Regardless, we believe the consistent nature of
these patterns across data sources are reflective, to some degree, of the actual trends in jaguar
Page 17 of 32
abundance over time, and the apparent recent increases in abundance since 2013 give hope for
the status of jaguars in Corcovado. The regular detection of individual jaguars, the potential
evidence of recent breeding (El Hijo), and the consistency of space use by resident jaguars are
also good signs for the jaguar population of Corcovado. Consistent presence of illegal gold
miners, poachers, and drug runners undermines the ability of ACOSA-SINAC to support
biodiversity conservation and ecotourism within the park. Efforts to reduce illegal activities in
the park should be maintained or increased. We base this overall assessment on the following
information:
1. Salom-Perez et al. (2007) documented a minimum known alive of 4 individuals for an 86
km2 study area with 363 trap nights in 2003. We documented a minimum known alive of
2 individuals in 2015 for an area nearly twice the size (153 km2) and nearly double the
number of trap nights (592 TN; Olson et al. 2016). In 2016, we documented a minimum
known alive of 3 individuals in 2016 for a ~170 km2 area over 793 trap nights (Olson et
al. 2017). For 2017, despite a reduction in camera stations, trap nights, and study area, 4
individuals were confirmed (study area of 240 km2 over 419 trap nights; Olson et al.
2018). For 2018, with a relatively low camera station density, Olson et al. (2020a)
observed 3 individuals and one unidentifiable individual. For 2019, with a relatively low
camera station density, but with a much greater survey effort, Olson et al. (2020b)
observed 7 individuals and 2 unidentifiable individuals. For 2020, with an intermediate
survey effort, in terms of TN’s, we observed 5 individuals and 1 unidentifiable
individual. This data suggests that jaguar abundance may have decreased slightly since
2003, and may be increasing in more recent years.
2. Carrillo et al. (2000) documented between roughly 0.05 and 0.15 jaguar tracks per km in
Corcovado National Park between 1990, 1992, and 1994. Saborío et al. (2021)
documented a mean of 0 (range=0-0), 0.035 (range=0-0.051), 0.0235 (range=0-0.103),
0.0210 (range=0-0.077), 0.0383 (range=0.029-0.061), 0.021 (range=0-0.10), 0.044
(range=0-0.061), and 0.039 (range=0-0.061), jaguar tracks per km calculated from
monthly track surveys throughout 2013, 2014, 2015, 2016, 2017, 2018, 2019, and 2020,
respectively (Figure 4). However, some researchers have highlighted issues associated
with track surveys as a method to assess jaguar abundance (de Angelo 2010; Harmsen et
al. 2010; Polisar et al. 2014). Carrillo et al. (2000) surveyed tracks in slightly different
areas with less temporal intensity than Azofeifa et al. (2021). Thus, as demonstrated by
the variation (see range for jaguar tracks per km based on monthly track surveys), we
would expect high variation in this type of data depending on the spatial and temporal
intensity of the survey. Monthly indices of track data from 2015 through 2020 suggest
strong seasonal variation in track abundance for both jaguars and white-lipped peccaries
(Azofeifa et al. 2021). Track data indicates that jaguar abundance is relatively low but
generally increasing from 2013-2019 (Azofeifa et al. 2019). This data suggests that
jaguars have likely declined since the 1990’s but appear to be increasing in abundance
over at least the last five years.
3. Based on the data presented in Carazo (2009) and Olson et al. (2016, 2017, 2018, 2020a,
2020b) we were able to compare the relative abundance (RAI, Carazo 2009) of jaguars
over time from 2003, 2008, 2015, 2016, 2017, 2018, 2019, and 2020 (Table 1). Over that
Page 18 of 32
time period jaguar abundance appeared to fluctuate, but showed no clear pattern
(2003=10.7, 2008=7.5, 2015=5.1, 2016=7.6, 2017=19.1, 2018=13.6, 2019=11.8,
2020=5.3; Table 1; Figure 5). It also indicates that jaguar abundance may be increasing in
more recent years (i.e., 2015-2019). The highest RAI was reported during the 2017 survey
period and the second highest was reported during the 2018 survey period. The third
highest was reported in 2019. In 2003, Salom-Perez et al. (2007) recorded the fourth
highest abundance, and their survey included camera traps sites in areas with nesting sea
turtles (Salom-Perez 2005; Salom-Perez et al. 2007). Jaguars in Costa Rica are well-
known for predating on sea turtles during the sea turtle nesting season (Salom-Perez
2005; Verissimo et al. 2012), which could have led to an inflated 2003 abundance index.
Jaguars were detected at 79% of camera trap sites in 2019, representing the highest
frequencies of occurrence (FOC or proportion of camera trap sites with presence)
reported from 2015-2020 (Table 2; Figure 6). However, survey effort and spatial
coverage are important factors to consider when comparing any of these results. More
recent surveys have covered larger areas and longer periods of time but were less
concentrated on detecting jaguars near beaches during sea turtle nesting season.
