Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024)

Impact of Collared Peccaries Dycotiles tajacu (Artiodactyla: Tayassuidae)

on understory vegetation in the tropical rainforest of

the Nogal-La Selva Biological Corridor, Costa Rica

Marco Herminio Osorto-Nuñez1*; https://orcid.org/0000-0003-2061-4950

Luis Diego Alfaro Alvarado2; https://orcid.org/0000-0001-9534-1948

Federico A. Chinchilla Romero3; https://orcid.org/0000-0001-5473-4307

Flávio H. Guimarães Rodrigues4; https://orcid.org/0000-0002-4797-0085

1. Instituto Internacional de Conservación y Manejo de Vida Silvestre. Universidad Nacional. Facultad de ciencias de la

Tierra y el Mar. Campus Omar Dengo, 40101, Provincia de Heredia, Heredia, Costa Rica; marco.osorto.nunez@est.una.

ac.cr (*Correspondence)

2. Facultad de Ciencias de la Tierra y el Mar, Escuela de Ciencias Ambientales, Universidad Nacional. Calle 9 y av. 1,

40101, Provincia de Heredia. Heredia, Costa Rica; luis.alfaro.alvarado@una.cr

3. Instituto Monteverde, Puntarenas, Costa Rica; federicoeap@gmail.com

4. Departamento de Genética, Ecologia e Evolução, Instituto de Ciências Biológicas, Universidade Federal de Minas

Gerais, Avenida Antônio Carlos 6627, Belo Horizonte, Minas Gerais, Brazil; rodriguesfhg@gmail.com

Received 15-VIII-2023. Corrected 20-II-2024. Accepted 22-II-2024.

ABSTRACT

Introduction: Evidence suggests that herbivores, such as peccaries, shape vegetation structure and diversity

through predation, trampling, dispersal, and rooting behavior.

Objective: To evaluate the impact of peccaries (Dycotiles tajacu) on the understory vegetation of the tropical

rainforest in the Nogal-La Selva Local Biological Corridor, Costa Rica, comparing a site with the absence of pec-

caries to another with the presence of these animals.

Methodology: From June to November 2021, 20 experimental exclusions and 20 free access plots, each measur-

ing 2 m2 were used to quantify herbivory, the number of leaf blades, damaged leaves, healthy leaves, sapling

height, and fallen biomass at both sites.

Results: A higher sapling density was found in the Nogal Reserve, but a lower sapling diversity, while in La Selva

there was a higher sapling diversity, but a lower density of seedlings. Herbivory and sapling height in La Selva

exceeded those in Nogal. The exclusion of peccaries reduced seedling damage but did not affect the dynamics

of fallen biomass.

Conclusion: For the design, implementation, and evaluation of the effectiveness of biological corridors, it is

crucial to consider plant-animal interactions to enhance the flow of ecological processes through functional and

structural connectivity, analyzed from interactions such as those presented in this paper.

Key words: Biological Station La Selva; herbivory; sapling height; natural regeneration; Nogal Reserve; Pecari

tajacu; recruitment.

https://doi.org/ 10.15517/rev.biol.trop..v72i1.53238

TERRESTRIAL ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024)

INTRODUCTION

Herbivory, dispersion, and seed preda-

tion by wildlife species are vital processes in

forest ecosystems. These processes are a key

feature that may significantly influence sapling

establishment, growth, composition, and for-

est recovery (Feng et al., 2021; Genes & Dirzo,

2022; Neuschulz et al., 2016; Norden, 2014;

Vallejo-Marín et al., 2006). Herbivory is a cru-

cial ecological process that contributes to the

individual adaptation of species (Janzen, 1971).

Environmental variables such as temperature

fluctuations, humidity, and sunlight also can

also impact forest recovery, affecting various

trophic levels (Kuprewicz, 2013; Powell et al.,

2015; Yong et al., 2011).

