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Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e20263589, enero-diciembre 2026 (Publicado Abr. 08, 2026)
Species richness and relative abundance of terrestrial mammals
in an area with different managements in Costa Rica
Olatz Erkizia-Suinaga1*; https://orcid.org/0009-0001-9832-7305
Keilor Cordero-Umaña2; https://orcid.org/0000-0002-4874-9577
Adam Yaney-Keller3; https://orcid.org/0000-0003-1538-664X
Víctor H. Montalvo4,5; https://orcid.org/0000-0002-3652-7694
Carolina Sáenz-Bolaños4,5; https://orcid.org/0000-0003-4855-4486
Juan C. Cruz5,6; https://orcid.org/0000-0001-7240-8230
Eduardo Carrillo4,5; https://orcid.org/0000-0002-5303-0858
Pilar Santidrián Tomillo7; https://orcid.org/0000-0002-6895-7218
1. Real Jardín Botánico-CSIC, Plaza Murillo 2, 28014 Madrid, Spain; erkiziaolatz@gmail.com(*Correspondance)
2. Menéndez Pelayo International University (UIMP-CSIC), C. Isaac Peral 23, 28040, Madrid, Spain;
kecordero95@gmail.com
3. School of Biological Sciences, Monash University, 25 Rainforest Walk, Clayton, 3800, Victoria, Australia; adamyks27@
gmail.com
4. Instituto Internacional en Conservación y Manejo de Vida Silvestre, Universidad Nacional, Apdo. 1350-3000, Heredia,
Costa Rica; vmontalvog@gmail.com, carolina.saenz.bolanos@una.ac.cr
5. Namá Conservation, Heredia 40101, Costa Rica; ecarrilloj@gmail.com
6. Amazon Conservation Team, Washington, 22046, United States; saasilbaalam@gmail.com
7. Centre Oceanogràfic de les Balears, Instituto Español de Oceanografía (IEO, CSIC), Moll de Ponent s/n, 07015 Palma
de Mallorca, Spain; mpilar.santidrian@ieo.csic.es
Received 06-XI-2025. Corrected 06-II-2026. Accepted 25-III-2026.
ABSTRACT
Introduction: While essential, protected areas are sometimes insufficient to conserve wildlife. Terrestrial mam-
mals often move between protected and unprotected sites, and the abundance of one species may influence the
occurrence of others.
Objectives: To compare mammal species richness and relative abundance between three nearby sites with differ-
ent management approaches in a tropical dry forest.
Methods: We placed 18 camera traps in three contiguous sites under different management schemes (a national
park – high protection; forest reserve – medium protection; and a non-protected site) in Northwest Costa Rica
between December 31, 2021, and March 31, 2022, and compared the richness and relative abundance of terres-
trial mammal species occurring in each site.
Results: We recorded 730 independent captures corresponding to 18 species. The highest number of species
was recorded in the forest reserve (16 species), but 10 species were found at all three locations. The most abun-
dant species were white-tailed deer, jaguars and tapirs in the National Park, ocelots in the Forest Reserve, and
white-faced capuchins, white-nosed coatis, common opossums, raccoons, and coyotes in the non-protected site.
Generalist species were more commonly detected in the unprotected site, whereas specialist species were highly
reported in the protected sites.
Conclusions: Different management restrictions could affect the presence and relative abundance of terrestrial
mammals. However, other factors such as the presence of rivers, trails, and/or roads could also affect movements
https://doi.org/10.15517/hgmw3k88
TERRESTRIAL ECOLOGY
2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e20263589, enero-diciembre 2026 (Publicado Abr. 08, 2026)
INTRODUCTION
The rates of global biodiversity loss due
to anthropogenic activities have been increas-
ing during the last few decades (Johnson et
al., 2017). Therefore, as a worldwide strategy
to protect wildlife, conservationists set a goal
of increasing the world’s network of protected
areas (PAs) (Naughton-Treves et al., 2005).
Creating PAs is one of the most efficient mech-
anisms to protect wildlife (González-Maya et
al., 2015; Pinheiro et al., 2020). With a clearly
defined geographical space, these areas might
be managed to achieve long-term protection
and conservation of species, with associated
ecosystem services and cultural values (Kareiva
& Marvier, 2015). But unfortunately, due tothe
extension and the lack of resources, many PAs
face uneven protection and external threats
that could limit their conservation success
(Mora & Sale, 2011). PAs are crucial inprovid-
ing refuge to terrestrial mammal species from
direct persecution and other anthropogenic
disturbances (Sáenz-Bolaños et al., 2020). Nev-
ertheless, species may require large areas for
foraging or migration, which may exceed the
boundaries of what PAs can provide (González-
Maya et al., 2015). Thus, the current extent of
many PAs is likely insufficient to protect many
vertebrate species across their entire range
(Wen et al., 2022).
One of the world’s most threatened eco-
systems is the tropical dry forest, particularly
in Central America, where it has been reduced
to less than 20 % of its original extent (Klemens
et al., 2011; Siyum, 2020). In the Northwest
of Costa Rica, large areas of dry forest were
converted to pastures for cattle grazing and
and influence distribution. A detailed analysis of the factors driving species abundance in sites with different
management could improve species protection throughout their range.
