Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Community-based identification of ecosystem services

in Ramsar wetlands: a socio-ecological network approach

Vladimir González-Gamboa1,2*; https://orcid.org/0009-0000-5283-8989

1. Programa Estado de la Nación, CONARE, San José, Costa Rica; vgonzalez@estadonacion.or.cr,

vladimir.gonzalezgamboa@ucr.ac.cr (*Correspondence)

2. Programa de Posgrado en Gestión Integrada de Áreas Costeras Tropicales, Universidad de Costa Rica, San José,

Costa Rica.

Received 08-VIII-2024. Corrected 17-XII-2024. Accepted 24-I-2025.

ABSTRACT

Introduction: Ramsar wetlands are recognized worldwide for the ecosystem services they provide to society,

for example they are source of food and water. However, the study of their socio-ecological complexity is often

not focused on the local communities that directly interact with the ecosystem. However, the participation and

involvement of local actors with wetland management authorities are key factors in achieving sustainability.

Objective: To study the socio-ecological relevance of Ramsar wetland ecosystem services at the community level

in Costa Rica.

Methods: Information was collected from one participatory workshop, 27 interviews with community leaders,

and a survey administered to 744 households. These methods were applied in 14 communities within seven

Ramsar wetlands where interaction between the ecosystem and the communities is significant.

Results: Provisioning ecosystem services were most frequently mentioned by local leaders and households. The

main product quantified was fish harvesting, except for the Térraba-Sierpe wetland where mollusks were the

most collected wetland product. The social network of wetland actors shows that public actors are the prin-

cipal actors interacting positively with wetland authorities. The socio-ecological network of wetland-collected

products identified central products. The fish families that are relevant simultaneously for many wetlands are

robalo (Centropomidae), roncador (Haemulidae), and pargos (Lutjanidae). Additionally, the Caribe Noreste,

Térraba-Sierpe, and Gandoca Manzanillo wetlands are those where more diversity and quantity of products were

identified.

Conclusions: The two socio-ecological networks highlight that two wetlands have conditions for high manage-

ment tension, four have moderate tension, and only one has low tension for management.

Key words: ecosystem services; socio-ecological network; natural resources management; local communities;

Ramsar wetlands.

RESUMEN

Identificación comunitaria de servicios ecosistémicos en humedales Ramsar:

un enfoque socioecológico hacia el manejo sostenible

Introducción: Los humedales Ramsar son reconocidos mundialmente por los servicios ecosistémicos que pro-

porcionan a la sociedad como por ejemplo son fuente de alimento, y agua. Sin embargo, el estudio de su com-

plejidad socio-ecológica a menudo no se centra en las comunidades locales que interactúan directamente con el

ecosistema. La participación e implicación de los actores locales con las autoridades de gestión de humedales son

factores clave para lograr la sostenibilidad.

https://doi.org/10.15517/rev.biol.trop..v73iS1.63638

SUPPLEMENT

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Wetlands General Insights: Wetlands are

commonly understood as complex ecologi-

cal communities and their associated environ-

ments, forming integral components of natural

ecosystems (Mitsch et al., 2009). Despite cove-

ring only 5 % to 7 % of the Earth’s land surface,

wetlands play a disproportionately significant

role in global ecological functions (Mitsch

et al., 2009). Wetlands contribute to various

aspects of human well-being, including the

provision of resources for human consump-

tion, maintenance of climate stability, enhan-

cement of water quality, coastal protection,

flood mitigation, and preservation of wildlife

habitats (Mitsch et al., 2009; Straton, 2006;

Van der Valk, 2012). Consequently, wetlands,

as ecological entities, not only play a crucial

role in maintaining ecosystem connectivity but

also deliver indispensable ecosystem services

to society (Straton, 2006). The social and eco-

nomic implications of these services contribute

significantly to the overall well-being of human

populations (Constanza et al., 1997; De Groot

et al., 2006; Turner et al., 2000). Hence, the

goal of the present study is to identify ecosys-

tem services from which the local population

benefits in Ramsar wetlands in Costa Rica. A

detailed explanation is provided in the final

part of this section.

Ecosystem services obtained from wet-

lands are exclusively related to hydrological

processes. Due to this nature, there are challen-

ges in assessing the benefits derived from these

processes. They often align with non-market

values, making it difficult to explicitly quantify

ecosystem services (Barbier, 2011). Thus, to

sustain the long-term benefits of wetlands, it is

urgent that society moves towards their protec-

tion and, when possible, rational use (Ramsar

Convention Secretariat, 2010).

In 1971, the Convention on Wetlands

(Ramsar, Iran) took place, creating an inter-

governmental treaty for the conservation and

wise use of all wetlands, promoting local and

national actions, as well as international coo-

peration. According to the Ramsar Convention

Secretariat, as of August 24, 2023, 172 countries

have subscribed to the Ramsar Convention, and

Costa Rica has been a signatory since April 27,

1992 (Ramsar Convention Secretariat, 2023a).

