Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e53245, enero-diciembre 2023 (Publicado Jun. 28, 2023)

Medium and large-sized mammals in a premontane

moist forest fragment, Atenas, Costa Rica

Milena Cambronero1*; https://orcid.org/0000-0002-8117-957X

Ricardo Sánchez-Calderón1,2; https://orcid.org/0000-0003-1303-8545

Rodney Lobo1,3; https://orcid.org/0009-0008-5161-1609

1. School for Field Studies (SFS), Center for Ecological Resilience Studies, Atenas, 20501, Alajuela, Costa Rica;

mcambronero@fieldstudies.org (Correspondence*), rsanchez@fieldstudies.org

2. Sistema de Estudios de Posgrado, Escuela de Biología, Universidad de Costa Rica, San Pedro de Montes de Oca,

11501-2060, San José, Costa Rica.

3. Asociación de Desarrollo Específica para la Conservación del Ambiente (ADECA), Atenas, 20501, Alajuela, Costa

Rica; rodmyml01@gmail.com

Received 26-XI-2022. Corrected 20-III-2023. Accepted 08-VI-2023.

ABSTRACT

Introduction: Medium and large-sized mammals are threatened globally due to habitat loss and fragmentation.

In Costa Rica, forest fragments that remain outside protected areas may serve as a refuge or corridor for these

animals. Mammal presence in fragmented landscapes may be influenced by resource availability, which varies

greatly in seasonal regions.

Objective: To determine the species richness of medium and large-sized mammals in a forest fragment, and to

examine if species diversity and detectability varied between the wet and dry season.

Methods: We established 26 camera trap stations throughout the Municipal Forest of Atenas (Alajuela, Costa

Rica), a 26.4 ha forest fragment that provides water to about 1 800 people. From August 2021 to April 2022, one

camera trap was active at each station for 12-21 days throughout each season (dry and wet), for a total of 810

camera trap days. We applied rarefaction and extrapolation curves to characterize mammal’s species diversity

and compare it between seasons.

Results: We detected 19 species of mammals. The most frequently detected species were the White-nosed Coati

(Nasua narica; 0.254 detections/camera trap day), and the Central American Agouti (Dasyprocta punctata;

0.163 detections/camera trap day). We also registered four species considered as endangered in Costa Rica, three

felines: Puma concolor, Herpailurus yagouaroundi, Leopardus pardalis, and the Neotropical River Otter, Lontra

longicaudis. Seasonality did not affect species diversity, but it influenced dominant species and detectability,

which was greater during the wet season.

Conclusions: Our study demonstrates how protecting small forest fragments, can help conserve endangered

medium and large-sized mammals in human-dominated landscapes.

Key words: Alajuela; Municipal Forest of Atenas; camera trapping; checklist; fragmentation; seasonality; water

protection.

RESUMEN

Mamíferos medianos y grandes en un fragmento de bosque húmedo premontano, Atenas, Costa Rica

Introducción: Los mamíferos medianos y grandes están amenazados globalmente debido a la pérdida y frag-

mentación de su hábitat. En Costa Rica, los fragmentos de bosque fuera de áreas protegidas pueden servir como

https://doi.org/10.15517/rev.biol.trop..v71i1.53245

ECOLOGÍA TERRESTRE

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e53245, enero-diciembre 2023 (Publicado Jun. 28, 2023)

INTRODUCTION

Medium (1-15 kg) and large-sized (> 15

kg) mammals are essential for the long-term

functioning of terrestrial ecosystems, serving

as seed dispersers and top predators, regulating

plant populations, and fostering bioturbation

(Lacher et al., 2019; McConkey et al., 2012).

These animals are particularly diverse in the

Neotropics (Burgin et al., 2018; Pillay et al.,

2022), where land use intensification (e.g.

agriculture and urban expansion) has led to

forest fragmentation, isolation, and habitat loss

(Tabarelli et al., 2004). These factors, along

with illegal hunting, roadkill, and poor farming

practices, pose significant challenges to the

survival of Neotropical mammals (Alroy, 2017;

Bogoni et al., 2020; Crooks et al., 2017; Ripple

et al., 2016; Vetter et al., 2011).

