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

29, 2024) 1

TERRESTRIAL ECOLOGY

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

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; [email protected].
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; [email protected]
3. Instituto Monteverde, Puntarenas, Costa Rica; [email protected]
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; [email protected]

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

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.

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

gest that herbivores may become dominant in
Herbivory, dispersion, and seed preda- trophic cascades (Borer et al., 2005). Neverthe-
tion by wildlife species are vital processes in less, analyzing the causes of mortality of sapling
forest ecosystems. These processes are a key is vital for understanding the processes that
feature that may significantly influence sapling maintain forest species diversity (Paine & Beck,
establishment, growth, composition, and for- 2007), since, besides herbivory, there are other
est recovery (Feng et al., 2021; Genes & Dirzo, factors restricting the undergrowth sapling
2022; Neuschulz et al., 2016; Norden, 2014; recruitment. These factors include dispersion
Vallejo-Marín et al., 2006). Herbivory is a cru- restriction, environmental filters, biotic and
cial ecological process that contributes to the abiotic factors, and the negative density depen-
individual adaptation of species (Janzen, 1971). dence (Ramírez-Mejía & Mendoza, 2010).
Environmental variables such as temperature Substantial evidence has been encoun-
fluctuations, humidity, and sunlight also can tered, indicating that large herbivores such as
also impact forest recovery, affecting various peccaries significantly contribute to physical
trophic levels (Kuprewicz, 2013; Powell et al., damage and mortality of undergrowth sapling
2015; Yong et al., 2011). due to their rooting and trampling behaviors
Wild mammals play a key role in the while searching for fruits and seeds (Beck,
conservation of neotropical systems (Curran 2005; Queenborough et al., 2012). However,
& Webb, 2000; Hermes et al., 2006), as they peccaries also contribute to the structure and
affect vegetation community diversity (Dirzo diversity of ecosystems and vegetation commu-
& Miranda, 1991; Ickes et al., 2001; Mendoza & nities since they serve as seed dispensers and
Dirzo, 2007; Terborgh & Wright, 1994), through predators (Beck, 2005; Beck et al., 2010; Clark
predation of reproductive and vegetation com- & Clark, 1989; Paine & Beck, 2007; Roldán
ponents, and recovery recruitment (DeMattia & Simonetti, 2001). Thus, relevance of wild
et al., 2004). They facilitate modifications in mammals in herbivory, such as peccaries, has
demography and plants composition in the been acknowledged; disturbing their densities
 Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024) 3