Regardless, this data suggests that jaguar abundance has been relatively stable since 2003
and that it may have increased in recent years.
4. We have captured one male, Macho Uno, every year, except for 2020, since 2015 – the
same male detected by Carazo (2009) in 2008. This individual has been living in the
same area for >13 years. Macho Uno’s resident status, small core home range area, and
relatively short distance movements (3.5-6km) are good indicators that Macho Uno likely
occupies high quality habitat (Olson et al., 2019a). However, more recently Macho Uno
has been detected outside of the park in 2019 (Bone-Guzmán and Chacón-Madrigal
2020). These observations may suggest that Macho Uno is being displaced by other
males as he ages. Two other individuals, Espejo and El Trotomundo were observed alive
for >5 yrs, Calvin observed to be alive for >4 yrs, and Vivi observed to be alive for >3
yrs. These potentially resident individuals are indicative of high-quality habitat and some
degree of stability (i.e., individual turnover doesn’t occur on an annual basis).
5. A minimum of eight males and two females were detected between 2015 and 2020
(excluding >7 unidentifiable jaguars), indicating that there is potential reproductive
capacity for the jaguars of Corcovado. Additionally, the detection of El Hijo, suggests
that jaguars within the Greater Corcovado Ecosystem are reproducing. El Hijo appears to
be one of the youngest jaguars detected between 2015 and 2020, likely ~2 yrs-of-age. We
also believe that Calvin may have been a relatively young jaguar when first detected (~2
to 3 yrs-of-age), further suggesting reproductive success for jaguars of the Greater
Corcovado Ecosystem.
Page 19 of 32
Figure 2. Images of seven of the nine jaguars (Panthera onca) a) Macho Uno (m) (2008, 2015, 2016, 2017, 2018, 2019), b) Dia (f)
(2016), c) Vivi (f) (2015, 2017), d) Espejo (m) (2016, 2018, 2019, 2020), e) Calvin (m) (2017, 2019, 2020), f) Don Álvaro (m) (2019),
g) El Trotomundo (m) (2016, 2017, 2018, 2019, 2020), h) Champeon (2019), and i) El Hijo (2019, 2020) detected during the 2015-
2020 camera trap surveys in Corcovado National Park. These pictures exclude one male (Coco, 2020) and multiple undeterminable
events of jaguars.
Page 20 of 32
Figure 3. Macho Uno, a male jaguar (Panthera onca), caught on camera in 2008 (mistakenly
identified as an adult female, though likely a juvenile male at the time; Carazo 2009), 2015
(Olson et al. 2016), 2016 (Olson et al. 2017), 2017 (Olson et al. 2018), 2018 (Olson et al. 2020a),
and 2019 (Olson et al. 2020b) in Corcovado National Park. Olson et al. (2019) recently published
a paper regarding Macho Uno in the journal Cat News (issue 69 spring, pp. 4-6) suggesting that
the resiliency of one of the oldest wild jaguars ever recorded is a sign of hope for the jaguars of
Corcovado National Park. Macho Uno was not detected via this survey in 2020.
2008
2015
2008
2016
2008
2017
2018
2019
Page 21 of 32
Figure 4. Jaguar (blue) and white-lipped peccary (orange) abundance (tracks/km) from Carrillo
(2012) and Azofeifa et al. (2021). R2 values are associated with dashed lines representing
estimates of a best fit line for the data. Data indicates that jaguars may have declined in
abundance (tracks/km) from 1990’s through early 2010’s but are now increasing. Data also
indicates that white-lipped peccary abundance is highly variable over time, but may be
increasing in recent years.
Figure 5. Abundance (RAI) from camera trap data (left; Carazo 2009; Olson et al. 2016, 2017,
2018, 2020, and this report; RAI=Events/1000TN) for jaguar (light blue), white-lipped peccary
(orange), tapir (dark blue), collared peccary (yellow), and puma (grey) (no data for years 2004-
2007 and 2009-2014). Frequency of occurrence from camera trap data (right; Olson et al. 2016,
2017, 2018, 2020, and this report) for jaguars (light blue) and white-lipped peccaries (orange)
with linear trend lines fit to the data. Note that camera trap station locations have changed
somewhat over time due to camera placement, camera malfunction, or theft – this especially
influences tapir detections.