Key words: tropical dry forest; camera trapping; Guanacaste Conservation Area; Santa Rosa National Park;
Horizontes Forest Reserve; Playa Cabuyal; protection level.
RESUMEN
Riqueza y abundancia relativa de mamíferos terrestres en un área con diferentes manejos en Costa Rica
Introducción: Las Áreas Protegidas son fundamentales para la conservación de la vida salvaje, pero podrían ser
insuficientes. Los mamíferos terrestres suelen moverse entre zonas protegidas y no protegidas, y la abundancia de
una especie puede influir en la presencia de otras.
Objetivos: Comparar la riqueza y abundancia relativa de mamíferos terrestres en tres sitios cercanos con diferen-
tes enfoques de manejo en un bosque seco tropical.
Métodos: Colocamos 18 cámaras trampa en tres localizaciones contiguas bajo diferente régimen de manejo
(parque nacional – alta protección; reserva forestal – protección media; y una localización sin protección) en el
noroeste de Costa Rica entre el 31 de diciembre de 2021 y el 31 de marzo de 2022, y comparamos la riqueza y
abundancia relativa de especies de mamíferos terrestres en cada localización.
Resultados: Registramos 730 capturas independientes correspondientes a 18 especies. El mayor número de espe-
cies se registró en la reserva forestal (16 especies), pero 10 especies fueron encontradas en las tres localidades. Las
especies más abundantes fueron el venado de cola blanca, jaguar y tapir centroamericano en el Parque Nacional,
el ocelote en la Reserva Forestal y el mono carablanca, coatí de nariz blanca, zarigüeya común, mapache y coyote
en la zona no protegida. Más especies generalistas fueron encontradas en la localización no protegida y más
especialistas en las zonas protegidas.
Conclusiones: Diferentes restricciones de manejo pueden afectar la presencia y abundancia de mamíferos terres-
tres. Sin embargo, otros factores como la presencia de ríos, senderos y/o carreteras también podrían influir en su
distribución. Un análisis detallado de los factores que determinan la abundancia en localizaciones de diferente
manejo podría mejorar la protección de especies a lo largo de su rango de distribución.
Palabras clave: bosque tropical seco; fototrampeo; Área de Conservación Guanacaste, Parque Nacional Santa
Rosa; Estación Experimental Forestal Horizontes; Playa Cabuyal; nivel de protección.
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to selective logging of high-value tree species
during the early XIX century (Klemens et al.,
2011), thereby reducing the extent of dry for-
est to a few isolated small patches (Janzen,
1986). Considering this, in 1971, the Santa Rosa
National Park (SRNP) was created, followed by
the Horizontes Forest Reserve (HFR) in 1989.
Both sites were designed to protect the largest
remnant of tropical dry forest in the Neotropi-
cal region (Janzen, 1986; Janzen & Hallwachs,
2020). Outside SRNP and HFR, the remaining
patches of dry forest are highly fragmented
(Yaney-Keller et al., 2019) and threatened by
anthropogenic fires, land conversion for hotel
developments, or livestock grazing (Reyes,
2012; Vargas, 2016).
In Costa Rica, the national system of PAs
(Sistema Nacional de Áreas de Conservación
[SINAC], for its Spanish acronym) has 12 man-
agement categories, each with specific restric-
tions based on conservation goals (Sistema
Nacional de Áreas de Conservación [SINAC],
2023). For example, SRNP is protected as a
“national park”, which allows tourism and low
anthropogenic alteration and prohibits extrac-
tion (Dudley, 2008). HFR is classified as a
“forest reserve”, which allows active forest res-
toration and sustainable timber extraction
(Organización de las Naciones Unidas para la
Alimentación y la Agricultura [FAO], 2010). It
is surrounded by “fincas” (farms for cattle and
agriculture) without use restrictions, creating a
landscape with differing management regimes
(SINAC, 2023). These differences in protection
level and adjacent land use suggest that PAs
may experience contrasting degrees of anthro-
pogenic pressure, with potential consequences
for wildlife communities. For example, Sáenz-
Bolaños et al. (2020) found that different levels
of protection affected the species richness and
relative abundance of mammal species in Costa
Rican tropical rainforests. In addition, Naing et
al. (2015) reported that within a protected area
(PA) in the subtropical rainforests of Myanmar,
locations with higher human infrastructure
had lower species richness thanthose that were
less impacted by humans. More recent studies
further indicate that anthropogenic pressure
negatively affects mammal assemblages, reduc-
ing species richness and occupancy even within
PAs (Gallego-Zamorano et al., 2020; Streicher
et al., 2025; Zwicker et al., 2025).
Mammal species play a key role in the
ecosystems, and conserving their populations is
essential to ensure thelong-term functionality
of PAs (González-Maya et al., 2015; Dorji et al.,
2019). Previous studies suggest that by compar-
ing mammal diversity inside and outside PAs,
or among sites with different levels of protec-
tion, decision-makers can get baseline informa-
tion to support conservation efforts (Dorji et
al., 2019; Sáenz-Bolaños et al., 2020).