can have detrimental effects on the forest and the direct relation of peccaries with recovery
organisms that depend on litterfall and dendrit- dynamics and their influence in the tropical
ic food webs (Beck, 2005; Reider et al., 2013). forest at La Selva (Clark & Clark, 1989). Most
On the other hand, peccary popula- studies with mammals have been conducted
tions around La Selva Biological Station have in other natural locations. (DeMattia et al.,
declined or become locally extinct (Kuprewicz, 2004; Dirzo & Miranda, 1991; Ickes et al.,
2013). For instance, there are no reports of 2001; Mendoza & Dirzo, 2007; Paine & Beck,
peccary presence in the Nogal Private Wildlife 2007; Roldán & Simonetti, 2001; Terborgh &
Refuge (Nogal) since 2004 by locals or based Wright, 1994).
on monitoring activities conducted by person- Nevertheless, the presence of peccaries and
nel of the reserve. Since 2004, wildlife tracking their relationship with the ecosystem should
has been conducted through field observation not be considered negative a priori because its
and camera traps at the site, without sight- natural distribution plays a crucial role within
ing any peccaries (pers. comm). In regard to the trophic chain or other ecological processes.
La Selva, studies on the historical and cur- Therefore, efforts must be channeled to pre-
rent abundance of peccaries suggest population serve the integrity of mammal communities
growth (Kuprewicz, 2013; Michel et al., 2015; and research the causes of sapling mortality to
Romero et al., 2013). Thus, comprehending the better understand the processes that maintain
role of this species in natural forest recovery is the diversity of forest species. In this way, we
essential, knowing that there is a possibility of can ensure the preservation of fauna and flora
a significant reduction in its population in neo- and the ecological processes that favor the
tropical areas (Beck, 2005; Gongora et al., 2011; recovery and maintenance at La Selva. Hence,
Ontiveros et al., 2021; Reider et al., 2013). This this study aims to measure the impact of pec-
reduction is attributable to rapid deforestation caries (Dycotiles tajacu) on the understory veg-
rates and excessive hunting, which may impact etation of the tropical rainforest at the Nogal-La
in the trophic cascade and the natural recovery Selva Biological Corridor, Costa Rica, in a site
(Reider et al., 2013; Stoner et al., 2007). without peccaries and another location with
Overall, recent studies have shown a ten- their presence.
dency of increasing peccary populations at La
Selva (Romero et al., 2013), leading to the per- MATERIALS AND METHODS
ception that peccaries are the source of nega-
tive impact on forest natural recovery (Michel Study Areas: The study was conducted
& Sherry, 2012). Based on this, a debate about within the biological corridor at Nogal-La
the management of this species at La Selva Selva, Heredia, Northwest Costa Rica. The
has emerged (Romero et al., 2013). Investiga- corridor was developed within the Natura-
tions have focused on direct trophic relations leza y Communidad Project located in Nogal
with one or more species in trophic cascades (Masis-Aguilar, 2019; Ubieta et al., 2009).
(Michel et al., 2014), effects on insectivorous It was created through collaborative efforts
birds and bats (Kalka et al., 2008; Van Bael & involving Chiquita Brands, the local commu-
Brawn, 2005), interaction and perturbation nity, the Sarapiquí local government, Rainfor-
between palms and peccaries (Avalos et al., est Alliance, German Technical Cooperation
2016; Queenborough et al., 2012), use of natu- Agency (GTZ), and the Swiss supermarket
ral and anthropized areas (Osorto-Nuñez & chain Migros (Ubieta et al., 2009). Ecological
Alvarado, 2023) peccaries as important agents restoration actions took place from 2004 to the
that impact litterfall structure, and the abun- present, aiming to connect the Nogal Private
dance of aquatic (anurans), and the terrestrial Wildlife Refuge (Nogal) with La Selva Biologi-
reptiles (Beck et al., 2010; Reider et al., 2013). cal Station (Masis-Aguilar, 2019). La Selva com-
However, few investigations have centered on prises an area of 1 600 hectares. It is classified
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as a wet tropical rainforest (Hartshorn, 1983). reserve adjoins the Río Sucio to the North,
The average daytime temperatures range from forming the Nogal-La Selva ecological corridor
24.7 to 27.1 °C and receives between 3 800–4 (Masis-Aguilar, 2019). There are two areas of 92
000 mm of rainfall annually (Armstrong et hectares of wet rainforest and riverine habitat.
al., 2020; Robinson et al., 2018). The rainfall The reserve is mainly dominated by second-
is slightly lower from January to April (Clark growth forest and dense scrubland reed (Arun-
et al., 2013), while higher rainfall occurs from do donax), and it is 7.8 km away from La Selva
June to July and from November to December (Masis-Aguilar, 2019; Rodríguez-Matamoros
(Salazar-Blanco, 2001). La Selva is connected et al., 2012). The site falls within the tropical
with Braulio Carrillo National Park, featuring rainforest life-zone (Holdridge, 1988) and has
primary rainforest, various stages of second a flat topography and elevations ranging from
growth, and forestry systems (Arroyo-Arce et 40 to 51 m.a.s.l. The climate is predominantly
al., 2013; Oviedo-Pérez, 2008; Raich et al., 2014; warm and humid, with temperatures fluctuat-
Romero et al., 2013). It is located on volcanic ing between 26 °C and 28 °C and an annual
origin soil, which provokes an extreme of high rainfall of 4 000 mm (Masis-Aguilar, 2019)
fertility of lowlands neotropical forests (Clark et (refer to Fig. 1).
al., 2013) Fig. 1).
Meanwhile, Nogal, owned by Chiquita Design of Exclosures and Control Plots:
Brand S.R.L, is in Puerto Viejo district, Sara- La Selva served as the control site with the
piquí, Heredia, North Caribbean region of presence of D. tajacu, while Nogal Refuge
Costa Rica (10°29’23’’ N & 83°56’15” W). This 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) 5