R² = 0.7324
R² = 0.8951
R² = 0.3002
R² = 0.1086
0
0.02
0.04
0.06
0.08
0.1
0.12
0.14
0.16
0.18
0.2
1985 1990 1995 2000 2005 2010 2015 2020
Tracks/km
0
50
100
150
200
250
2003 2008 2013 2018
RAI (events/1000TN)
jaguar wlp puma
clp tapir
R² = 0.5789
R² = 0.6479
0
20
40
60
80
100
2015 2016 2017 2018 2019 2020
Frequency of occurence
jaguar wlp
Linear (jaguar) Linear (wlp)
Page 22 of 32
Figure 6. Jaguar distribution within 4X4 km grid cells as determined by camera trap or track survey data collected in Corcovado
National Park, Costa Rica from 2013-2020 (areas without any survey effort are not shaded any color). Camera trap locations for 2020
are shown for reference (15 of 23 (65%) grid cells have evidence of jaguars)
Page 23 of 32
Status of the White-lipped Peccary
Our research indicates that trends in white-lipped peccary abundance varies with time. It appears
as though white-lipped peccary abundance fluctuates over multi-year periods of time. Based on
Carrillo’s track data white-lipped peccary abundance appears to have declined from 1990 to
2004 and then increased back to previous abundance levels by 2007. Abundance appears to have
fluctuated since 2007, but generally remained relatively high since at least 2013. 2020 appears to
have been another good year for white-lipped peccaries within Corcovado as all metrics of
abundance are relatively high for both track and camera trap surveys (e.g., Table 2, Figure 4, part
2 below). The co-occurrence of white-lipped peccaries and jaguars within the park is striking and
suggests a strong association between the two species (Olson et al. 2018). Consistent presence of
illegal gold miners, poachers, and drug runners undermines the ability of ACOSA-SINAC to
support biodiversity conservation and ecotourism within the park. Efforts to reduce illegal
activities in the park should be maintained and enhanced. We base this assessment on the
following information:
1. Based on RAI from camera trap data, white-lipped peccary abundance appears to have
increased since 2003 (Table 1). RAI for white-lipped peccaries was 50.4, 82.3, 158.8,
51.7, 74.0, 121.3, 80.1, and 98.6 for 2003, 2008, 2015, 2016, 2017, 2018, 2019, and
2020, respectively (Figure 5). Frequency of camera stations with white-lipped peccaries
present appeared to remain relatively stable between 2015 and 2017 and has increased
between 2018 and 2020 – especially if we consider the fact that a number of the new sites
established in 2017 did not detect white-lipped peccaries and that many of our historical
camera stations (some of which had white-lipped peccaries present) did not collect data
due to the reasons described earlier (damage, theft, opportunity costs). However, as
mentioned above, 2019 saw an extension of the survey period length, which would
increase the proportion of camera trap sites with white-lipped peccaries detected due to
greater survey effort at each site. Yet, we detected an even greater increase in abundance
in 2020, despite a reduced survey effort. Regardless, this data suggests that white-lipped
peccary abundance has likely fluctuated annually since 2003 and may have increased
more recently.
2. Since 2017, we have changed our data management technique for white-lipped peccaries.
For 2019 and 2020, we used the maximum number of individuals of any age-class
reported to estimate the minimum number of individuals of each age-class after
attempting to identify mega-herds that used more than one camera trap station. Thus, our
2019 and 2020 estimates are not perfectly comparable to those of 2015-2017 (Olson et al.
2018) – but are worth comparing as the methods are relatively similar. We detected
between 4 and 7 mega-herds at our stations (4, 6, 5, and 6-9 in 2015, 2016, 2017, [no
estimates were made for 2018] and 2019, respectively) and between 258 and 384
individuals which is slightly higher than previous estimates, but slightly lower than the
2019 estimate (204-312, 144-267, 158-243, and 429-581 individuals for 2015, 2016,
2017, [no estimates were made for 2018] and 2019, respectively) (Olson et al. 2018,
2020). In 2019, two relatively independent sites alone had estimates of over 100
individuals each. In 2020, at least two relatively independent sites had estimates of over
60 individuals each. All mega-herds had evidence of recent reproduction (juveniles or
Page 24 of 32
neonates). For 2020, we observed between 32 and 48 juveniles (31-63, 17-29, 24-33, and
78-113 in 2015, 2016, 2017, [no estimate was made in 2018] and 2019, respectively) and
between 38 and 41 neonates (12-31, 21-33, 22-24, and 45-59 in 2015, 2016, 2017, [no
estimate was made in 2018] and 2019, respectively) based on the maximum number of
each age class detected in any event for the 5 or 7 mega-herds detected in 2020 (i.e.,
conservative estimate). Our estimates for 2019 indicated that white-lipped peccaries did
very good that year. This data also tends to track well with white-lipped peccary
abundance (RAI) and occurrence (FOC) at camera trap stations, with lowest abundance
and occurrence occurring in 2016 and 2017. This suggests, at the very least, that the
species is not in immediate threat of extirpation within the park.
3. Carrillo et al. (2000) documented between roughly 0.11 and 0.19 white-lipped peccary
tracks per km in Corcovado National Park between 1990, 1992, and 1994. Olson et al.
(2016) documented 0.1 white-lipped peccary tracks per km in 2015. However, their track
surveys were more opportunistic. Azofeifa et al. (2021) documented a mean of 0.10
(range=0-0.231), 0.12 (range=0-0.308), 0.083 (range=0.051-0.205), 0.09 (range=0-
0.154), 0.18 (range=0.061-0.556), 0.10 (range=0-0.195), 0.13 (range=0.051-0.244), and
0.17 (range=0.070-0.308) white-lipped peccary tracks per km, calculated from monthly
track surveys throughout 2013, 2014, 2015, 2016, 2017, 2018, 2019, and 2020,
respectively (Figure 4). This suggests that white-lipped peccary abundance based on track
data has declined since the 1990’s until 2004 and then increased through present and
maybe now fluctuating annually. We suspect more recent trends in abundance are likely
associated with weather patterns and food availability.