Wild mammals play a key role in the

conservation of neotropical systems (Curran

& Webb, 2000; Hermes et al., 2006), as they

affect vegetation community diversity (Dirzo

& Miranda, 1991; Ickes et al., 2001; Mendoza &

Dirzo, 2007; Terborgh & Wright, 1994), through

predation of reproductive and vegetation com-

ponents, and recovery recruitment (DeMattia

et al., 2004). They facilitate modifications in

demography and plants composition in the

forest (Romero et al., 2016). Some findings sug-

gest that herbivores may become dominant in

trophic cascades (Borer et al., 2005). Neverthe-

less, analyzing the causes of mortality of sapling

is vital for understanding the processes that

maintain forest species diversity (Paine & Beck,

2007), since, besides herbivory, there are other

factors restricting the undergrowth sapling

recruitment. These factors include dispersion

restriction, environmental filters, biotic and

abiotic factors, and the negative density depen-

dence (Ramírez-Mejía & Mendoza, 2010).

Substantial evidence has been encoun-

tered, indicating that large herbivores such as

peccaries significantly contribute to physical

damage and mortality of undergrowth sapling

due to their rooting and trampling behaviors

while searching for fruits and seeds (Beck,

2005; Queenborough et al., 2012). However,

peccaries also contribute to the structure and

diversity of ecosystems and vegetation commu-

nities since they serve as seed dispensers and

predators (Beck, 2005; Beck et al., 2010; Clark

& Clark, 1989; Paine & Beck, 2007; Roldán

& Simonetti, 2001). Thus, relevance of wild

mammals in herbivory, such as peccaries, has

been acknowledged; disturbing their densities

RESUMEN

Impacto del pecarí de collar, Dycotiles tajacu (Artiodactyla: Tayassuidae) en la vegetación del sotobosque

del bosque tropical húmedo del Corredor Biológico Local Nogal-La Selva, Costa Rica

Introducción: Existe evidencia que herbívoros, como los saínos, dan forma a la estructura y diversidad de la

vegetación a través del comportamiento de depredación, pisoteo, dispersión y enraizamiento.

Objetivo: Evaluar el impacto de los saínos (Dycotiles tajacu) en la vegetación del sotobosque del bosque tropical

húmedo en el Corredor Biológico Local Nogal-La Selva, Costa Rica, en un sitio con ausencia y en otro con pre-

sencia de saínos.

Métodos: De junio a noviembre de 2021 se utilizaron 20 exclusiones experimentales y 20 parcelas de acceso libre

de 2 m2, se cuantifico la herbivoría, número de láminas foliares, hojas dañadas, hojas sanas, altura de brinzales y

biomasa caída en ambos sitios.

Resultados: Se encontró una mayor densidad de brinzales en Reserva Nogal pero una menor diversidad, contra-

rio en La Selva donde se encontró una mayor diversidad de brinzales, pero una menor densidad de plántulas. La

herbivoría y la altura de brinzales en La Selva fue mayor que en Nogal. La exclusión de los saínos disminuyó el

daño a las plántulas, pero no afectó la dinámica de la biomasa caída.

Conclusión: Es necesario contemplar para el diseño, implementación y evaluación de la efectividad de corredores

biológicos, las interacciones planta-animal, para potencializar el flujo de procesos ecológicos mediante la conecti-

vidad funcional y estructural, analizada a partir de interacciones como las presentadas en este trabajo.

Palabras claves: Estación Biológica La Selva; herbivoría; altura de brinzales; regeneración natural; Reserva Nogal;

Pecari tajacu; reclutamiento.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024)

can have detrimental effects on the forest and

organisms that depend on litterfall and dendrit-

ic food webs (Beck, 2005; Reider et al., 2013).

On the other hand, peccary popula-

tions around La Selva Biological Station have

declined or become locally extinct (Kuprewicz,

2013). For instance, there are no reports of

peccary presence in the Nogal Private Wildlife

Refuge (Nogal) since 2004 by locals or based

on monitoring activities conducted by person-

nel of the reserve. Since 2004, wildlife tracking

has been conducted through field observation

and camera traps at the site, without sight-

ing any peccaries (pers. comm). In regard to

La Selva, studies on the historical and cur-

rent abundance of peccaries suggest population

growth (Kuprewicz, 2013; Michel et al., 2015;

Romero et al., 2013). Thus, comprehending the

role of this species in natural forest recovery is

essential, knowing that there is a possibility of

a significant reduction in its population in neo-

tropical areas (Beck, 2005; Gongora et al., 2011;

Ontiveros et al., 2021; Reider et al., 2013). This

reduction is attributable to rapid deforestation

rates and excessive hunting, which may impact

in the trophic cascade and the natural recovery

(Reider et al., 2013; Stoner et al., 2007).