In this study, we used automatic camera
traps to compare terrestrial mammal richness
and relative abundance among contiguous sites
of different conservation management catego-
ries: 1) Santa Rosa National Park, 2) Horizontes
Forest Reserve, and 3) an adjacent unprotected
site: Playa Cabuyal. Previous studies have shown
that there is a lower number of mammal species
in disturbed areas and a generally higher num-
ber of species in PAs, including in regions of
Costa Rica (Sáenz-Bolaños et al., 2020; Vargas-
Soto et al., 2022; Cambronero et al., 2023; John-
son et al., 2023). Thus, we hypothesized that
richness and relative abundance of terrestrial
mammals would differ between our study sites.
Specifically, we predicted higher richness and
relative abundance in sites with greater protec-
tion, with progressively lower values in the less
protected sites.
MATERIALS AND METHODS
Study area: Our research was conducted
in the Guanacaste Conservation Area (GCA),
located in Northwestern Costa Rica along the
Pacific coast (10°53’01” N & 85°46’30” W). This
area encompasses one of the largest remnants of
tropical dry forest ecosystems in Central Amer-
ica, and the vegetation community is character-
ized by a mosaic of evergreen and deciduous
forests, mixed with grasslands and secondary
growth at different regeneration stages (Jan-
zen & Hallwachs, 2020; Kalacska et al., 2004).
The precipitation regime is highly seasonal,
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with anannual average rainfall of 1 600 mm,
a well-defined dry season from December to
April (months ≤ 100 mm of rain), and a rainy
season from May to November (Fuller et al.,
2020). The mean annual temperature is 25 °C,
with maximum temperatures of 31 °C during
the rainy season and above 35 °C during the
dry season (Jiménez et al., 2016; Fuller et al.,
2020). We selected three sites inside the GCA
corresponding to three different management
categories: 1) the Santa Rosa National Park
(hereinafter SRNP, 10°50’35.2” N & 85°42’13.0”
W), a site with high protection restrictions
since 1971; 2) the Horizontes Forest Reserve
(hereinafter HFR, 10°42’56.2” N & 85°34’0.7”
W), a site with intermediate protection, and
regulated human activities and use since 1989;
3) Playa Cabuyal (hereinafter PC, 10°40’32” N
& 85°39’11” W), a site without protection des-
ignation, with a relatively high anthropogenic
presence on the beach during peak tourist sea-
son (December to April) but low during the rest
of the year (Fig. 1).
SRNP is a site in the process of restora-
tion since the1980s that currently includes the
largest remnant of dry forest ecosystem in the
country, covering 387 km2 with a variety of suc-
cessional stages (Klemens et al., 2011; Janzen &
Hallwachs, 2020). The HFR comprises an area
of 72.93 km2, and it is in the early successional
stage compared with SRNP, where intensive
forest uses are carried out (Rigg, 2013). Playa
Cabuyal (PC) includes an important nesting
beach for sea turtles (Santidrián et al., 2015),
surrounded by a mangrove swamp and frag-
ments of tropical dry forest and cattle farms
(Yaney-Keller et al., 2019). Although the man-
grove forest is protected under the Law on the
Maritime Terrestrial Zone No. 6043 (Sistema
Costarricense de Información Jurídica [SCIJ],
1977), PC lacks official protection and has a
relatively high influx of tourists during the day.
Data collection: We used 18 camera traps,
of which six cameras were deployed in PC,
seven in HFR, and five in SRNP, covering a
total area of 2.67 km2, 5.02 km2, and 4.65 km2,
respectively (Table 1). We georeferenced each
camera trap location using a Global Position-
ing System unit (GPS; model Garmin eTrex
Fig. 1. Study area and location of the camera traps established between December 31, 2021 and March 31, 2022. In dark
grey are national parks (Santa Rosa and Guanacaste National Park, the latest not included in this study), in medium grey is
the forest reserve (Horizontes Forest Reserve), in light grey is protected marine site and in white is site non-protected, all of
them within the Guanacaste Conservation Area. Circles indicate the location of camera traps at Santa Rosa National Park,
Horizontes Forest Reserve and Playa Cabuyal.
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10, Olathe, Kansas, USA) and placed all the
cameras at a minimum distance of 1-2 km apart
and 30-40 cm above the ground, following rec-
ommendations by Montalvo (2020). We set the
cameras to be active for 24 hours per day and
to take three photos per shot at 15 s intervals
when triggered to avoid recording the same
event (Naing et al., 2015). The cameras used a
medium-sensitivity passive infrared (PIR) sen-
sor to minimize false activations. Videos were
recorded over 15 or 30 s per shot in three of
Santa Rosa’s cameras. We checked the cameras
every 10-20 days. During each visit, we down-
loaded the information, replaced the memory
card, and changed the batteries as necessary.
We collected data between December 31, 2021
and March 30, 2022 in PC, January 16, 2022
and March 31, 2022 in HFR, and December 31,
2021 and March 31, 2022 in SRNP.