4. Carrillo’s (2012) report of declining abundance of white-lipped peccaries was based on a
linear-fit model for what is clearly a bimodal dataset. A linear-fit model is statistically
inappropriate in this case. However, this does not invalidate Carrillo’s (2012) cause for
concern, because he detected more recent declines that he believed were associated with
changes in park management. However, recent track data (Azofeifa et al. 2021) suggests
that this abundance index may have recently increased to levels similar to those in 2010.
Track data indicates that white-lipped peccary abundance is relatively high, and generally
stable from 2013-2020 (Azofeifa et al. 2019; Azofeifa et al. 2021). Alternatively, because
there are many factors that could be influencing the abundance of this species (see list
below), it would be inappropriate to suggest any decline was linked to poaching without
data on poaching. Note: we did observe at least one group of poachers during our camera
trap survey in 2015, 2016, 2017, 2018, and 2019. In 2017, we also had six cameras
destroyed or stolen due to poachers and drug runners. In 2020, we had five camera
stations destroyed or stolen.
Factors to consider when interpreting abundance indices for this species:
a. appears to be both relatively stationary (Carrillo et al. 2002; i.e., relatively small
home ranges and limited seasonal movement) and migratory (Altrichter &
Almeida 2003; relatively large home ranges and strong seasonal movements),
b. commonly exhibits strong temporal and spatial population fluctuations (Fragoso
2004; Hansen et al. 2013),
Page 25 of 32
c. appears to be sensitive to anthropogenic killing (Fuller et al. 2002; Fragoso 2004;
Hansen et al. 2013), and
d. is likely very sensitive to changes in food availability (Kiltie & Terborgh 1985),
which could be associated with the shifts in seasonal weather patterns associated
with El Nino (noaa.gov).
Pumas in 2020
We documented 36 camera trap events of pumas (RAI=21.3) at 62% of camera trap sites (Tables
1 & 2). Based on the data presented in Carazo (2009) and Olson et al. (2016, 2017, 2018, 2019,
2020a, 2020b, and this report), puma abundance appears to be stable over time (2003=25.8,
2008=22.5, 2015=52.4, 2016=29, 2017=21.5, 2018=32.9, 2019=16.4; 2020=21.3; Table 1;
Figure 5). Track data indicates that puma abundance is relatively high, but generally declining
from 2013-2018 (Azofeifa et al. 2019; Azofeifa et al. 2021). Based on this data puma abundance
has maintained at relatively high levels and may have increased since 2003.
Collared Peccary in 2020
We documented 46 camera trap events (RAI=27.2) of collared peccaries at 54% of camera trap
sites (Tables 1 & 2). Based on camera trap abundance indices from 2003, 2008, 2015, 2016,
2017, 2018, 2019, and 2020 (Carazo 2009; Olson et al. 2016, 2017, 2018, 2020a, 2020b, and this
report), it appears as though collared peccary abundance has increased since 2003 (Figure 5).
Track data indicates that collared peccary abundance is relatively high, but generally declining
from 2013-2019, but may have increased in 2020 (Azofeifa et al. 2019; Azofeifa et al. 2021).
Baird’s Tapir in 2020
We documented 109 camera trap events of tapirs (RAI=64.3) at 85% of camera trap sites (Tables
1 & 2). Based on the data presented in Carazo (2009) and Olson et al. (2016, 2017, 2018, 2019,
2020a, 2020b, and this report), tapir abundance appears to have increased from 2003 to 2016 and
started to decline more recently – with the notable exception of 2020 (2003=49.5, 2008=70.4,
2015=206.1, 2016=128.6, 2017=93.1, 2018=48.3, 2019=38.2, 2020=64.3; Table 1; Figure 5).
However, these changes in RAI may be influenced by the changes in the distribution of camera
traps and the length of the survey period for each time period. For example, camera traps in 2003
and 2008 were in the same relative area; camera traps from 2015 through 2019 saw a general
reduction in the concentration of cameras within the bajura and along the beach. Including the
track data from Carrillo (2012), tapir track data from 1990 to 2007 (1.3-1.9 tracks/km) were
substantially higher than tapir track data from 2007 to 2019 (0.6-1.4 tracks/km). Due to the
unknown methods that were used for Carrillo (2012) it is hard to determine if there was a
legitimate drop in tapir tracks/km between these two time periods or a change in methodology. If
we look at the track data from 2013-2020 from Azofeifa et al. (2021), we see that tapir tracks/km
were relatively stable over that time period. Camera trap FOC from 2015-2020 (Olson et al.