D. tajacu. In both areas, 20 paired plots of 2 between plots to ensure the rain-induced seed
x 2 meters were established, consisting of 10 independence from the same parent tree, which
exclosures treatments and 10 control treat- is significantly reduced after 50 meters (Ceccon
ments with free access. Sampling periods were & Hernández, 2009; Cole et al., 2010; Martínez-
conducted from June 9th, 2021, to January 31st, Ramos & Soto-Castro, 1993) (Fig. 2).
2022. Before implementing the treatments, sev- The paired treatments were separated by
eral sampling areas were created to select the a 5 m distance as shown in Fig. 3. Exclosure
precise site where plots were located. Polygons treatments were surrounded by a galvanized
were selected in Tres Ríos, Las Vegas, and metallic mesh extending up to 1.10 m in height,
Arriera-Zompopa trails. Subsequently, random supported by stakes of flat iron rods with cor-
points were created using QGIS 3.10 software, ners angled at 40 degrees. Plots had an opening
considering 50 meters of separation distance 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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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.

of small mammals and simulate differential a specific range of consumed leaf area: 0-0 %,
extinction of mammals (Galetti et al., 2015; 1-6 %, 6-12 %, 12-25 %, 25-50 % and 50-100 %
Mendoza & Dirzo, 2007). In the control treat- (Dirzo & Dominguez, 1995). The percentage of
ments, plots of 2 m2 were delimited using a few lost leaf area was calculated using the herbivory
PVC stakes placed in the corners. Any damage index as defined by the following formula:
to the trap structures in the exclosure plots was
repaired during each visit. H = (Ci * ni) /N

Vegetation Sampling in Exclosure and Where Ci = mean point of each category.

Control Plots A taxonomy identification was ni = leaflets in the category of damages;
conducted on saplings (0.30–1.50 m height) and N = total leaflets rates.
to determine the most specific taxon pos-
sible (genus and species) based on dendro- Lastly, every leaf underwent visual analysis
logical features. This identification process was to identify browsing signs by mammals, par-
carried out with the assistance of botanical ticularly, if leaves were eaten, or partially or
experts from La Selva OET and Juvenal Valerio entirely removed.
Rodríguez from the Herbarium at Universidad
Nacional. Height measurements were recorded Litter Fall Dynamics: Litter fall was
for all individuals of forest species in both sites monthly measured by using a PVC sampling
(exclosure and control plots), following Orozco frame of 50 cm x 50 cm. Plots were divided into
& Brumer (2002). quadrants of equal size, numbered clockwise
from one to four. Litter fall was collected from
Herbivory: Herbivory levels were mea- three of the four randomly selected quadrants
sured in both treatment sites. The number of in the exclosure and control plots (Reider
leaf sheets of each plant, as well as healthy and et al., 2013), without using random mecha-
damaged leaves were also quantified at both nisms (Chavarria-Bolaños et al., 2012; Reider
sites. The quantification of herbivory occurred et al., 2013; Sousa-Neto et al., 2016; Zhu et al.,
during three periods: the first on July 9th, the 2019). With this method, there is a possibility
second period on September 9th, and the third of repeating two of the four quadrants in the
on November 9th, 2021. The extent of leaf nesting sampling period and selecting a two-
consumption by herbivores was estimated by month quadrant of deposited litter fall. Three
considering six categories of visual damage in samples were collected for each treatment every
 Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024) 7