Overall, recent studies have shown a ten-

dency of increasing peccary populations at La

Selva (Romero et al., 2013), leading to the per-

ception that peccaries are the source of nega-

tive impact on forest natural recovery (Michel

& Sherry, 2012). Based on this, a debate about

the management of this species at La Selva

has emerged (Romero et al., 2013). Investiga-

tions have focused on direct trophic relations

with one or more species in trophic cascades

(Michel et al., 2014), effects on insectivorous

birds and bats (Kalka et al., 2008; Van Bael &

Brawn, 2005), interaction and perturbation

between palms and peccaries (Avalos et al.,

2016; Queenborough et al., 2012), use of natu-

ral and anthropized areas (Osorto-Nuñez &

Alvarado, 2023) peccaries as important agents

that impact litterfall structure, and the abun-

dance of aquatic (anurans), and the terrestrial

reptiles (Beck et al., 2010; Reider et al., 2013).

However, few investigations have centered on

the direct relation of peccaries with recovery

dynamics and their influence in the tropical

forest at La Selva (Clark & Clark, 1989). Most

studies with mammals have been conducted

in other natural locations. (DeMattia et al.,

2004; Dirzo & Miranda, 1991; Ickes et al.,

2001; Mendoza & Dirzo, 2007; Paine & Beck,

2007; Roldán & Simonetti, 2001; Terborgh &

Wright, 1994).

Nevertheless, the presence of peccaries and

their relationship with the ecosystem should

not be considered negative a priori because its

natural distribution plays a crucial role within

the trophic chain or other ecological processes.

Therefore, efforts must be channeled to pre-

serve the integrity of mammal communities

and research the causes of sapling mortality to

better understand the processes that maintain

the diversity of forest species. In this way, we

can ensure the preservation of fauna and flora

and the ecological processes that favor the

recovery and maintenance at La Selva. Hence,

this study aims to measure the impact of pec-

caries (Dycotiles tajacu) on the understory veg-

etation of the tropical rainforest at the Nogal-La

Selva Biological Corridor, Costa Rica, in a site

without peccaries and another location with

their presence.

MATERIALS AND METHODS

Study Areas: The study was conducted

within the biological corridor at Nogal-La

Selva, Heredia, Northwest Costa Rica. The

corridor was developed within the Natura-

leza y Communidad Project located in Nogal

(Masis-Aguilar, 2019; Ubieta et al., 2009).

It was created through collaborative efforts

involving Chiquita Brands, the local commu-

nity, the Sarapiquí local government, Rainfor-

est Alliance, German Technical Cooperation

Agency (GTZ), and the Swiss supermarket

chain Migros (Ubieta et al., 2009). Ecological

restoration actions took place from 2004 to the

present, aiming to connect the Nogal Private

Wildlife Refuge (Nogal) with La Selva Biologi-

cal Station (Masis-Aguilar, 2019). La Selva com-

prises an area of 1 600 hectares. It is classified

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as a wet tropical rainforest (Hartshorn, 1983).

The average daytime temperatures range from

24.7 to 27.1 °C and receives between 3 800–4

000 mm of rainfall annually (Armstrong et

al., 2020; Robinson et al., 2018). The rainfall

is slightly lower from January to April (Clark

et al., 2013), while higher rainfall occurs from

June to July and from November to December

(Salazar-Blanco, 2001). La Selva is connected

with Braulio Carrillo National Park, featuring

primary rainforest, various stages of second

growth, and forestry systems (Arroyo-Arce et

al., 2013; Oviedo-Pérez, 2008; Raich et al., 2014;

Romero et al., 2013). It is located on volcanic

origin soil, which provokes an extreme of high

fertility of lowlands neotropical forests (Clark et

al., 2013) Fig. 1).