2016, 2017, 2018, 2020a, 2020b and this report) also indicates that tapir distribution within the
park is relatively widespread and consistent (Table 2). Tapirs in Corcovado appear to be in good
body condition relative to captive and other populations (Olson et al., in prep). Based on track
survey data (Azofeifa et al. 2021), tapirs are often one of the most abundant species detected,
Page 26 of 32
similar to agoutis and coatis (typically in the top three of most abundant species). Altogether, the
data suggests that tapirs are broadly distributed throughout the park, are relatively abundant
throughout the park, and are in good relative health. It is likely that Corcovado National Park is
functioning as a source population of tapirs, sending dispersing individuals outside of the park.
Figure 7. White-lipped peccary distribution within 4X4 grid cells as determined by camera trap
or track survey data collected in Corcovado National Park, Costa Rica from 2013-2020 (areas
without any survey effort are not shaded any color). Camera trap locations for 2020 are shown
for reference.
Page 27 of 32
Margays and Ocelots
Ocelots were documented in 29 separate camera trap events (RAI=17.1) at 77% of camera trap
sites in 2020 (Tables 1 & 2). During 2020, we identified 15 individual ocelots and 7
unidentifiable ocelot events at 10 camera trap stations. 11 ocelots were of an unknown sex, 4
were female, and 7 were male. Ocelots have been documented every year since 2015. Over the
span of this project a total of 88 unique individual ocelots have been detected (52 captured only
once and 36 that have been recaptured at least once), and there were 41 events in which the
image quality was too low to identify to the individual-level. There are several ocelots of
significance. Gordito was first detected in Corcovado in 2008 as a sexually mature male by
Corazo (2008). He was subsequently detected by this project in 2015 and 2016, so his minimum
age is 10 years old. Ocelots are estimated to live 7 to 10 years, thus, Gordito is likely one of the
oldest wild ocelots ever recorded. Estrella, a female found to be at least seven years old (detected
in 2015, 2016, 2018, and 2020) has been captured on video successfully hunting a rodent.
Finally, Margarita, a suspected female, was first captured as an older juvenile traveling with her
mother in 2019, and subsequently captured in 2020 traveling alone.
Margays were documented in 14 separate camera trap events (RAI=8.3) at 38% of camera trap
sites in 2020 (Tables 1 & 2). During 2020, we identified 11 individual margays and three
unidentifiable margay events at 6 camera trap stations. 8 margays were of an unknown sex, 4
were female, and 2 were male. Our detailed analysis of ocelot and margay images for all years of
the study resulted in the need to adjust ocelot and margay events for prior years due to accidental
misidentification of margays as ocelots in prior years by research technicians. Margays were first
captured in 2017 and have been seen every year henceforth. Since 2017, we have detected at
least 24 individual margays (1 margay has been recaptured, 23 margays were only detected only
once), and there were 8 events in which the image quality was too low to identify to the
individual-level. One female (Cazedora) was first seen in 2019 with a neonate, was recaptured in
2020.
Other species of note in 2020
Based on camera trap RAI and FOC, we can explore potential trends in species abundance since
2003. The following species appear to have increased in abundance and distribution since 2003:
crested guan Penelope pupurascens, red brocket deer Mazama americana, nine-banded
armadillo Dasypus novemcinctus, and agouti paca Cuniculus paca. The following species appear
to have decreased and then increased more recently: striped hog-nosed skunk Conepatus
semistriatus, jaguarundi Puma yaguarondi, and the great tinamou Tinamus major.
Osa Red: Ridge to Reef Mega Transect
In 2020, we contributed our survey efforts in the park to support the Osa Red: Ridge to Reef
effort creating a continuous camera trap mega transect that ran from La Amistad National Park
through Piedras Blancas National Park and Corcovado National Park, and ending on the tip of
the Osa Peninsula.
Future Work
Page 28 of 32
Our 2021 survey work is currently underway. For 2021, we attempted to increase the spatial
coverage of the park by installing cameras in three separate phases and adding new camera sites.
However, due to issues associated with the global pandemic and camera shortages from theft or
damage, we were not able to follow our proposal as well as anticipated. Staff availability has
improved dramatically with three staff dedicated to supporting this monitoring effort and their
work to ensure the project’s success has already led to some exciting results for 2021.
We are already entering camera trap some of the data from the 2021 survey effort in an attempt
to produce a summary report as soon as possible. Preliminary review of the data available thus
far indicates that we detected Vivi, a female who hasn’t been detected since 2017, and Macho
Uno.
We aim to continue collaborating with other researchers within MINAE-SINAC and nonprofit
organizations to better understand the status of wildlife within the Osa Peninsula to improve
conservation efforts. Erik Olson currently has funding to partially support the current survey
efforts in Corcovado and Piedras Blancas for two more years (i.e., through 2023) and plans to
seek additional funding to purchase more cameras. While the goals of these future surveys are
still developing – it is important to focus on the development of a long-term monitoring plan that
encompasses the full extent of Corcovado National Park as well as the neighboring wild lands
adjacent to the park.