Costa Rica occupies 0.03 % of the world’s

continental area (51 100 km²), but is home

to 256 species of mammals, accounting for

approximately four percent of the world’s mam-

malian biodiversity (Ramírez-Fernández et al.,

2023). This country serves as a corridor for

North and South American species, there-

fore, maintaining habitat connectivity in this

region is essential for the long-term survival

of Neotropical mammals. Costa Rica is well-

known for its conservation efforts, with more

than a quarter of its land area under some form

of protection (Sistema Nacional de Áreas de

Conservación, 2022). Nonetheless, protected

areas have become increasingly isolated, with

forest patches embedded in agricultural and

urban matrices dominating the surrounding

landscape (Morera & Sandoval, 2019; Stan &

Sánchez-Azofeifa, 2019).

Regional efforts have been implemented

to maintain and promote connectivity between

protected areas. For instance, the country has

been part of the Mesoamerican Biological

Corridor since 1997 (Holland, 2012) and of

the National Program of Biological Corridors

since 2006 (Sistema Nacional de Áreas de

Conservación, 2018). However, human activi-

ties continue to expand in critical areas such as

the Central Valley, where the capital is located,

and where forest connectivity is either poor or

non-existent.

Tropical forest fragments play a crucial

role connecting protected areas, facilitating

gene flow, and providing ecosystem services

in agricultural and urbanized landscapes (Beca

et al., 2017; Coulon et al., 2004; Edwards et

al., 2014; Williams et al., 2022). The presence

refugio o sitio de paso para estos animales. La presencia de mamíferos en ambientes fragmentados puede estar

influenciada por la disponibilidad de recursos, la cual puede variar ampliamente en sitios con una estacionalidad

marcada.

Objetivo: Determinar la riqueza de especies de mamíferos medianos y grandes en un fragmento de bosque, y

examinar si la diversidad y detectabilidad de estas especies variaron entre la época seca y lluviosa.

Métodos: Establecimos 26 estaciones de cámaras trampa en el Bosque Municipal de Atenas (Alajuela, Costa

Rica), un fragmento de bosque de 26.4 ha, que brinda agua potable a cerca de 1 800 personas. Desde agosto de

2021 hasta abril de 2022, una cámara trampa estuvo activa en cada estación durante 12-21 días en cada época

(seca y lluviosa), para un total de 810 días cámara trampa. Aplicamos curvas de rarefacción y extrapolación para

caracterizar la diversidad de especies y compararla entre la época seca y lluviosa.

Resultados: Registramos 19 especies de mamíferos medianos y grandes, siendo las más detectadas el pizote

(Nasua narica; 0.254 detecciones/día cámara trampa) y la guatusa (Dasyprocta punctata; 0.163 detecciones/

día cámara trampa). También, registramos cuatro especies consideradas en peligro de extinción en Costa Rica,

tres felinos: Puma concolor, Herpailurus yagouaroundi, Leopardus pardalis y la nutria, Lontra longicaudis. No

observamos un efecto de la estacionalidad en la diversidad, pero sí en la detectabilidad, la cual fue mayor durante

la época lluviosa.

Conclusiones: Nuestro estudio demuestra cómo la protección de parches de bosque pequeños puede contribuir

a la conservación de mamíferos en paisajes fragmentados.

Palabras clave: Alajuela; Bosque Municipal de Atenas; foto trampeo; lista de especies; fragmentación; estacio-

nalidad; protección del agua.

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and persistence of mammals in these areas are

influenced by the size of the forest patches,

their structural characteristics (e. g. floristic

composition, successional stage), and resource

availability, which varies greatly in seasonal

regions (Stoner & Timm, 2011). Therefore,

seasonality can be linked to behavioral changes

causing mammals to move and actively seek for

resources (Stoner & Timm, 2011). This could

be crucial in small fragments (< 30 ha) where

opportunities to find food, water, and suitable

shelters are limited, especially for large-sized

mammals (Crooks 2002; Crooks et al., 2017;

Morera et al., 2021). For instance, Ocelots

(Leopardus pardalis), Greater Grison (Galictis

vittata) and Collared Peccaries (Pecari taja-

cu) expand their home ranges during the dry

season (Dillon & Kelly, 2008; Haro-Carrión

et al., 2021).