The Ramsar List is the world’s largest net-

work of wetland protected areas with interna-

tional importance. According to this list, there

are over 2 400 Ramsar sites covering more than

2.5 million square kilometers. A wetland can be

designated as Ramsar if it is a site containing

representative rare or unique wetland types

or the site is of international importance for

Objetivo: Estudiar la relevancia socio-ecológica de los servicios ecosistémicos de los humedales Ramsar a nivel

comunitario en Costa Rica.

Métodos: Se recopiló información de un taller participativo, 27 entrevistas con líderes comunitarios y una encues-

ta aplicada a 744 hogares. Estos métodos se aplicaron en 14 comunidades dentro de siete humedales Ramsar

donde la interacción entre el ecosistema y las comunidades es significativa.

Resultados: Los servicios ecosistémicos de aprovisionamiento fueron mencionados con mayor frecuencia por

líderes locales y hogares. El producto principal cuantificado fue la pesca, excepto en el humedal Térraba-Sierpe

donde los moluscos fueron el producto más recolectado. La red social de actores de los humedales muestra que

los actores públicos son los principales actores que interactúan con las autoridades del humedal. La red socio-

ecológica de productos recolectados en el humedal identificó productos centrales. Las familias de peces que son

relevantes simultáneamente para muchos humedales son robalo (Centropotamidae), roncador (Haemulidae) y

pargos (Lutjanidae). Además, los humedales Caribe Noreste, Térraba-Sierpe y Gandoca Manzanillo son aquellos

donde se identificó mayor diversidad y cantidad de productos.

Conclusiones: Las dos redes socio-ecológicas resaltan que dos humedales tienen condiciones de alta tensión de

gestión, cuatro tienen tensión moderada y solo uno tiene baja tensión para la gestión.

Palabras clave: servicios ecosistémicos; red socio-ecológica; gestión de recursos naturales; comunidades locales;

humedales Ramsar.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

conserving biological diversity (Ramsar Con-

vention Secretariat, 2016). Costa Rica alone

has 12 Ramsar sites, covering 569 742 hectares

(Ramsar Convention Secretariat, 2023b).

Types of ecosystem services: One example

of an ecosystem service classification system is

provided by Barbier (2011). This system cate-

gorizes services into three groups: provisioning,

regulating, and cultural. Provisioning services

include goods, which are products obtained

from ecosystems, such as crops and water. A

common example is food for the population,

which can be provided by fisheries. Regulating

services encompass many processes that regu-

late the environment, such as soil retention that

helps to reduce erosion control. Cultural bene-

fits encompass spiritual, religious, and aesthetic

values that contribute to human well-being.

Another detailed classification of wetland

ecosystem services has been proposed based

on their relationship with human development

(De Groot et al., 2006). This classification cate-

gorizes services into four types: provisioning,

regulating, cultural, and supporting. Suppor-

ting services are those that underpin the pro-

vision of all other ecosystem services. They

include processes such as nutrient cycling and

biodiversity, which is essential for maintaining

habitat for resident and migratory species.

Complexity of wetland socio-ecolo-

gical interactions: There are many studies

that analyze the ecosystem services provided

by wetlands to society. However, due to the

complexity involved in analyzing interactions

between the biophysical and socio-economic

systems at the community level, the local social

dimension tends to be overlooked. An approach

to viewing and analyzing the complexity of the

biophysical and ecological system interacting

with the socio-economic system is through the

complex systems framework (Reyes et al., 2013;

Straton, 2006).

According to Reyes et al. (2013), identi-

fying and measuring ecosystem services and

changes in them is particularly difficult due to

the interaction of social and ecological factors.

Limburg et al. (2002) argue that the socio-eco-

nomic system indicates the human preferences

at a specific moment; this occurs within a

socio-economic system where tastes, preferen-

ces, and needs are changing (Chopra & Adhika-

ri, 2004; Limburg et al., 2002; Straton, 2006).

Therefore, it is advisable to analyze the socio-

economic and environmental systems jointly,

considering them as part of a co-evolutionary

process, where the economic system may exert

significant pressure on the environment (Tur-

ner et al., 2000). To pursue this, identifying

the interconnections between ecosystems and

their beneficiaries is the first step (Gunderson

& Holling, 2002).

Mapping ecosystem services: In map-

ping ecosystem services, Burkhard et al. (2013)

highlight that it is fundamental for decision-

making processes as it provides information

about the relevance of conservation investment

and land use planning since these require

robust data. Moreover, the identification and

geospatial location of ecosystem services are

prerequisites for future economic valuation.