Meanwhile, Nogal, owned by Chiquita

Brand S.R.L, is in Puerto Viejo district, Sara-

piquí, Heredia, North Caribbean region of

Costa Rica (10°29’23’’ N & 83°56’15” W). This

reserve adjoins the Río Sucio to the North,

forming the Nogal-La Selva ecological corridor

(Masis-Aguilar, 2019). There are two areas of 92

hectares of wet rainforest and riverine habitat.

The reserve is mainly dominated by second-

growth forest and dense scrubland reed (Arun-

do donax), and it is 7.8 km away from La Selva

(Masis-Aguilar, 2019; Rodríguez-Matamoros

et al., 2012). The site falls within the tropical

rainforest life-zone (Holdridge, 1988) and has

a flat topography and elevations ranging from

40 to 51 m.a.s.l. The climate is predominantly

warm and humid, with temperatures fluctuat-

ing between 26 °C and 28 °C and an annual

rainfall of 4 000 mm (Masis-Aguilar, 2019)

(refer to Fig. 1).

Design of Exclosures and Control Plots:

La Selva served as the control site with the

presence of D. tajacu, while Nogal Refuge

was designated as the exclusion site without

Fig. 1. Location of experimental plots (exclosure and control) at La Selva Biological Station and Nogal Private Wildlife

Refuge.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024)

D. tajacu. In both areas, 20 paired plots of 2

x 2 meters were established, consisting of 10

exclosures treatments and 10 control treat-

ments with free access. Sampling periods were

conducted from June 9th, 2021, to January 31st,

2022. Before implementing the treatments, sev-

eral sampling areas were created to select the

precise site where plots were located. Polygons

were selected in Tres Ríos, Las Vegas, and

Arriera-Zompopa trails. Subsequently, random

points were created using QGIS 3.10 software,

considering 50 meters of separation distance

between plots to ensure the rain-induced seed

independence from the same parent tree, which

is significantly reduced after 50 meters (Ceccon

& Hernández, 2009; Cole et al., 2010; Martínez-

Ramos & Soto-Castro, 1993) (Fig. 2).

The paired treatments were separated by

a 5 m distance as shown in Fig. 3. Exclosure

treatments were surrounded by a galvanized

metallic mesh extending up to 1.10 m in height,

supported by stakes of flat iron rods with cor-

ners angled at 40 degrees. Plots had an opening

of 15 x 10 cm of wall base to allow the entrance

Fig. 2. Random points and points selected for exclosures and control plots in A. Nogal Private Wildlife Reserve and B. La

Selva Biological Station.

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of small mammals and simulate differential

extinction of mammals (Galetti et al., 2015;

Mendoza & Dirzo, 2007). In the control treat-

ments, plots of 2 m2 were delimited using a few

PVC stakes placed in the corners. Any damage

to the trap structures in the exclosure plots was

repaired during each visit.

Vegetation Sampling in Exclosure and

Control Plots A taxonomy identification was

conducted on saplings (0.30–1.50 m height)

to determine the most specific taxon pos-

sible (genus and species) based on dendro-

logical features. This identification process was

carried out with the assistance of botanical

experts from La Selva OET and Juvenal Valerio

Rodríguez from the Herbarium at Universidad

Nacional. Height measurements were recorded

for all individuals of forest species in both sites

(exclosure and control plots), following Orozco

& Brumer (2002).

Herbivory: Herbivory levels were mea-

sured in both treatment sites. The number of

leaf sheets of each plant, as well as healthy and

damaged leaves were also quantified at both

sites. The quantification of herbivory occurred

during three periods: the first on July 9th, the

second period on September 9th, and the third

on November 9th, 2021. The extent of leaf

consumption by herbivores was estimated by

considering six categories of visual damage in

a specific range of consumed leaf area: 0-0 %,

1-6 %, 6-12 %, 12-25 %, 25-50 % and 50-100 %

(Dirzo & Dominguez, 1995). The percentage of

lost leaf area was calculated using the herbivory

index as defined by the following formula:

H = (Ci * ni) /N

Where Ci = mean point of each category.

ni = leaflets in the category of damages;

and N = total leaflets rates.