Monitoring mammal presence in frag-

ments within a human-dominated landscape

and understanding seasonal variations is essen-

tial for ensuring connectivity and conservation

of endangered species. Thus, the objectives of

this study were to determine the species rich-

ness of medium and large-sized mammals in

a forest fragment, and to examine if species

diversity and detectability varied according to

seasonality. Considering that rainfall patterns

and tree phenology vary drastically in the stud-

ied location, we hypothesized that seasonality

affected these animals’ diversity and detect-

ability. Since there are more resources available

during the wet season (e. g., fruits, insects, and

water) (Basset et al., 2015; da Silva et al., 2011;

Darosci et al., 2021; Valenzuela & Ceballos

2000), we expected a higher diversity and num-

ber of detections of medium and large-sized

mammals during this season. However, we

expected to find a higher species richness dur-

ing the dry season, when predators and other

large mammals may be detected when traveling

long distances in search of resources.

MATERIALS AND METHODS

Study site: We carried out this study in the

Municipal Forest of Atenas (MFA) Anselmo

Murillo Jiménez, a 26.4 ha premontane moist

forest fragment, located in the province of

Alajuela, in the Central Valley of Costa Rica

(950-1 060 m.a.s.l.; Fig. 1). This area was

previously used for cattle ranching, coffee,

and sugar cane plantations, until the 1930s,

when the Municipality of Atenas purchased the

property to protect it as a source of drinking

water for neighboring communities. Currently,

this forest is part of the Montes del Aguacate

Biological Corridor, and it is co-managed by

the Municipality of Atenas, the community

development association of Alto del Monte and

the Administrative Association for Aqueducts

and Sewers of Plancillo, providing drinking

water to about 1 800 people as well as serving

as a recreational area.

The forest is surrounded mainly by shade

coffee plantations, Highway 3 and riparian

forest (Avalos et al., 2006), and the nearest

large, forested areas (> 2 000 ha) are about 25

km away. The MFA maintains approximately

119 tree species, the most abundant being the

densely flowered Inga or “guabo salado” (Inga

densiflora), the invasive species Syzygium jam-

bos (known as Rose Apple), and secondary

successional species such as the “espavel”

(Anacardium excelsum), the kapok tree (Ceiba

pentandra), and Luehea seemannii (Avalos et

al., 2006; Brassil et al., 2000). Atenas has a

strong seasonality, with the rainy season last-

ing from May to November, and receiving an

averaging annual precipitation of 3 000 mm.

During the dry season (December to April),

seasonal streams in the MFA dry out for 2-3

months. Permanent springs are used as sources

of drinking water.

Data collection: We established 26 camera

trap stations throughout the MFA (Fig. 1). From

August 2021 to April 2022, one camera trap

(Blazevideo SL112) was active at each station

for 12-21 days throughout each season (dry and

wet). We placed 18 cameras near human-made

trails, four cameras off trails (> 20 m away)

and four near streams, aiming to detect species

that avoid trails or that move along streams.

Each site was separated at least 50 m from

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e53245, enero-diciembre 2023 (Publicado Jun. 28, 2023)

the nearest site. Cameras were set to record

ten second videos, with a five second interval

before resetting the trigger, and were active

24 hours a day. We strapped each camera to a

tree at approximately 0.5 m above the ground

and recorded its location with a GPS (Garmin

etrex 10). We also accounted for direct obser-

vations in the field and the use of three camera

traps placed in the canopy for a one month of

exploratory and aleatory sampling. These cam-

eras were placed approximately 12 m above

the ground and were active between March and

April 2022, for a total of 89 camera trap days.

Data analysis: We used digiKam (ver-

sion 7.3.0) (digiKam Developers Team, 2020)

to tag each video with the observed species.

We used ExifTool (version 12.3.2.0) (Harvey,

2022) and the package camtrapr (Niedballa et

al., 2016) in R (version 4.2.0) (R Development

Core Team, 2022) to extract the metadata. Delta

time was set to 120 minutes so that detec-

tions of the same species within a 120-minute

timespan would be considered as one detection.

Considering the small study area (26.4 ha) and

the home range of the medium and large-sized

mammals (from ~ 2 ha for Cuniculus paca to

2 600 ha for Leopardus pardalis), a detection

was not necessarily an independent detection

(Beck-King et al., 1999; Dillon & Kelly, 2008).

Frequency of detection for each species was

calculated as the number of detections per

camera trap day.