Species identification and sampling
effort: We used CPW Photo Warehouse
(https://cpw.state.co.us) to archive, identify,
summarize, and analyse photo data (Newkirk,
2016). Species identification was conducted
following Reid and Gómez-Zamora (2022).
We considered that photos were independent
captures when: 1) photos were separated by
more than 60 min, 2) in consecutive photos of
the same species, we could identify animals at
an individual level, and 3) photos of individuals
of the same species were separated by photos of
another species (Montalvo, 2020; O’Brien et al.,
2003). To estimate sampling effort, we counted
the number of nights each camera was active,
adjusting those periods when cameras were
inactive or malfunctioned. If the camera was
not pointing in the correct direction or height,
the photos were excluded, and the correspond-
ing nights were subtracted from the sampling
effort. We plotted accumulation curves with a
logistic adjustment to compare species richness
(number of species within a defined region) at
the three sample sites, counting the number
of camera trap nights required to reach the
expected number of species in each site, follow-
ing Naing et al. (2015) and Sáenz-Bolaños et
al. (2020). We assessed inventory completeness
using incidence-based nonparametric richness
estimators with camera nights as the sampling
unit (presence/absence per species per opera-
tional camera-night) following Rovero et al.
(2016). Expected richness was estimated using
Chao2, and completeness was calculated as the
ratio between observed and expected richness.
Inventory completeness ranges from 0 to 1,
with values closer to 1 indicating a more com-
plete species inventory.
We calculated the relative abundance index
(RAI, number of independent captures / 100
trap nights) (Montalvo et al., 2023; Rovero &
Marshall, 2009) of each species for each camera
and averaged the RAI for each species for all
cameras at each location. Statistical analyses
and comparisons were made considering the
mean RAI. For species with more than 10
detections in total, we performed a Chi-square
test to compare RAI between the three loca-
tions (Naing et al., 2015; Sáenz-Bolaños et al.,
2020), using the R “stats” package (R Core
Team, 2017).
To quantify the differences between mam-
mal communities under the three manage-
ment categories, we used two dissimilarity
indices: (1) the Chao estimator of shared spe-
cies, which estimates the number of shared spe-
cies between each site (Chen et al., 1995), and
(2) the Chao-Jaccard (C-J) similarity estimator,
which is a measure of beta-diversity and ranges
from 0 (completely distinct communities) to
Tabl e 1
Minimum, maximum, and mean distances (km) between
the cameras placed between December 31, 2021 and March
31, 2022 and the total extension (km2) that comprised the
cameras in each locality.
PC HFR SRNP
Min. Dist. (km) 1.83 1.17 1. 87
Max. Dist. (km) 3.43 5.47 6.38
Mean Dist. (km) 1.94 2.08 6.64
Total extension (km2) 2.67 5.02 4.65
Min. Dist.: Minimum distance, Max. Dist.: Maximum
distance, Mean Dist.: Mean distance, PC: Playa Cabuyal,
HFR: Horizontes Forest Reserve, SRNP: Santa Rosa
National Park.
6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e20263589, enero-diciembre 2026 (Publicado Abr. 08, 2026)
1 (identical communities; Chao et al., 2000).
Both metrics correct for under-sampling bias
by estimating the number of ‘unseen’ shared
species between sites (Chao et al., 2005). To
visualize the difference in mammal assemblages
between sites, we performed a non-metric mul-
tidimensional scaling (NMDS) ordination with
two axes, using the C-J similarity estimator
and metaMDS function of the vegan package
version 2.6.10 (Oksanen et al., 2014). Statisti-
cal differences in species composition between
the three sites were tested using permutational
multivariate analysis of variance (PERMANO-
VA), with the adonis function from the vegan
package (Oksanen et al., 2014). Finally, we used
the envfit function from the vegan package and
applied 999 permutations to the species detec-
tions and ordination axis scores to identify the
key species that contributed most to variation
in assemblage structure among camera trap
locations (Oksanen et al., 2014).
RESULTS
We recorded a total of 17 470 pho-
tos with an effort of 842 camera trap nights.
Additionally, we obtained 2 290 photos of ter-
restrial mammals of which, 558 photos were
obtained at PC over 286 nights, 771 photos at
HFR over 330 nights, and 961 photos at SRNP
over 226 nights.
We recorded 728 independent captures
and identified 18 species of terrestrial mam-
mals between all sites. We detected 14 species
of mammals in PC from 11 families and six
orders, 16 species in HFR from 12 families and
seven orders, and 13 species in SRNP from nine
families and six orders (SMT 1).
The number of shared species was simi-
lar between all sites (Table 2). Among them,
ten species were present in PC, HFR, and
SRNP: Central American agoutis (Dasyprocta
punctata), white-nosed coatis (Nasua narica)
(Fig. 2A), raccoons (Procyon lotor), white-faced
monkeys (Cebus imitator), ocelots (Leopardus
pardalis), pumas (Puma concolor) (Fig. 2B),
Baird’s tapirs (Tapirus bairdii) (Fig. 2C), tayras
(Eira barbara), white-tailed deers (Odocoileus
virginianus), and common opossums (Didelphis
marsupialis). The C-J similarity index indicates
that HFR and SRNP had the highest similar-
ity in species composition (0.93 C-J similarity
Fig. 2. Photo captures. A. A white-nosed coati (Nasua narica) in PC. B. A puma (Puma concolor) in HFR. C. A Baird’s tapir
(Tapirus bairdii) in PC. D. A jaguar (Panthera onca) in HFR (d).