Lastly, every leaf underwent visual analysis

to identify browsing signs by mammals, par-

ticularly, if leaves were eaten, or partially or

entirely removed.

Litter Fall Dynamics: Litter fall was

monthly measured by using a PVC sampling

frame of 50 cm x 50 cm. Plots were divided into

quadrants of equal size, numbered clockwise

from one to four. Litter fall was collected from

three of the four randomly selected quadrants

in the exclosure and control plots (Reider

et al., 2013), without using random mecha-

nisms (Chavarria-Bolaños et al., 2012; Reider

et al., 2013; Sousa-Neto et al., 2016; Zhu et al.,

2019). With this method, there is a possibility

of repeating two of the four quadrants in the

nesting sampling period and selecting a two-

month quadrant of deposited litter fall. Three

samples were collected for each treatment every

Fig. 3. Diagram of experimental plots conducted at La Selva Biological Station and Nogal Private Wildlife Reserve. A.

Exclosure plots allowing the entrance of only small mammals. B. Control plots located 5 m away from the experimental

exclosure, with unrestricted access.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024)

month (n = September 7th, n = October 7th,

n = November 6th, and n = December 6th)

and placed in paper bags. The collected plant

litter material included leaves, part of leaves,

flowers, small fruits, seed pods and branches.

Branches larger than 20 mm in diameter were

excluded as they were considered woody debris

(Muller-landau & Wright, 2010). Subsequently,

the organic matter samples were weighed in

the laboratory. The humid litter fall was dried

for 48 hours at 65 °C, and its dry weight

was measured.

Fauna Activity within Plots: From July

9th to December 9th, 2021, three camera traps

were installed (Bushnell Trophy Cam Hd 12MP

model: 119739 and 119736) per site. Cameras

were placed only in the control plots because

our interest was to observe if medium to large

size mammals, such as peccaries, tapirs, and

deer, predate or damage saplings. Each camera

was mounted on a tree at a height of 50 cm

and set to operate for 24 h d – 1, in video mode

with a minimum delay of 60 seconds after

detecting an animal within its sensor reach.

Once the cameras were set up, the videos were

reviewed within the first 24 hours to change

the SD memory card and to check for species

recorded at the study site. Subsequently, the

cameras were reviewed monthly to replace the

SD memory card or check the batteries. For

each camera, the study recorded the follow-

ing data: display, location, camera functioning

dates, trap night number, and the number of

videos for each species. The cameras remained

in each plot for a month, before being moved

to the next plot until all plots in La Selva and

Nogal were covered.

Data Analysis: The normality of the vari-

ables was verified using the Shapiro-Wilk test

(Shapiro & Wilk, 1965), and the homogeneity

of variance was assessed using the Bartlett test

(Bartlett, 1951). Following these tests, a two-

way analysis of variance (ANOVA) was con-

ducted to confirm the hypothesis of differences

between means of two or more groups (Stahle

& Wold, 1989), to ultimately detect potential

differences between treatments (exclosure and

control). Finally, the study conducted an analy-

sis of between-sites interactions and experi-

mental treatments considering the following

variables: rate of herbivory, number of leaf

sheets, healthy and damaged leaves, sapling

height, and litterfall. This analysis was per-

formed using an analysis of variance to observe

if there was evidence of contrasts between

each variable in each study site (La Selva and

Nogal Reserve).

Fauna Activity within Plots: Species cap-

tured on video by the camera traps served as

an index of activity in the control plots. All

graphs and data were analyzed with Python

3.9 through Google Collaboratory, and geo-

spatial databases were processed using QGIS

3.10. software.

RESULTS

Vegetation Abundance and Diversity: A

total of 208 saplings were quantified in the two

study sites. In La Selva, 98 individuals represent

33 species from 18 families, and 110 in Nogal

Reserve comprising 30 species from 24 families

(Table 1). The most numerous families in the

Nogal Reserve were Moraceae (n = 30) and

Rhamnaceae (n = 8), whereas Rubiaceae (n =

28), Primulaceae (n = 13), and Moraceae (n =

16) were the predominant families in La Selva.