We characterized mammal’s species diver-

sity by applying rarefaction and extrapolation

curves using Hill numbers for sample-based

incidence (Chao et al., 2014; Hsieh et al.,

2016). We used species detectability (number

of detections per species per camera trap day)

as the relative incidence when performing

the rarefaction/extrapolation curves. Hill num-

bers represent the effective number of species

within a community (Jost, 2006). The first Hill

number (q0) is equivalent to species richness.

The second one (q1) corresponds to the effec-

tive number of equally common species. The

Fig. 1. Distribution of camera trap stations to survey medium (1-15 kg) and large-sized mammals (> 15 kg) at the Municipal

Forest of Atenas (Alajuela, Costa Rica), between August 2021 and April 2022. Sources: Google Satellite 2022, Centro

Nacional de Alta Tecnología, 2022.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e53245, enero-diciembre 2023 (Publicado Jun. 28, 2023)

third one (q2) includes detectability as well, but

it prioritizes the species with the higher fre-

quency of detections (Chao et al., 2014; Hsieh

et al., 2016).

We applied the same analysis to compare

dry and wet season mammal diversity, consid-

ering the confidence interval (95 %) around

the rarefaction and extrapolation curves. We

compared the curves to test the hypothesis that

seasonality affected diversity. We also com-

pared the average number of detection/camera

trap day between seasons using a Wilcoxon

test. Analyses were performed using R, and the

package iNEXT (Hsieh et al., 2016). Data from

direct observations and the canopy camera

traps were not included in these analyses.

RESULTS

Medium and large-sized mammals of the

MFA: We registered 19 species of medium and

large-sized mammals distributed in 12 families

and five orders (Table 1, Fig. 2.) (Hereafter

we refer to species detected by their common

names). Fifteen species were recorded using

the camera traps in the understory, whereas we

recorded four species by direct observations

and by using camera traps in the canopy (Table

1). Understory camera traps were active for

810 camera days. We registered 557 detections.

The general frequency of detections was 0.688

detections/camera trap day (mean = 0.046 ±

0.073 detections/camera trap day). The average

number of species detected per station was 3.14

± 1.21. Dominant species in the medium and

large-sized mammals assemblage included the

White-nosed Coati (0.254 detections/camera

trap day), the Central American Agouti (0.163

detections/camera trap day), the Lowland Paca

(0.139 detections/camera trap day), the Com-

mon Opossum (0.069 detections/camera trap

day), and the Northern Raccoon (0.059 detec-

tions/camera trap day) (Fig. 3).

In the rarefaction/extrapolation curve, the

effective number of common (q1 = 6) and

dominant species (q2 = 4) reached an asymptote

(Fig. 4A). However, because some uncommon

species have not yet been detected, the species

Fig. 2. Medium (1-15 kg) and large-sized (> 15 kg) mammal species classified as “endangered” in Costa Rica (Ministerio de

Ambiente y Energía, 2017), detected at the Municipal Forest of Atenas (Alajuela, Costa Rica). A. Puma (Puma concolor). B.

Ocelot (Leopardus pardalis). C. Jaguarundi (Herpailurus yagouaroundi). D. Neotropical River Otter (Lontra longicaudis).

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e53245, enero-diciembre 2023 (Publicado Jun. 28, 2023)

Table 1

Medium (1-15 kg) and large mammals (> 15 kg) recorded at the Municipal Forest of Atenas (Alajuela, Costa Rica) between

August 2021 and April 2022

Family Species Common name Guild IUCN MINAE

Mustelidae Eira barbara

(Linnaeus 1758)

Tayra O Least concern Least concern

Galictis vittata

(Schreber 1776)*

Greater Grison CLeast concern Vulnerable

Lontra longicaudis

(Olfers 1818)

Neotropical River Otter CNear threatened Endangered

Felidae Leopardus pardalis

(Linnaeus 1758)

Ocelot C Least concern Endangered

Puma concolor

(Linnaeus 1771)

Puma C Least concern Endangered

Herpailurus yagouaroundi

(É. Geoffroy

Saint-Hilaire 1803)

Jaguarundi C Least concern Endangered

Procyonidae Nasua narica

(Linnaeus 1766)

White-nosed Coati OLeast concern Least concern

Procyon lotor

(Linnaeus 1758)