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Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e20263589, enero-diciembre 2026 (Publicado Abr. 08, 2026)
index). PC and SRNP also had a relatively high
similarity (0.74 C-J similarity index, Table 2),
although it was lower than that of HFR and
SRNP. Three species were only found in PC and
HFR: coyotes (Canis latrans), collared peccaries
(Dicotyles tajacu), and grey foxes (Urocyon
cinereoargenteus); and two species in HFR and
SRNP: striped hog-nosed skunks (Conepatus
semistriatus) and jaguars (Panthera onca) (Fig.
2D). Also, three species were unique to each
site: spotted skunk (Spilogale angustifrons) in
PC, tamandua (Tamandua mexicana) in HFR,
and jaguarundi (Herpailurus yagouaroundi) at
SRNP. These differences in species composition
were further confirmed by permutation tests
(PERMANOVA: F = 2.45, p < 0.001). Addition-
ally, NMDS ordination revealed high correla-
tions between several species and specific sites.
For example, coyotes (R2 = 0.60, p < 0.01) and
raccoons (R2 = 0.32, p < 0.05) were associated
with PC, while ocelots (R2 = 0.37, p < 0.05),
striped hog-nosed skunks (R2 = 0.39, p < 0.05),
and Central American agoutis (R2 = 0.30, p <
0.05) were associated with HFR (Fig. 3).
Differences in the number of recorded spe-
cies were illustrated by the species accumulation
Tabl e 2
Number of shared taxa and Chao-Jaccard similarity index
between three areas with different management categories.
Study sites PC HFR SRNP
PC 10 (14.9) 13 (17.8)
HFR 0.74 11 (12.7)
SRNP 0.88 0.93
Playa Cabuyal (PC), Horizontes Forest Reserve (HFR),
and Santa Rosa National Park (SRNP). Upper diagonal:
observed number of shared taxa (Chao-estimated number
of shared taxa in parentheses). Lower diagonal: Chao–
Jaccard similarity index (0–1; higher values indicate greater
community similarity). Gray diagonal cells indicate within-
site comparisons.
Fig. 3. Non-metric multidimensional scaling (NMDS) ordination of the terrestrial mammal species composition detected
by camera traps in Playa Cabuyal, Horizontes Forest Reserve, and Santa Rosa National Park. The NMDS plot area is derived
from relative abundance data for each species at each site. The species represented in the graph contributed the most to the
difference in species composition between the sites.
8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e20263589, enero-diciembre 2026 (Publicado Abr. 08, 2026)
curve (Fig. 4). The curves indicated that HFR
had the highest species richness (n = 16, R2 =
0.97), followed by SRNP (n = 13, R2 = 0.95),
and PC (n = 14, R2 = 0.91) (Fig. 4). None of
the curves reached an asymptote. Neverthe-
less, inventory completeness was high for HFR
(0.89) and SRNP (0.85), suggesting that sam-
pling captured most of the species present at
these sites. In addition, Chao2 estimated 18.0
and 15.2 species respectively, suggesting that
only 2-3 species may have been undetected. In
contrast, PC exhibited a lower completeness
(0.52), with Chao2 estimating 25.2 species,
compared with 14 observed, implying that ~ 12
species may have been undetected.
The most abundant species across the
sample sites were white-tailed deer and com-
mon opossums. We found statistically signifi-
cant differences in relative abundances among
locations for eight species: white-nosed coatis,
white-faced monkeys, common opossums, rac-
coons, and coyotes were significantly more
abundant in PC, whereas white-tailed deer, jag-
uars, and tapirs were more abundant in SRNP,
and ocelots in HFR (p-value < 0.05 all cases)
(Table 3, SMF 1).
Though we did not find statistically sig-
nificant differences considering all the analysed
species, we recorded a higher number of cap-
tures of Central American agoutis in HFR (RAI
= 7.54) than in SRNP (RAI = 5.70) and PC (RAI
= 0.61). Pumas detection was higher in SRNP
(RAI = 3.44), followed by HFR (RAI = 2.68)
and PC (RAI = 0.43). We detected more striped
hog-nosed skunks in HFR (RAI = 4.75), fol-
lowed by SRNP (RAI = 2.16), with no records
of presence in PC (Table 3). The observed
differences in species RAI between sites are
further supported by the results of the NMDS
ordination (Fig. 3).
Six of the species we detected were listed
as endangered (ocelot, puma, tapir, jaguar,
jaguarundi, and spotted skunk), and one addi-
tional species (peccary) was locally threatened
according to the official list of endangered
species in Costa Rica (SCIJ, 2017). Tapirs and
jaguars were also listed as endangered and near
threatened, respectively, under the IUCN Red
List of Threatened Species (IUCN, 2024).