month (n = September 7th, n = October 7th, differences between treatments (exclosure and
n = November 6th, and n = December 6th) control). Finally, the study conducted an analy-
and placed in paper bags. The collected plant sis of between-sites interactions and experi-
litter material included leaves, part of leaves, mental treatments considering the following
flowers, small fruits, seed pods and branches. variables: rate of herbivory, number of leaf
Branches larger than 20 mm in diameter were sheets, healthy and damaged leaves, sapling
excluded as they were considered woody debris height, and litterfall. This analysis was per-
(Muller-landau & Wright, 2010). Subsequently, formed using an analysis of variance to observe
the organic matter samples were weighed in if there was evidence of contrasts between
the laboratory. The humid litter fall was dried each variable in each study site (La Selva and
for 48 hours at 65 °C, and its dry weight Nogal Reserve).
was measured.
Fauna Activity within Plots: Species cap-
Fauna Activity within Plots: From July tured on video by the camera traps served as
9th to December 9th, 2021, three camera traps an index of activity in the control plots. All
were installed (Bushnell Trophy Cam Hd 12MP graphs and data were analyzed with Python
model: 119739 and 119736) per site. Cameras 3.9 through Google Collaboratory, and geo-
were placed only in the control plots because spatial databases were processed using QGIS
our interest was to observe if medium to large 3.10. software.
size mammals, such as peccaries, tapirs, and
deer, predate or damage saplings. Each camera RESULTS
was mounted on a tree at a height of 50 cm
and set to operate for 24 h d – 1, in video mode Vegetation Abundance and Diversity: A
with a minimum delay of 60 seconds after total of 208 saplings were quantified in the two
detecting an animal within its sensor reach. study sites. In La Selva, 98 individuals represent
Once the cameras were set up, the videos were 33 species from 18 families, and 110 in Nogal
reviewed within the first 24 hours to change Reserve comprising 30 species from 24 families
the SD memory card and to check for species (Table 1). The most numerous families in the
recorded at the study site. Subsequently, the Nogal Reserve were Moraceae (n = 30) and
cameras were reviewed monthly to replace the Rhamnaceae (n = 8), whereas Rubiaceae (n =
SD memory card or check the batteries. For 28), Primulaceae (n = 13), and Moraceae (n =
each camera, the study recorded the follow- 16) were the predominant families in La Selva.
ing data: display, location, camera functioning In terms of species, Ardisia nigropunctata (n =
dates, trap night number, and the number of 13), Sorocea pubivena (n = 10) and Psychotria
videos for each species. The cameras remained B. (n = 10) were more numerous in La Selva,
in each plot for a month, before being moved and Sorocea pubivena (n = 28), and Colubrina
to the next plot until all plots in La Selva and spinosa (n = 8) were the more common species
Nogal were covered. in Nogal. A dead individual (Pentaclethra mac-
roloba) was documented in the Nogal Reserve
Data Analysis: The normality of the vari- during this study. Furthermore, during the
ables was verified using the Shapiro-Wilk test second sampling period in La Selva, a 30-meter
(Shapiro & Wilk, 1965), and the homogeneity tree fell onto an exclosure plot; consequently,
of variance was assessed using the Bartlett test the plot was modified as it was difficult to con-
(Bartlett, 1951). Following these tests, a two- tinue measuring individuals (Table 1).
way analysis of variance (ANOVA) was con-
ducted to confirm the hypothesis of differences Sapling Traits and Litterfall: Strong evi-
between means of two or more groups (Stahle dence was found for differences in herbivo-
& Wold, 1989), to ultimately detect potential ry rates (La Selva only) and the number of
8 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024)