Northern Raccoon OLeast concern Least concern

Potos flavus

(Schreber 1774)**

Kinkajou O Least concern Least concern

Canidae Canis latrans

(Say 1823)

Coyote O Least concern Least concern

Urocyon cinereoargenteus

(Schreber 1775)

Gray Fox OLeast concern Least concern

Mephitidae Conepatus semistriatus

(Boddaert 1785)

Striped Hog-nosed

Skunk

CLeast concern Least concern

Cuniculidae Cuniculus paca

(Linnaeus 1766)

Lowland Paca HLeast concern Vulnerable

Dasyproctidae Dasyprocta punctata

(Gray 1842)

Central American Agouti HLeast concern Least concern

Erethizonthidae Coendou mexicanus

(Kerr 1792)**

Mexican Hairy Dwarf

Porcupine

HLeast concern Least concern

Dasypodidae Dasypus novemcinctus

(Linnaeus 1758)

Nine-banded Armadillo ILeast concern Least concern

Cebidae Cebus imitator

(Thomas 1903)**

Panamanian White-faced

Capuchin

O Vulnerable Least concern

Myrmecophagidae Tamandua mexicana

(Saussure 1860)

Northern Tamandua ILeast concern Least concern

Didelphidae Didelphis marsupialis

(Linnaeus 1758)

Common Opossum OLeast concern Least concern

Guild: O = Omnivore, Carnivore, H = Herbivore, I = Insectivore. IUCN = International Union for Conservation of Nature’s

Red List of Threatened Species (https://www.iucnredlist.org/). MINAE = National Ministry of Environment and Energy,

Costa Rica (Ministerio de Ambiente y Energía, 2017). *Species detected by observation. **Species detected by camera traps

placed in the canopy (12 m above the ground).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e53245, enero-diciembre 2023 (Publicado Jun. 28, 2023)

richness (q0 = 15) could increase slightly with

additional sampling effort (Fig. 4A). We have

already recorded the following rare species:

Jaguarundi (0.001 detections/camera trap day),

Puma (0.003 detections/camera trap day), Neo-

tropical River Otter (0.004 detections/camera

trap day) and Gray Fox (0.003 detections/cam-

era trap day). Except for the Gray Fox, all of

them are listed as endangered species in Costa

Rica (Table 1).

Seasonality and mammal diversity in

the MFA: The sampling effort during the dry

season consisted of 465 camera days and 295

detections, which accounted for 86.6 % of the

reported mammals in the MFA. The sampling

effort during the wet season was 345 camera

days and 262 mammal detections, with 73.3

% of the mammals detected in this study (Fig.

3). Detection frequencies were higher during

the wet season (Dry season = 0.042 ± 0.080

detections/camera trap day; Wet season = 0.051

± 0.082 detections/camera trap day; W = 417,

P = 0.04).

Seasonality influenced the dominating

species (q2) but did not affect species richness

(q0) or the number of effective common spe-

cies (q1) (Fig. 4). However, if sampling effort is

increased, the overall species richness may also

increase (Fig. 4A). Between the dry and wet

seasons, the identity and prevalence of domi-

nant species varied little (Fig. 3). The most

frequent species recorded during the dry season

was the White-nosed Coati (0.314 detections/

camera trap day), while the most frequent

mammal during the wet season was the Central

American Agouti (0.273 detections/camera trap

day) (Fig. 3). During the dry season, there were

more detections of the Lowland Paca (Fig. 3).

The presence of rare species was the main

difference between wet and dry seasons. The

Striped Hog-nosed Skunk, the Puma, the Neo-

tropical River Otter, and the Jaguarundi were

only observed during the dry season. The Tayra

and the Gray Fox were exclusively detected

during the wet season. Except for the Striped

Hog-nosed Skunk (0.013 detections/camera

trap day), all of them had frequencies lower

than 0.007 detections each camera day (Fig. 3).

Despite these variations, because they are rare

species, they could have been detected in both

seasons with an increased sampling effort.

Fig. 3. Frequency of detections of medium (1-15 kg) and large-sized mammals (> 15 kg) at the Municipal Forest of Atenas

(Alajuela, Costa Rica) during the wet (August-November 2021) and dry season (December 2021 - April 2022).