DISCUSSION
We hypothesized that terrestrial mammal
richness and relative abundance would dif-
fer among the studied sites, predicting higher
Fig. 4. Species accumulation curves. Trend lines for the total cumulative number of species based on the number of camera
trap nights (effort) in Horizontes Forest Reserve (HFR: black line and dots), Santa Rosa National Park (SRNP: grey line and
dots), and Playa Cabuyal site (PC: dashed line and white dots) between December 2021 – March 2022 sampling period.
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values in areas under stricter protection. Our
results partially supported this prediction.
Although mammal composition was similar
between sites, we found differences in species
richness and relative abundance. Contrary to
our prediction, the highest species richness
was recorded in the forest reserve, followed
by the national park and the unprotected site.
However, several species showed higher relative
abundance in the national park compared to
the other sites. Species typically associated with
disturbed habitats were more frequent in the
unprotected area.
Our results indicate that the composition
of mammal communities exhibits little varia-
tion across the studied sites, as evidenced by
a C-J similarity index greater than 0.70 in all
cases. This high similarity is likely driven by
the fact that all sites are located within the same
tropical dry forest matrix (Montalvo et al., 2019;
Yaney-Keller et al., 2022). In contrast, species
richness varied across sites, with both PAs hav-
ing higher richness than the unprotected site,
with the highest richness in HFR, followed by
SRNP. However, inventory completeness was
low in PC, indicating that the observed rich-
ness estimates for this site should be interpreted
with caution. The reduced detectability of some
species in disturbed, human-transited areas
and the presence of transient species at PC
may have contributed to an underestimation
of richness. Consistent with this interpretation,
Tabl e 3
Relative abundance indices (RAI) for each sampled site and the Chi-square test results.
Scientific name Common name
Conservation
status RAI
X2p-value
CR IUCN PC
n = 286
HFR
n = 330
SRNP
n = 226
Didelphis marsupialis Common opossum NC LC 24.30* 1.36 7.42 25.63 2.70E-06
Cebus imitator White-faced monkey NC VU 15.09* 1.94 0.36 22.09 1.26E-05
Nasua narica White-nosed coati NC LC 7.93* 1.13 1.44 8.42 0.01
Canis latrans Coyote NC LC 6.63* 3.90 0 6.33 0.04
Procyon lotor Northern raccoon NC LC 5.13* 0.21 0.72 7.25 0.03
Leopardus pardalis Ocelot EN LC 0.28 9.94* 1.74 13.60 0.001
Odocoileus virginianus White-tailed deer NC LC 0.61 12.59 100.99* 157.93 2.2E-16
Panthera onca Jaguar EN NT 0 2.99 12.81* 17.06 0
Tapirus bairdii Baird’s tapir EN EN 0.28 2.93 7.86* 0.018 0.02
Dasyprocta punctata Central American agouti NC LC 0.61 7.54 5.70 5.58 0.06
Puma concolor Puma EN LC 0.43 2.68 3.44 2.24 0.33
Conepatus semistriatus Striped hog-nosed skunk NC LC 0 4.75 2.16 4.91 0.09
Dicotyle tajacu Collared peccary TLC 0.38 2.47 0 3.72 0.16
Eira barbara Tayra NC LC 0.19 0.84 1.01 - -
Herpailurus yagourandi Jaguarundi EN LC 0 0 0.36 - -
Urocyon cinereoargenteus Grey fox NC LC 0.61 4.23 0 - -
Tamandua mexicana Northern tamandua NC LC 0 0.31 0 - -
Spilogale angustifrons Spotted skunk EN LC 0.28 0 0 - -
PC: Playa Cabuyal, HFR: Horizontes Forest Reserve, SRNP: Santa Rosa National Park. Statistically significant differences
between the three locations are highlighted with an asterisk (*). The n value was the number of camera trap nights. The
conservation status of each species followed the official list of species in danger of extinction and with reduced and threatened
populations of Costa Rica (referenced as CR in the table; abbr.: EN – Endangered, T – Threatened, and NC – Not catalogued;
SCIJ, 2017) and The IUCN Red List of Threatened Species (referenced as IUCN in the table; abbr.: EN – Endangered, VU
– Vulnerable, NT – Near threatened, and LC – Least concerned; International Union for Conservation of Nature [IUCN],
2024).
10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e20263589, enero-diciembre 2026 (Publicado Abr. 08, 2026)
Yaney-Keller et al. (2022) reported seven mam-
mal species at PC that were not detected in
the present study (e.g., jaguarundi, Virginia
opossum, jaguar, Eastern cottontail, variegated
squirrel, Northern tamandua), suggesting that
additional sampling effort would be required to
detect rare or transient species. Similarly coy-
otes and collared peccaries were not detected in
SRNP despite the high inventory completeness,
even though both species are known to be pres-
ent (Montalvo et al., 2015).
Despite higher richness in protected areas,
greater richness is not necessarily aligned with
stricter protection. Although stricter protection
often enhances habitat preservation, sites with
lower levels of protection can sometimes host
more species due to their diverse management
practices and land-use mosaics (Naughton-
Treves et al., 2005; Geldmann et al., 2013).
Accordingly, mixed-use landscapes, such as
HFR, may substantially contribute to conserva-
tion by providing complementary habitats that
support both generalist and specialist species.