In terms of species, Ardisia nigropunctata (n =

13), Sorocea pubivena (n = 10) and Psychotria

B. (n = 10) were more numerous in La Selva,

and Sorocea pubivena (n = 28), and Colubrina

spinosa (n = 8) were the more common species

in Nogal. A dead individual (Pentaclethra mac-

roloba) was documented in the Nogal Reserve

during this study. Furthermore, during the

second sampling period in La Selva, a 30-meter

tree fell onto an exclosure plot; consequently,

the plot was modified as it was difficult to con-

tinue measuring individuals (Table 1).

Sapling Traits and Litterfall: Strong evi-

dence was found for differences in herbivo-

ry rates (La Selva only) and the number of

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Table 1

Frequency of plant species found in the plots (exclosure and control) in Nogal Private Wildlife Refuge and La Selva Biological

Station.

Study sites

Nogal Private Wildlife Refuge La Selva Biological Station

Species Count Species Count

Sorocea pubivena 28 Ardisia nigropunctata 13

Colubrina spinosa 8Sorocea pubivena 10

Symphonia globulifera 5Psychotria B.10

Lauraceae 5 Virola sebifera 9

Heisteria macrophylla 5Psychotria A.8

Guatteria diospyroides 4Pentaclethra macroloba 4

Protium pittieri 4Palicourea chiapensis 3

Virola sebifera 4Casearia corymbosa 3

Hasseltia floribunda 4Vochysia guatemalensis 3

Gloeospermum diversipetalum 4Dilleniaceae 2

Ardisia nigropunctata 3Nectandra reticulata 2

Annonaceae 2Brosimum alicastrum 2

Tetragastris panamensis 2Trophis racemose 2

Sloanea A.2Perebea angustifolia 2

Hernandia stenura 2Syzygium jambos 2

Herrania purpurea 2Psychotria marginata 2

Pachira aquatica 2Pentagonia monocaulis 2

Mollinedia pinchotiana 2solanaceae 2

Sorocea sp. 2Protium pittieri 1

Virola koschnyi 2Inga sapindoides 1

Pentagonia sp. 2Inga sp. 1

Spondias mombin 1Ocotea cernua 1

Garcinia sp. 1Rhodostemonodaphne kunthiana 1

Sloanea B. 1Theobroma cacao 1

Inga sp.1Guarea Guidonia 1

Pentaclethra macroloba 1Eugenia selvana 1

Papilionaceae 1Piperaceae 1

Piperaceae 1Pentagonia sp.1

Rubiaceae 1Psychotria cyanococca 1

Chione venosa 1Psychotria C. 1

Salicaceae 1Meliosma glabrata 1

Lunania sp. 1Hasseltia floribunda 1

Allophylus psilospermus 1Paullinia sp. 1

Pouteria sp. 1Chrysophyllum cainito 1

Simarouba glauca 1Cestrum schlechtendalii 1

Cuatresia exiguiflora 1

Violaceae 1

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024)

damaged leaves between the exclosure and

control groups. Moderate evidence was found

for number of leaves (La Selva only) and sapling

height (La Selva only) between exclosure and

control groups. Additionally, strong evidence

was found for the effect of this interaction

on herbivory, sapling height and litter fall,

confirming that these variables depend on the

treatment and the site. No interaction effect was

found for the number of leaves and the number

of damaged leaves. Finally, no evidence of dif-

ference was found for the number of healthy

leaves in the treatment and interaction effect

(Table 2, Fig. 4).

Fig. 4. Effects of the two-way interaction between treatment factors (exclosure and control) and the study sites La Selva

Biological Station and Nogal Private Wildlife Refuge, Costa Rica, for the following variables: A. Herbivory, B. Number of leaf

sheets, C. Number of healthy leaves, D. Sapling height, E. Number of damaged leaves and F. Litterfall.