Finally, the current global pandemic has also stressed our ability to maintain the project –
creating uncertainty and making it impossible for researchers and student research assistants
from the US to travel to Corcovado to support the monitoring efforts. We hope to collaboratively
address these challenges as we move this historical and critically important survey effort
forward.
Professional Activities
Peer-reviewed publications since 2015 (*indicates Northland College student)
Vargas Soto, J.S., Beirne, C., Flatt, E., Pillco-Huarcaya, R., Whitworth, A., Olson, E.R., Saborio-
R, G., Espinoza-Munoz, D., Salom-Perez, R., Cruz Diaz, J.C., Whittaker, L., Roldan, C.,
North Broadbent, E., Molnar, P.K. 2021. Human disturbance shifts the vertebrate
community from large-bodied threatened species to small-bodied generalists in a
neotropical biodiversity hotspot. Conservation Biology. https://doi.org/10.1111/cobi.1381
*Beal, M.R.W., *Matzinger, P.J., Saborio-R, G., Bristan, J.N., Olson, E.R. 2020. A survey of
medium and large mammals of Piedras Blancas National Park, Costa Rica, 2016-18.
Checklist. 16(4): 939-950. https://doi.org/10.15560/16.4.939.
Olson, E.R., Saborío-R, G., Salazar, J.C. 2019. Age of the jaguar: A novel approach to
evaluating the lifespan of a rare carnivore. Cat News. 70:36-38.
Olson, E.R., *Matzinger, P.J., Saborío-R, G., Carazo-Salazar, J.C. 2019. Macho Uno: a sign of
hope for the jaguars of Corcovado National Park, Costa Rica. Cat News. 69:4-6.
Other publications & reports since 2015
Page 29 of 32
Azofeifa, A., Montes, W., Olmos, E., Herrera, D., Solano, E., Olson, E.R., Saborío-R, G. 2021.
Monitoreo de mamiferos medianos y grandes por medio de huellas y rastros, Parque
Nacional Corcovado. Technical Report. SINAC-MINAE.
Olson, E.R., Chevalier, V., Franke, J., Kolasch, J., Saborío-R, G., Azofeifa, A., Olmos, E.,
Montes, W. 2020b. Wildlife monitoring report for Corcovado National Park, Costa Rica -
2019. Technical Report. Pp. 30. Northland College, Ashland, Wisconsin, USA.
https://doi.org/10.13140/RG.2.2.33583.76966/1.
Olson, E.R., *Beal, M.R.W., Saborío-R, G., Azofeifa, A., Olmos, E., Montes, W. 2020a.
Wildlife monitoring report for Corcovado National Park, Costa Rica - 2018. Technical
Report. DOI: 10.13140/RG.2.2.28664.67845
Azofeifa, A., Montes, W., Olmo, E., Olson, E.R., Saborío-Rodríguez, S. 2019. Monitoreo de
mamíferos terrestres medianos y grandes en el parquet nacional Corcovado: Resumen
resultados 2013-2018. SINAC-MINAE Governmental Technical Report – FOI-004-004.
Collaborator. Sistema Nacional de Áreas de Conservación (SINAC). 2018. Estado de
conservación del jaguar (Panthera onca) en Costa Rica a través de la integración de datos
de registro de la especie y modelaje del hábitat idóneo. Technical Report.
Olson, E., *Beal, M., Saborío-R, G., Azofeifa, A., Montes W. 2018. Wildlife Monitoring Report
for Corcovado National Park, Costa Rica – 2017. Technical Report for MINAE-SINAC.
Olson, E., *Matzinger, P., Saborío-R, G. 2016. Wildlife Monitoring Report for Corcovado
National Park, Costa Rica – 2015. Technical Report for MINAE-SINAC.
Olson, E., *Matzinger, P., Saborío-R, G., Azofeifa, A., Montes, W. 2017. Wildlife Monitoring
Report for Corcovado National Park, Costa Rica – 2016. Technical Report for MINAE-
SINAC.
Professional presentations since 2015
Chevalier, V., Azofeifa, A., Saborio-R, G., Olson, E.R. 2021. Use of HotSpotter to assess the
status of ocelots (Leopardus pardalis) and margays (Leopardus weidii) within Corcovado
National Park, Costa Rica. 2021 Wisconsin Chapter of The Wildlife Society Winter
Meeting (virtual).
Beal, M., Niermann, B., Matzinger, P., Saborio-R, G., and E.R. Olson. Poster. 2019. Impacts of
development on presence and abundance of terrestrial species in Piedras Blancas and
Corcovado National Park, Costa Rica. 2019 Wisconsin Chapter of The Wildlife Society
Winter Meeting.
Niermann, B., Saborio-R, G., Matzinger, P., and E.R. Olson. Poster. 2018. Distribution and body
condition scores in wild Baird’s tapir (Taprius bairdii) of Corcovado National Park,
Costa Rica. Midwest Fish & Wildlife Conference. Milwaukee, WI.
Beal, M., Niermann, B., Matzinger, P., Saborio-R, G., and E.R. Olson. Poster. 2018. Diversity
and abundance of terrestrial wildlife in Piedras Blancas National Park, Costa Rica.