We found differences in the occurrence
and relative abundance of some species among
the study sites, suggesting that differing levels
of protection may shape mammal presence and
abundance across the landscape (Jordan et al.,
2016; Pérez-Solano et al., 2018; Vargas-Soto et
al., 2022). However, it is important to note that
although differences were found between sites
with varying management, our results do not
indicate causality. A more detailed analysis of
the different factors that could drive wildlife
distributions would help to better pinpoint driv-
ers of abundance. For instance, rivers, streams,
trails, or roads can facilitate or impede animal
distributions across the landscape (Cusack et
al., 2015; Hill et al., 2021), thereby conditioning
their presence or absence at each site.
Regardless of causality, differences were
detected between sites based on management
type. For instance, some large herbivores and
carnivores, such as white-tailed deer, tapirs, and
jaguars, were more abundant in the National
Park. On the other hand, generalist species
such as common opossums, raccoons, coyotes,
white-nosed coatis, and white-faced monkeys
were observed regularly in PC. Sáenz-Bolaños
et al. (2020) found a higher abundance of large
herbivores and large carnivores in Barbilla
National Park compared to other PAs with a
lower level of protection (Indigenous territory
and forest reserve) in the Northern Talamanca
Mountains of Costa Rica. Our study indicated
similar results in the PAs, where there was likely
less anthropic pressure and high food avail-
ability, possibly providing resources to special-
ist species. On the contrary, generalist species
and mesopredators such as white-nosed coatis,
raccoons, and common opossums, which can
tolerate or thrive in proximity to human settle-
ments and anthropogenic activities, showed
higher occurrences outside PAs (i.e., PC), where
human impacts are expected to be higher.
Our results showed the same patterns
found by Montalvo (2012), who reported low
records of coyote, grey fox, collared peccary,
and spotted skunks with camera traps in the
core of SRNP. On the contrary, Yaney-Keller et
al. (2022), also with camera traps, found abun-
dant records of grey foxes, collared peccaries,
andspotted skunks at PC. Coyotes are a native
species in Costa Rica and have been spotted at
the three study sites at different times. It has
been suggested that coyote populations are
expanding in Central America (Hody & Kays,
2018), which may influence their distribu-
tion across the region. However, this seems
insufficient to explain the variation in local
detection rates.
Disturbed and urbanized sites can increase
the number of individuals of generalist species
(Prange & Gehrt, 2004; Prange et al., 2004).
Our results support that generalist species and
mesopredators occur mostly in the most per-
turbed site among the studied sites. This occur-
rence, known as mesopredator release, happens
when the absence of top predators leads to an
increase in generalist species due to reduced
predation pressure (Crooks & Soulé, 1999;
Prugh et al., 2009). Human-derived food sourc-
es may also enhance the success of mesopreda-
tors and generalist species (Oro et al., 2013).
Among the sample sites, PC was the one with
the highest level of anthropogenic activity, as
11
Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e20263589, enero-diciembre 2026 (Publicado Abr. 08, 2026)
there is ahigh transit of tourists that frequently
visit the beach during the daytime, especially
on weekends. As a result, garbage accumulates
during tourist peak season (from November to
April), providing food sources to generalist spe-
cies, such as coyotes and raccoons. On the other
hand, predation of sea turtle eggs by raccoons
has become a major problem at Las Baulas
National Park (Cordero-Umaña et al., 2026),
and predation by coyotes has also been con-
firmed at Cabuyal (unpublished data). These
species can rapidly increase their abundance
and suppress smaller competitor populations,
such as striped hog-nosed skunks or grey foxes
(Jachowski et al., 2020). Such dynamics illus-
trate the critical role that top predators play in
maintaining ecological balance and highlight
how anthropogenic activities can favor gener-
alist species over specialized ones (Ritchie &
Johnson, 2009).
Keystone species with specific ecological
requirements, such as large carnivores (e.g., jag-
uars and pumas) and herbivores (e.g., tapirs and
white-tailed deer), were more abundant inside
PAs. These species require extensive areas of
well-conserved forests that provide shelter from
human disturbance (Laidlaw, 2000; Stoner &
Timm, 2004). In addition, large species and
top predators naturally occur in low popula-
tion densities and have low reproductive rates,
which makes them more vulnerable to habitat
degradation and anthropogenic pressures. This
makes them at the same time good indica-
tors of the quality of the environment and its
change over time (bioindicators; Díaz-Pulido
et al., 2015). Therefore, the higher abundance
of specialist species, top predators, and large
herbivores we found in the PAs could indicate a
recovery of the forests and the wild populations
that inhabit there compared to the unprotected
site (i.e., PC).