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024)

Fauna Activity within Plots: A total of

152 trap days were recorded in the control

plots. Omnivorous and frugivorous mammals

were the predominant guilds captured on video

in both study sites; for instance, 120 captures

involving 16 species were documented in La

Selva; while in Nogal Reserve, 88 captures

of 13 species were recorded. Dycotiles tajacu

accounted for only 12 % (n = 14) of the video

captures in the control plots, ranking as the

fourth most frequently recorded species. In

contrast, agouti was the species with the most

captures on video (37 %, n = 44). No traces,

direct sighting reports, or fecal evidence of T.

bairdii and O. virginianus were found near the

plots in La Selva during the period of this study.

In Nogal Reserve, the most frequently captured

species on video was Nasua narica (n = 36),

representing 42 % of the captures, followed

by Cuniculus paca which accounted for 17 %

(n = 15). No captures of Dycotiles tajacu were

reported on this site.

DISCUSSION

Floristic Composition: Peccaries affect

the recruitment of individuals in the natural

understory regeneration in La Selva, since in

Nogal S. pubivena (consumed by peccaries)

accounts for 25.45 % of the individuals (Table

1). In addition, individuals belonging to the

Moraceae family were more abundant in Nogal

than in La Selva. The species of this family are

one of the most important in the diet of pecca-

ries according to the meta-analysis conducted

by Beck (2005) and the one found by Osorto-

Nuñez et al. (2023) in La Selva, Costa Rica.

Defaunated understories tend to have a higher

plant density and lower diversity compared

to forests with a higher degree of conserva-

tion (Dirzo & Miranda, 1991). The defauna-

tion of vertebrates, especially mammals, can

alter those ecological mechanisms allowing

the coexistence of thousands of plants in tropi-

cal forests (Wright, 2003). This is considered

as an indirect effect for the impoverishment

of floristic diversity (Dirzo & Miranda, 1991;

Kurten, 2013; Leigh et al., 1993; Terborgh, 1992;

Terborgh & Wright, 1994).

The absence of wild mammals may nega-

tively affect certain plant species while ben-

efiting others (Kurten, 2013). In areas where

peccary populations have decreased, there is

a top-down effect on (Michel et al., 2014)

plant-animal interaction. This is because seed

predators and herbivores limit the abundance

of slow- to moderate-growing forest species

that form the upper layers in a natural for-

est (Camargo-Sanabria et al., 2015; Kurten,

2013; Wright, 2003). However, there are other

biotic and abiotic factors that hinder regenera-

tion recruitment such as temperature variation,

humidity, light, wind, water availability, soil

properties, diseases, energy or nutrient reserves,

and storms, among others (Augspurger, 1984;

Bullied et al., 2012; Cano & Stevenson, 2009;

Kéfi et al., 2012; Leigh et al., 1993; Osunkoya et

al., 1993; Paine & Beck, 2007; Ramírez-Mejía &

Mendoza, 2010). If there is a selective behavior

in the consumption of vegetation by herbivo-

rous mammals, its absence could allow certain

Table 2

Statistics summary for variables between exclosure and

control plots at La Selva Biological Station and Nogal

Private Wildlife Refuge, Sarapiquí, Costa Rica.

Vari abl e Mean Anova p

Exclosure Control

Herbivory rates

La Selva 0.1923 0.1516 0.002

Nogal 0.1516 0.1571 0.773

Number of leaf sheets

La Selva 13.22 20.06 0.01

Nogal 14.69 14.19 0.84

Number of damaged leaves

La Selva 9.17 14.47 0.002

Nogal 7.49 10.14 0.007

Healthy leaves

La Selva 3.97 5.56 0.21

Nogal 7.18 4.05 0.10

Sapling height

La Selva 63.74 72.55 0.006

Nogal 62.95 61.51 0.632

Litter fall

La Selva 147.48 137.62 0.479

Nogal 185.33 162.06 0.124

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024)

species to be freed from herbivory, favoring the

dominance of these species in the ecosystem

(Roldán & Simonetti, 2001); this could be the

case of S. pubivena in Nogal.