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 8 Sorocea pubivena 10
Symphonia globulifera 5 Psychotria B. 10
Lauraceae 5 Virola sebifera 9
Heisteria macrophylla 5 Psychotria A. 8
Guatteria diospyroides 4 Pentaclethra macroloba 4
Protium pittieri 4 Palicourea chiapensis 3
Virola sebifera 4 Casearia corymbosa 3
Hasseltia floribunda 4 Vochysia guatemalensis 3
Gloeospermum diversipetalum 4 Dilleniaceae 2
Ardisia nigropunctata 3 Nectandra reticulata 2
Annonaceae 2 Brosimum alicastrum 2
Tetragastris panamensis 2 Trophis racemose 2
Sloanea A. 2 Perebea angustifolia 2
Hernandia stenura 2 Syzygium jambos 2
Herrania purpurea 2 Psychotria marginata 2
Pachira aquatica 2 Pentagonia monocaulis 2
Mollinedia pinchotiana 2 solanaceae 2
Sorocea sp. 2 Protium pittieri 1
Virola koschnyi 2 Inga sapindoides 1
Pentagonia sp. 2 Inga sp. 1
Spondias mombin 1 Ocotea cernua 1
Garcinia sp. 1 Rhodostemonodaphne kunthiana 1
Sloanea B. 1 Theobroma cacao 1
Inga sp. 1 Guarea Guidonia 1
Pentaclethra macroloba 1 Eugenia selvana 1
Papilionaceae 1 Piperaceae 1
Piperaceae 1 Pentagonia sp. 1
Rubiaceae 1 Psychotria cyanococca 1
Chione venosa 1 Psychotria C. 1
Salicaceae 1 Meliosma glabrata 1
Lunania sp. 1 Hasseltia floribunda 1
Allophylus psilospermus 1 Paullinia sp. 1
Pouteria sp. 1 Chrysophyllum cainito 1
Simarouba glauca 1 Cestrum schlechtendalii 1
Cuatresia exiguiflora 1
Violaceae 1
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damaged leaves between the exclosure and confirming that these variables depend on the
control groups. Moderate evidence was found treatment and the site. No interaction effect was
for number of leaves (La Selva only) and sapling found for the number of leaves and the number
height (La Selva only) between exclosure and of damaged leaves. Finally, no evidence of dif-
control groups. Additionally, strong evidence ference was found for the number of healthy
was found for the effect of this interaction leaves in the treatment and interaction effect
on herbivory, sapling height and litter fall, (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)