The presence of prey influences thepres-
ence of predators (Karanth et al., 2004; Mon-
talvo et al., 2015), and in our study, the high
abundance of white-tailed deer and the season-
al large availability of sea turtles in SRNP could
also explain the high abundance of jaguars
(Montalvo, 2012). Jaguars are regular predators
of sea turtles in SRNP (Alfaro et al., 2016), par-
ticularly at Nancite (Fonseca et al., 2020), where
synchronized arribadas of hundreds of turtles
occur, providing high-energy and readily acces-
sible prey (Carrillo et al., 2009; Fonseca et al.,
2009; Montalvo et al., 2020). This could also
explain the high abundance of jaguars in SRNP
compared to HFR and PC. In fact, the number
of jaguars has increased in recent years, but not
the percentage of sea turtles predated (Fon-
seca et al., 2020). In addition, no jaguars were
detected in PC during the study, despite being
Cabuyal a sea turtle nesting beach, possibly due
to the relatively high level of tourist inflow. In
PC, forest patches are structurally disconnected
from HFR and SRNP and are dominated by
anthropogenic activities (farmlands). Although
we did not find jaguars in PC, they have been
observed before (Yaney-Keller et al., 2022). One
jaguar recorded in HFR in this study was pre-
viously spotted in PC in 2020 (Fonseca, pers.
Communication). Likewise, a jaguar previously
seen in PC was originally identified at SRNP
(Yaney-Keller et al., 2022). This may indi-
cate that at least some jaguars naturally move
between the three sites, even if their abundance
is far greater in the site with the highest level
of protection and greatest forest cover and sea
turtle abundance.
Interspecific competition and seasonal-
ity can also define the composition of mam-
mal communities in the tropical dry forest.
Therefore, the harsh conditions that are typical
during the dry seasons can limit food avail-
ability (Stoner & Timm, 2011). For instance,
jaguars are known to directly compete with
pumas (Montalvo, 2012; Montalvo et al., 2015;
Montalvo et al., 2023), while ocelots can also
be outcompeted and displaced by larger car-
nivore species (Montalvo, 2012; Oliveira et al.,
2010). Hence, Montalvo (2012) suggested that
the high abundance of jaguars in SRNP pushes
pumas to consume smaller prey species, which
would normally be species that ocelots prey
upon, having a cascade effect on the ecosystem.
Our results also support Oliveira et al. (2010)
conclusions about interspecific competition
since we found a low abundance of ocelots
12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e20263589, enero-diciembre 2026 (Publicado Abr. 08, 2026)
in SRNP, in contrast with HFR, where ocelot
populations were possibly high due to the lower
frequency of jaguars and pumas.
Tapirs generally require large areas of well-
preserved forest (Chassot et al., 2009; Stoner
& Timm, 2011). Thus, a higher abundance
obtained in the SRNP compared to other sam-
ple sites was expected. Tapirs are listed as
endangered in Costa Rica (SCIJ, 2017) and at
a global level (IUCN, 2024) and are considered
keystone species for the tropical dry forest, due
to their important role in seed dispersal and
vegetation regeneration (Chassot et al., 2009;
O’Farrill et al., 2013). We detected tapirs in
all three sites during this study, being the first
scientific record of this species in PC. Tapirs are
highly dependent on waterholes during the dry
season, and their observations are not common
outside waterhole sites (Montalvo et al., 2019).
The observation of tapirs in PC may, therefore,
indicate that non-protected sites could work
as transit or occasional use sites for large her-
bivore species, as suggested by the detection
of a white-tailed deer in this study and previ-
ous records in 2017 (Yaney-Keller et al, 2022).
This indicates that PAs’ adjacent sites could be
important to the preservation of this threatened
species.
Finally, endangered or threatened species
were encountered more frequently at SRNP
than at HFR or PC. Though differences in
relative abundance between sites were not sta-
tistically significant for some of the evaluated
species, a larger sampling effort would be nec-
essary to obtain more independent captures of
rare species. SRNP might act as a source for
populating other sites. Previous records indi-
cate that some endangered species (e.g., jagua-
rundi and jaguars) also utilize PC (Yaney-Keller
et al., 2022), though their use is infrequent and
was not detected during this study.
In conclusion, our findings suggest that
different management strategies could affect
the presence and relative abundance of terres-
trial mammals at the Guanacaste Conservation
Area, likely reflecting a positive impact of PAs
on the conservation of the tropical dry forest.
The records of top predators, large herbivores,
and endangered species at PC suggest that
attention should also be paid to unprotected
sites in relation to protected ones. A greater
effort is needed to conduct more thorough
analyses on the different factors that drive wild-
life distributions in the Guanacaste Conserva-
tion Area and on how management categories
could specifically condition their abundance.
Ethical statement: The authors declare
that they all agree with this publication and
made significant contributions; that there is no
conflict of interest of any kind; and that we fol-
lowed all pertinent ethical and legal procedures
and requirements. All financial sources are fully
and clearly stated in the acknowledgments sec-
tion. A signed document has been filed in the
journal archives.
Ver material suplementario
a16v74n1-suppl. 1
ACKNOWLEDGMENTS
We thank Ricardo Bedoya Arrieta from
FONAFIFO (Costa Rica’s Forest Financing
Fund) and The Leatherback Trust for their
contributions to this study. We also thank the
Horizontes Forest Station team for their sup-
port during this research, as well as all the col-
leagues who assisted during the fieldwork. We
finally thank Roger Blanco and the Guanacaste
Conservation Area for providing the research
permits necessary to conduct this study.
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