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e53245, enero-diciembre 2023 (Publicado Jun. 28, 2023)

DISCUSSION

Medium and large-sized mammals of

the MFA: We detected 19 species of medium

and large-sized mammals at the MFA. Com-

pared to similar studies (Botelho et al., 2012;

Michalski & Peres, 2007), we found a relatively

high richness given that this forest fragment

is only 26.4 ha and is primarily surrounded

by agricultural fields and forest fragments.

It is known that there is a tolerance gradient

to forest-cover change, ranging from species

that are highly sensitive to fragmentation to

species that are tolerant or that even benefit

from it (Garmendia et al., 2013). For instance,

Jaguars (Panthera onca) or White-lipped Pec-

caries (Tayassu pecari) are highly sensitive

to habitat loss and prefer continuous forest,

whereas species such as the White-nosed Coati,

the Common Opossum and the Northern Rac-

coon are known to be species that adapt easily

to fragmented environments (Daily et al., 2003;

Garmendia et al., 2013).

Central American Agoutis and Lowland

Pacas are common rodents in fragmented land-

scapes. These animals are considered important

seed predators and dispersers. Agoutis are

primarily diurnal and are commonly found

in human altered landscapes, including agro-

forestry systems such as coffee plantations

Fig. 4. Rarefaction and extrapolation curves for the medium (1-15 kg) and large-sized (> 15 kg) mammals detected at the

Municipal Forest of Atenas (Alajuela, Costa Rica). A. Effective number of species using the whole sampling period and

the three first orders of the Hill numbers. B. Effective number of species, C. Effective number of common species and D.

Effective number of dominant species, all of them comparing wet and dry seasons.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e53245, enero-diciembre 2023 (Publicado Jun. 28, 2023)

(Sánchez-Brenes & Monge, 2021), which is

a common land use at the studied site. Pacas

are nocturnal animals which are considered

vulnerable in Costa Rica, due to heavy hunting

pressure, which although illegal, is a common

practice in the study site. The high number of

detections of these species, while not necessari-

ly related to their abundance, is a good indicator

of prey availability for predators such as coy-

otes, ocelots and pumas (Emsens et al., 2014).

Although small fragments (< 30 ha) are

unlikely to support a viable population of large

predators such as pumas, there is evidence that

they may be crucial to facilitate their mobility,

providing the connectivity that is critical to

sustain viable populations, since these patches

are used as “stepping stones” between larger

protected areas (Dalecky et al., 2002; Garmen-

dia et al., 2013; Sampaio et al., 2010). Popula-

tion viability also depends on habitat matrix

surrounding forest fragments (Garmendia et

al., 2013). Agroforestry systems are known

to increase connectivity across altered land-

scapes, providing habitat for generalist species

and facilitating animal movements, especially

those of forest-dependent species (Perfecto &

Vandermeer, 2008).

The study area is surrounded by a com-

plex matrix that includes residential areas,

secondary forest patches, and agricultural sites

dominated by shade coffee plantations and pas-

turelands. Shade trees in these coffee planta-

tions are capable of natural regeneration (Häger

et al., 2014), which may facilitate connectivity

among forest patches. Previous studies have

shown that larger patches surrounded by sec-

ondary forests and tree crops maintain a higher

number of species of medium and large-sized

terrestrial mammals (Garmendia et al., 2013).

These landscape characteristics may explain

the relatively high mammal richness observed

at the MFA, which can be considered as a

refuge for threatened species and an important

transit area for large mammals.

Seasonality and mammal diversity in

the MFA: As we hypothesized, detectability

was higher during the wet season, when more

resources are available (Basset et al., 2015; da

Silva et al., 2011; Darosci et al., 2021; Valen-

zuela & Ceballos 2000). We also observed a

seasonal turnover in species, which may be

related to food habits and available resources.

For instance, herbivores such as the Central

American Agouti and the Lowland Paca were

detected up to three times more frequently

during the wet season, while the White-nosed

Coati, the Northern Raccoon and the Ocelot

increased their detectability during the dry

season (Fig. 3). This pattern can be related

to changes across space and time that gener-

ates a greater heterogeneity of micro-habitats

inside a small forest fragment (Basset et al.,

2015; Haro-Carrión et al., 2021; Valenzuela

& Macdonald, 2002).