Midwest Fish & Wildlife Conference. Milwaukee, WI.
Olson, E.R., Sabario-R, G. and P. Matzinger. 2017. Jaguars and white-lipped peccaries of
Corcovado National Park, Costa Rica. For – Chequamegon Audubon Society.
Page 30 of 32
Matzinger, P., Saborio-R, G., and E.R. Olson. 2016. Poster. Assessing non-invasive monitoring
techniques in Corcovado National Park, Costa Rica. North American Congress for
Conservation Biology. Madison, WI.
Matzinger, P., Olson, E.R., and G. Saborio-R. 2016. Efficacy of combined non-invasive
monitoring techniques for assessing terrestrial mammalian abundance and presence.
2016 Annual Winter Meeting of the Wisconsin Chapter of The Wildlife Society. Stevens
Point, WI.
Acknowledgements
We thank SINAC administration and staff, especially, Sirena Ranger Station administration,
kitchen and maintenance staff. Thanks to local guides from both Drake Bay and Puerto Jimenez
for their cooperation and assistance. Thanks to numerous undergraduates of Northland College
for entering and managing camera trap data. Special thanks to Gordy Scott and Hope Sandlin.
There are many more individuals we are indebted to for their support and encouragement
throughout this project – to all of you – we are grateful.
Page 31 of 32
References (if not listed above)
Apps, P.J., and McNutt, J.W. 2018. How camera traps work and how to work them. African Journal of
Ecology. 56:702-709.
Altrichter, M., and R. Almeida. 2002. Exploitation of white-lipped peccaries (Tayassu pecari
(Artiodactyla: Tayassuidae) on the Osa Peninsula, Costa Rica. Oryx. 36:126-132.
Azofeifa, A., Montes, W., Olmo, E., Olson, E.R., Saborío-Rodríguez, S. 2019. Monitoreo de mamíferos
terrestres medianos y grandes en el parquet nacional Corcovado: Resumen resultados 2013-2018.
SINAC-MINAE Governmental Technical Report – FOI-004-004.
Bone-Guzmán, R. and E. Chacón-Madrigal. 2020. From jaguars to margays: spatial distribution and
conservation of five feline endangered species and their prey in Golfo Dulce Forest Reserve,
Costa Rica. Wild Felid Monitor. 13(2):21-22.
Carazo Salazar, J. 2009. Camios en las poblaciones de jaguares (Panthera onca), sus presas potenciales y
manigordos (Leopardus pardalis), en dos periodos de tiempo sujetos a diferentes esfuerzos de
control de caceria en el Parque Nacional Corcovado, Costa Rica. Thesis, Universidad Nacional –
Sistem de Estudios de Posgrado, Instituto Internacional en Conservacion y Manejo de Vida
Silvestre.
Carrillo, E. 2012. Formal letter to Director of ACOSA and Corcovado dated 14 September 2012.
Carrillo, E., Saenz, J.C., and T.K. Fuller. 2002. Movements and activities of white-lipped peccaries in
Corcovado National Park, Costa Rica. Biological Conservation. 108:317-324.
Carrillo, E., Wong, G., and A.D. Cuarón. 2000. Monitoring mammal populations in Costa Rican
protected areas under different hunting restrictions. Conservation Biology. 14:1580-1591.
De Angelo, C., Paviolo, A., and M.S. Di Bitetti. 2010. Traditional versus multivariate methods for
identifying jaguar, puma, and large canid tracks. Journal of Wildlife Management. 74:1141-1153.
Fragoso J.M.V. 2004. A long-term study of white-lipped peccary (Tayassu pecari) population
fluctuations in northern Amazonia—anthropogenic versus “natural” causes. In K.M. Silvius, R.E.
Bodmer and J.M.V. Fragoso (eds.). People in Nature: Wildlife Conservation in South and Central
America (pp. 286-296). Columbia University Press, New York, USA.
Fuller, T.K., Carrillo, E., and J.C. Saenz. 2002. Survival of protected white-lipped peccaries in Costa
Rica. Can. J. Zool. 80: 586-589.
Fuller, T.K., Judas, J. and O. Henry. 1999. Seasonal variation of home range of collared peccary in
tropical rain forests of French Guiana. The Journal of Wildlife Management. 63:546-552.
Harmsen B.J., Foster R.J., Sanchez E., Gutierrez-Gonzalez C.E., Silver S.C., Ostro L.E.T., Kelly M.J.,
Kay E., & Quigley H. 2017. Long term monitoring of jaguars in the Cockscomb Basin Wildlife
Sanctuary, Belize: Implications for camera trap studies of carnivores. PLOS One
https://doi.org/10.1371/journal.pone.0179505.
Kiltie, R.A. and Terborgh, J. 2010. Observations of the behavior of rain forest peccaries in Perú: Why do
white-lipped peccaries form herds? Ethology. 62:241-255.
Long, R.A., MacKay, P., Zielinski, W.J., Ray, J.C. 2008. Noninvasive survey methods for carnivores, pp.