Table 2 DISCUSSION
Statistics summary for variables between exclosure and
control plots at La Selva Biological Station and Nogal Floristic Composition: Peccaries affect
Private Wildlife Refuge, Sarapiquí, Costa Rica.
the recruitment of individuals in the natural
Mean understory regeneration in La Selva, since in
Variable Anova p
Exclosure Control Nogal S. pubivena (consumed by peccaries)
Herbivory rates accounts for 25.45 % of the individuals (Table
La Selva 0.1923 0.1516 0.002 1). In addition, individuals belonging to the
Nogal 0.1516 0.1571 0.773 Moraceae family were more abundant in Nogal
Number of leaf sheets than in La Selva. The species of this family are
La Selva 13.22 20.06 0.01 one of the most important in the diet of pecca-
Nogal 14.69 14.19 0.84 ries according to the meta-analysis conducted
Number of damaged leaves by Beck (2005) and the one found by Osorto-
La Selva 9.17 14.47 0.002 Nuñez et al. (2023) in La Selva, Costa Rica.
Nogal 7.49 10.14 0.007 Defaunated understories tend to have a higher
Healthy leaves
plant density and lower diversity compared
La Selva 3.97 5.56 0.21
to forests with a higher degree of conserva-
Nogal 7.18 4.05 0.10
tion (Dirzo & Miranda, 1991). The defauna-
Sapling height
tion of vertebrates, especially mammals, can
alter those ecological mechanisms allowing
La Selva 63.74 72.55 0.006
the coexistence of thousands of plants in tropi-
Nogal 62.95 61.51 0.632
cal forests (Wright, 2003). This is considered
Litter fall
as an indirect effect for the impoverishment
La Selva 147.48 137.62 0.479
of floristic diversity (Dirzo & Miranda, 1991;
Nogal 185.33 162.06 0.124
Kurten, 2013; Leigh et al., 1993; Terborgh, 1992;
Terborgh & Wright, 1994).
Fauna Activity within Plots: A total of The absence of wild mammals may nega-
152 trap days were recorded in the control tively affect certain plant species while ben-
plots. Omnivorous and frugivorous mammals efiting others (Kurten, 2013). In areas where
were the predominant guilds captured on video peccary populations have decreased, there is
in both study sites; for instance, 120 captures a top-down effect on (Michel et al., 2014)
involving 16 species were documented in La plant-animal interaction. This is because seed
Selva; while in Nogal Reserve, 88 captures predators and herbivores limit the abundance
of 13 species were recorded. Dycotiles tajacu of slow- to moderate-growing forest species
accounted for only 12 % (n = 14) of the video that form the upper layers in a natural for-
captures in the control plots, ranking as the est (Camargo-Sanabria et al., 2015; Kurten,
fourth most frequently recorded species. In 2013; Wright, 2003). However, there are other
contrast, agouti was the species with the most biotic and abiotic factors that hinder regenera-
captures on video (37 %, n = 44). No traces, tion recruitment such as temperature variation,
direct sighting reports, or fecal evidence of T. humidity, light, wind, water availability, soil
bairdii and O. virginianus were found near the properties, diseases, energy or nutrient reserves,
plots in La Selva during the period of this study. and storms, among others (Augspurger, 1984;
In Nogal Reserve, the most frequently captured Bullied et al., 2012; Cano & Stevenson, 2009;
species on video was Nasua narica (n = 36), Kéfi et al., 2012; Leigh et al., 1993; Osunkoya et
representing 42 % of the captures, followed al., 1993; Paine & Beck, 2007; Ramírez-Mejía &
by Cuniculus paca which accounted for 17 % Mendoza, 2010). If there is a selective behavior
(n = 15). No captures of Dycotiles tajacu were in the consumption of vegetation by herbivo-
reported on this site. rous mammals, its absence could allow certain
 Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e53238, enero-diciembre 2024 (Publicado Feb. 29, 2024) 11

species to be freed from herbivory, favoring the reproduction in relation to the less damaged
dominance of these species in the ecosystem plants (Hawkes & Sullivan, 2001; Maschinski &
(Roldán & Simonetti, 2001); this could be the Whitham, 1989; McNaughton, 1983).
case of S. pubivena in Nogal. In the short term, mammals alter the con-
dition of plant species that are part of their diet
Herbivory: Sounders of peccaries, when and influence other components of food webs,
they are abundant, such as those found in La which can affect insect herbivores (Firn et al.,
Selva, influence natural regeneration through 2017; Vandegehuchte et al., 2017) by reducing
herbivory, given the differences observed with the amount of food resources (Teichman et al.,
Nogal (Fig. 2 A), herbivore exclusions reduce 2013). Thus, the exclusion of mammals could
foliar herbivory (Pearson et al., 2003), and pop- lead to positive effects for herbivorous insects
ulations of medium and large mammals have (Vandegehuchte et al., 2017). However, in the
substantial effects on regeneration (Belovsky & long term, mammals such as peccaries can
Slade, 2000; Medinaceli et al., 2004). Accord- modify the vegetation composition, includ-
ing to the camera records in the plots, the ing the relative abundance of preferred plants,
most abundant species was D. punctata, which which in turn may affect herbivore insects asso-
coincides with (Kuprewicz, 2013), who in turn ciated with these plants (Beck, 2005; Huntly,
points to the peccary as the other most abun- 1991; Kurten, 2013; Rumiz, 2010).
dant mammal species. This combination of In the context of biological corridors, the
agouti and peccary may contribute to higher analysis of ecological dynamics becomes highly
herbivory rates in La Selva compared to Nogal.
relevant given the particularities of a landscape
According to the sampling data, the most abun-
that is confronted with changing conditions
dant species in Nogal was N. narica, which
at a faster rate compared to areas of absolute
primarily feeds on fruits and insects and, there-
protection, such as national parks. This study
fore, it would not have a significant impact on
conducted in two key areas of the Nogal - La
herbivory rates (Valenzuela, 1998). It is worth
Selva biological corridor shows the complexity
noting that the agouti is a predator and seed
that governs plant-animal interactions, taking
disperser, but it does not eat or trample vegeta-
tion (Kuprewicz, 2013). the D. tajacu’s influence on the regeneration
The growth in height of saplings contrasts of species as an example at the sapling level
with the studies by Osunkoya et al. (1993) and in lowland tropical rainforests. Therefore, the
Wahungu et al. (2002). These researchers found design, implementation, and evaluation of the
that seedlings in excluded areas grew at a faster effectiveness of biological corridors should con-
rate than those in unprotected areas. However, sider this type of complexities in order to maxi-
this could be related to the total number of mize the success of biodiversity conservation,
leaves and branches since there was evidence of ecosystem services to human communities, and
disparity in the number of leaf laminae biased the flow of ecological processes through func-
toward the open access plots and is a covariate tional and structural connectivity, the latter
related to seedling size (Arteaga, 2006; Gross- being analyzed from interactions such as those
nickle & MacDonald, 2018; Seiwa & Kikuzawa, presented in this study.
1991). The peccary trampling could affect the
height of saplings, but the differences in height Ethical statement: The authors declare
in La Selva compared to Nogal (Fig 2B) indicate that they all agree with this publication and
that there was a compensation as a positive made significant contributions; that there is no
plant response to herbivory, since the dam- conflict of interest of any kind; and that we fol-
aged seedlings alter their resource allocation, lowed all pertinent ethical and legal procedures
physiology, and phenology in order to reduce and requirements. All financial sources are
the impact of the damage in their growth and 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 Bartlett, M. S. (1951). The effect of standardization on a χ2
approximation in factor analysis. Biometrika, 38(3-4),
the journal archives. 337–344.