Herbivory: Sounders of peccaries, when

they are abundant, such as those found in La

Selva, influence natural regeneration through

herbivory, given the differences observed with

Nogal (Fig. 2 A), herbivore exclusions reduce

foliar herbivory (Pearson et al., 2003), and pop-

ulations of medium and large mammals have

substantial effects on regeneration (Belovsky &

Slade, 2000; Medinaceli et al., 2004). Accord-

ing to the camera records in the plots, the

most abundant species was D. punctata, which

coincides with (Kuprewicz, 2013), who in turn

points to the peccary as the other most abun-

dant mammal species. This combination of

agouti and peccary may contribute to higher

herbivory rates in La Selva compared to Nogal.

According to the sampling data, the most abun-

dant species in Nogal was N. narica, which

primarily feeds on fruits and insects and, there-

fore, it would not have a significant impact on

herbivory rates (Valenzuela, 1998). It is worth

noting that the agouti is a predator and seed

disperser, but it does not eat or trample vegeta-

tion (Kuprewicz, 2013).

The growth in height of saplings contrasts

with the studies by Osunkoya et al. (1993) and

Wahungu et al. (2002). These researchers found

that seedlings in excluded areas grew at a faster

rate than those in unprotected areas. However,

this could be related to the total number of

leaves and branches since there was evidence of

disparity in the number of leaf laminae biased

toward the open access plots and is a covariate

related to seedling size (Arteaga, 2006; Gross-

nickle & MacDonald, 2018; Seiwa & Kikuzawa,

1991). The peccary trampling could affect the

height of saplings, but the differences in height

in La Selva compared to Nogal (Fig 2B) indicate

that there was a compensation as a positive

plant response to herbivory, since the dam-

aged seedlings alter their resource allocation,

physiology, and phenology in order to reduce

the impact of the damage in their growth and

reproduction in relation to the less damaged

plants (Hawkes & Sullivan, 2001; Maschinski &

Whitham, 1989; McNaughton, 1983).

In the short term, mammals alter the con-

dition of plant species that are part of their diet

and influence other components of food webs,

which can affect insect herbivores (Firn et al.,

2017; Vandegehuchte et al., 2017) by reducing

the amount of food resources (Teichman et al.,

2013). Thus, the exclusion of mammals could

lead to positive effects for herbivorous insects

(Vandegehuchte et al., 2017). However, in the

long term, mammals such as peccaries can

modify the vegetation composition, includ-

ing the relative abundance of preferred plants,

which in turn may affect herbivore insects asso-

ciated with these plants (Beck, 2005; Huntly,

1991; Kurten, 2013; Rumiz, 2010).

In the context of biological corridors, the

analysis of ecological dynamics becomes highly

relevant given the particularities of a landscape

that is confronted with changing conditions

at a faster rate compared to areas of absolute

protection, such as national parks. This study

conducted in two key areas of the Nogal - La

Selva biological corridor shows the complexity

that governs plant-animal interactions, taking

the D. tajacu’s influence on the regeneration

of species as an example at the sapling level

in lowland tropical rainforests. Therefore, the

design, implementation, and evaluation of the

effectiveness of biological corridors should con-

sider this type of complexities in order to maxi-

mize the success of biodiversity conservation,

ecosystem services to human communities, and

the flow of ecological processes through func-

tional and structural connectivity, the latter

being analyzed from interactions such as those

presented in this study.

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

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024)

section. A signed document has been filed in

the journal archives.

ACKNOWLEDGMENTS

To German Academic Exchange Service

(DAAD) for the scholarship granted to M. H.

Osorto-Nuñez, to study the Master’s degree in

Wildlife Conservation and Management. To

Universidad Nacional de Costa Rica (UNA) for

the funding granted for materials and logistics.

To the Organization for Tropical Studies (OTS)

and Glaxo Centro America Fellowships (Glaxo-

SmithKline) for the funds to carry out the field

work at the La Selva Biological Station (scholar-

ship code 1417). To Chiquita Brands Costa Rica

LTDA for their support and approval to con-

duct my study at Nogal Private Wildlife Refuge.

To Idea Wild by the research equipment pro-

vided. To O. Vargas for assistance in identifying

plant species. To three anonymous reviewers

that contributed to improve the manuscript.

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