385. Island Press, Washington DC, USA.
Maffei, L., Noss, A.J., Silver, S.C., and M.J. Kelly. 2011. Abundance/density case study: Jaguars in the
Americas. In: .F. O’Connell et al. (eds.). Camera Traps in Animal Ecology: Methods and
Analyses (pp.119-144). Springer.
Naing, H., Fuller, T.K., Sievert, P.R., Randhir, T.O., Tha Po, S.H., Maung, M., Lynam, A.J., Htun, S.,
Naing, W., and Myint, T. 2015. Assessing large mammal and bird richness from camera-trap
records in the Hukaung Valley of Northern Myanmar. Raffles Bulletin of zoology. 63:376-388.
O’Brien, T.G., Kinnaird, M.F., and Wibisono, H.T. 2003. Crouching tigers, hidden prey: sumatran tiger
and prey populations in a tropical forest landscape. Anim. Conserv., 6: 131-139. Doi:
10.1017/s1367943003003172
Page 32 of 32
Olson, E.R., Beal, M.R.W., Saborío-R, G., Azofeifa, A., Olmos, E., Montes, W. 2020a. Wildlife
monitoring report for Corcovado National Park, Costa Rica - 2018. Technical Report.
DOI: 10.13140/RG.2.2.28664.67845
Olson, E., Beal, M., Saborío-R, G., Azofeifa, A., Montes W. 2018. Wildlife Monitoring Report for
Corcovado National Park, Costa Rica – 2017. Technical Report for MINAE-SINAC.
Olson, E.R., Chevalier, V., Franke, J., Kolasch, J., Saborío-R, G., Azofeifa, A., Olmos, E., Montes, W.
2020b. Wildlife monitoring report for Corcovado National Park, Costa Rica - 2019. Technical
Report. Pp. 30. Northland College, Ashland, Wisconsin, USA.
https://doi.org/10.13140/RG.2.2.33583.76966/1.
Olson, E., Matzinger, P., Saborío-R, G. 2016. Wildlife Monitoring Report for Corcovado National Park,
Costa Rica – 2015. Technical Report for MINAE-SINAC.
Olson, E., Matzinger, P., Saborío-R, G., Azofeifa, A., Montes, W. 2017. Wildlife Monitoring Report for
Corcovado National Park, Costa Rica – 2016. Technical Report for MINAE-SINAC.
Olson, E., Matzinger, P., Saborío-R, G., Carazo-Salazar, J. 2019. Macho Uno: a sign of hope for the
jaguars of Corcovado National Park, Costa Rica. Cat News. 69:4-6.
Palmer, M.S., Swanson, A., Kosmala, M., Arnold, T., Packer, C. 2018. Evaluating relative abundance
indices for terrestrial herbivores from large-scale camera trap surveys. African Journal of
Ecology. 56:791-803.
Parsons, A.W., Forrester, T., McShea, W.J., Baker-Whatton, M.C., Millspaugh, J.J., Kays, R. 2017. Do
occupancy or detection rates from camera traps reflect deer density? Journal of Mammalogy.
98(6):1541-1557.
Plhal, R., Kamler, J., Homolka, M., and Z. Adamec. 2011. An assessment of the applicability of photo
trapping to estimate wild boar population density in a forest environment. Folia Zool. 60: 237-
246.
Polisar, J., O’Brien, T.G., Matthews, S.M., Beckman, J.P., Sanderson, E.W., Rosas-Rosas, O.C., and C.A.
Lopez-González. 2014. Jaguar survey and monitoring techniques and methodologies: A review.
The Wildlife Conservation Society. Contract F13PX01563.
Rivera, C.J. 2014. Facing the 2013 Gold Rush: A population viability analysis for the Endangered white-
lipped peccary (Tayassu pecari) in Corcovado National Park, Costa Rica. Natural Resources, 5:
1007-1019.
Salom-Perez, R. 2005. Ecologia del jaguar (Panthera onca) y del manigordo (Leopardus pardalis)
(Carnivora:Felidae) en el Parque Nacional Corcovado, Costa Rica. Thesis, Universidad de Costa
Rica, Sistema de Estudios de Postgrado.
Salom-Perez, R., Carrillo, E., Sáenz, J.C., and J.M Mora. 2007. Critical condition of the jaguar Panthera
onca population in Corcovado National Park, Costa Rica. Oyrx. 41:51-56.
Sollmann, R. 2018. A gentle introduction to camera-trap data analysis. African Journal of Ecology.
56:740-749.
Stewart, F.E.C., Fisher, J.T., Burton, A.C., and Volpe, J.P. 2018. Species occurrence data reflect the
magnitude of animal movements better than the proximity of animal space use. Ecosphere:
9(2):e02112.10.1002/ecs2.2112.
Tobler, M.W. and G.V.N. Powell. 2013. Estimating jaguar densities with camera traps: Problem with
current designs and recommendations for future studies. Biological Conservation. 159:109-118.