Beck, H. (2005). Seed predation and dispersal by peccaries

ACKNOWLEDGMENTS throughout the neotropics and its consequences: a
review and synthesis. In P. M. Forget, J. E. Lambert,
To German Academic Exchange Service P. E. Hulme, & S. B. Vander Wall (Eds.), Seed fate:
predation, dispersal and seedling establishment (pp.
(DAAD) for the scholarship granted to M. H. 77–115). CABI.
Osorto-Nuñez, to study the Master’s degree in
Wildlife Conservation and Management. To Beck, H., Thebpanya, P., & Filiaggi, M. (2010). Do Neo-
tropical peccary species (Tayassuidae) function as
Universidad Nacional de Costa Rica (UNA) for ecosystem engineers for anurans? Journal of Tropical
the funding granted for materials and logistics. Ecology, 26(4), 407–414.
To the Organization for Tropical Studies (OTS) Belovsky, G. E., & Slade, J. B. (2000). Insect herbivory
and Glaxo Centro America Fellowships (Glaxo- accelerates nutrient cycling and increases plant pro-
SmithKline) for the funds to carry out the field duction. Proceedings of the National Academy of Scien-
ces, 97(26), 14412–14417. https://doi.org/10.1073/
work at the La Selva Biological Station (scholar- PNAS.250483797
ship code 1417). To Chiquita Brands Costa Rica
Borer, E. T., Seabloom, E. W., Shurin, J. B., Anderson, K.
LTDA for their support and approval to con- E., Blanchette, C. A., Broitman, B., Cooper, S. D., &
duct my study at Nogal Private Wildlife Refuge. Halpern, B. S. (2005). What determines the strength
To Idea Wild by the research equipment pro- of a trophic cascade? Ecology, 86(2), 528–537.
vided. To O. Vargas for assistance in identifying Bullied, J. W., van Acker, R. C., & Bullock, P. R. (2012).
plant species. To three anonymous reviewers Review: Microsite characteristics influencing weed
that contributed to improve the manuscript. seedling recruitment and implications for recruitment
modeling. Canadian Journal of Plant Science, 92(4),
627–650.

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