Seasonality and rainfall pattern fluctua-

tions directly affect plant phenology and there-

fore, fruit availability (Wright et al., 1999). The

densely flowered Inga (I. densiflora) is one of

the most common trees at the MFA (Silva-Fors-

berg, 2008). This species fructifies during the

wet season (Koptur, 1983) and is eaten by the

Central American Agouti (Sánchez-Brenes &

Monge, 2021). Along with I. densiflora fruits,

other common trees, like the introduced Rose

Apple (S. jambos), and the shrub Allophylus

occidentalis, whose fructification period also

coincides with the rainy season, may be linked

to the high detectability of herbivores at this

time of year (Avalos et al., 2006; Gargiullo et

al., 2008; Patel et al., 2017). On the other hand,

it is well known that ocelot home-range size

is larger during the dry season when food is

thought to be scarcer (Dillon & Kelly, 2008).

Since a small forest fragment cannot support a

viable feline population, detectability of these

carnivores depends on the use of corridors for

movement (Morera et al., 2021; Salom-Pérez

et al., 2022).

Finally, the White-nosed Coati was one of

the most frequently detected species along the

sampling period, but during the dry season it

represented 50 % of the detections, overshad-

owing other common species such as the Cen-

tral American Agouti and the Lowland Paca

that were dominant during the rainy season. As

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e53245, enero-diciembre 2023 (Publicado Jun. 28, 2023)

this species tends to have a larger home-range

and be more widely dispersed on the area dur-

ing the dry season, this could be associated

to a greater detection in the MFA during this

season, where we were able to record at least

40-50 individuals in the same band (Valenzuela

& Ceballos, 2000; Valenzuela & Macdonald,

2002). This pattern may also be connected to

the coati’s parental care behavior, as females

tend to stay longer in a refuge area caring for

the young, and thus they are less mobile during

the wet season (Valenzuela & Ceballos, 2000).

Implications for water management and

mammal conservation: Although, most of the

mammalian species we registered are classi-

fied as “least concern” by the IUCN red list, in

Costa Rica two of these species are considered

vulnerable (Lowland Paca and Greater Grison)

and four are endangered (Puma, Jaguarundi,

Ocelot and Neotropical River Otter). The pres-

ence of these species in a relatively small

forest fragment (< 30 ha) surrounded by an

agricultural matrix, highlights the importance

of the MFA in the protection and connectivity

of species, contributing to gene flow between

isolated populations. Despite its importance,

the MFA lacks a management plan. Informa-

tion presented in this document contributes to

a better understanding of mammal ecology, the

implementation of proper forest management

strategies, and the strengthening of commu-

nity organization capacities centered around

water preservation.

Like the MFA, in Costa Rica there are

about 1 400 local associations that manage for-

est fragments that provide water to about 25.5

% of the population (1.1 million people) (Insti-

tuto Costarricense de Acueductos y Alcantaril-

lados, 2016). Assessment of the water sources

and monitoring of the area around springs are

therefore essential for efficient water manage-

ment. Due to their sensitivity to human interfer-

ence, medium and large-sized mammals serve

as powerful bioindicators and an important tool

to assess the integrity of those forests (Cove

et al., 2013; Salom-Pérez et al., 2021). It is

critical to examine the relationship between

mammal conservation and water protection.

Conservation efforts should be multipurpose

while also showing the tangible benefits of

protecting forested areas in the provision of

critical environmental services. This study gen-

erates baseline information that shows how

protecting forest fragments to secure drinking

water also helps to restore and maintain stable

animal communities.

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

followed all pertinent ethical and legal proce-

dures and requirements. All financial sources

are fully and clearly stated in the acknowled-

gments section. A signed document has been

filed in the journal archives.

ACKNOWLEDGMENTS

We would like to thank The School for

Field Studies (SFS) for providing key financial

and logistical support to carry out this project.

We thank SFS students for collaborating in data

collection and processing. We are grateful to the

Municipality of Atenas, the Community Devel-

opment Association of Alto del Monte and

the Administrative Association for Aqueducts

and Sewers (ASADA) of Plancillo, for their

logistical support. We also thank Asociación de

Desarrollo Específica para la Conservación del

Ambiente (ADECA) which provided the three

camera traps that were placed in the canopy.

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