Therefore, it is necessary to understand where

and what wetland goods and services benefit

the population (Burkhard et al., 2013).

There are ecosystem services mapping

tools such as InVEST, ARIES, and GUMBO

that are mostly used to map regional and natio-

nal scales. In this context, regulating services

mapping dominates, integrating biophysical

information, and less frequently, social values

(Crossman et al., 2013). For instance, Crossman

et al. (2013) indicate that the sub-national level

is the most common scale for mapping ecosys-

tem services. The use of empirical data, inclu-

ding social actors, to map local socio-ecological

relations with ecosystem services is rare (Page-

lla & Sinclair, 2014).

Ecosystem services can be identified and

prioritized with stakeholders, including the

local population. For this, Cárcamo et al. (2014)

propose a network approach. The relevance

of identifying ecosystem services based on

the population is that people’s perceptions are

correlated with the activities and livelihoods

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

directly associated with the wetland. Thus, the

socio-ecological system can be better unders-

tood (Cárcamo et al., 2014). Furthermore, this

approach incorporates the knowledge of the

local population and creates local legitimacy

for ecosystem services maps, thus helping to

enhance any policy intervention to improve

wetland management (Pagella & Sinclair, 2014).

Identifying ecosystem services via the

population focuses on the social demand side,

instead of the capacity of the system to provide

those services (Castro et al., 2013). The relevan-

ce of this approach is that it shows the societal

dependence on ecological life. This is highly

relevant since the status of ecosystem services

is not only influenced by ecosystem properties

but also by societal needs. Hence, there is a

need to record and spatially localize the actual

ecosystem services collected and consumed by

the population (Castro et al., 2013). At the local

scale, this approach requires fine-resolution

datasets, with the advantage that local farmers

and fishermen identify their natural context

(Pagella & Sinclair, 2014).

Therefore, the present study contributes

to the identification and mapping of ecosystem

services at the local level. More specifically,

the goal of the present study is to empirically

and spatially identify ecosystem services that

the local population directly demands from

Ramsar wetlands in Costa Rica. For this, we

approach the socio-ecological system similarly

to Cárcamo et al. (2014) and as Barnes et al.

(2019) who analyze the socio-ecological net-

work. Barnes et al. (2019) defined this network

as the linked structures between social actors

and ecological resources. The socio-ecological

network takes place as long as it models the

social and ecological relationships at some

level, independently if the discussion is focused

on one of both dimensions (Sayles et al., 2019).

The case of Costa Rica’s wetlands: Accor-

ding to Xu et al. (2020), no scientific studies

on wetland ecosystem services in Costa Rica at

the national level have been documented in the

international literature. This reflects that the

identification and quantification of wetlands

in Costa Rica have been major challenges.

On this concern, we could identify several,

mostly local and regional, research studies done

on Costa Rica’s wetlands. The actual number

of hectares of registered wetlands is 268 703

(Sistema Nacional de Áreas de Conservación

[SINAC], 2023). An examination of the socio-

economic contributions of Palo Verde National

Park estimates how the park quantitatively and

qualitatively benefits society at local, regional,

national, and international levels (Moreno et

al., 2010). Another study focusing on the live-

lihoods of communities around Medio Queso

wetland analyzes interactions by concentrating

on different types of capital (Gutiérrez & Siles,

2008). Alvarado (2008) carried out a compre-

hensive assessment to estimate interactions

among economic, social, ecological, cultural,

and institutional components using a multicri-

teria analysis for Gandoca-Manzanillo wetland.

In the Térraba-Sierpe wetland, several

studies have been conducted. For instance,

the BIOMARCC-SINAC-GIZ Project, (2012)

estimates the capture and fixation of carbon

through different ecosystems. Sanchez et al.

(2013) identified and valued ecosystem services

including tourism, artisanal fishing, “piangua”

(a type of mollusk) extraction; in the producti-

ve part: oil palm (Elaeis guineensis), pineapple

(Ananas comosus), rice (Oryza sativa), dual-

purpose livestock, and forestry. A similar

approach was used by Barton (1995), who iden-

tified and quantified a wide range of ecosystem

services. Kocian et al. (2010), relying on litera-

ture for quantification, given the lack of local

data on ecosystem services. They identified

ecosystem services such as flood protection,

natural protection against droughts, nutrient

cycling, biodiversity, providing “piangua”, and

aesthetic value (Kocian et al., 2010). Reyes et

al. (2004) assessed the ecosystem services of

the Térraba-Sierpe wetland, focusing primarily

on three services: fishing, “piangua” extraction,

and tourism (including hospitality). Aguilar &

Moulaert (2011) proposed the use of multicri-

teria valuation as a form of participatory valua-

tion, political influence, and conflict resolution

for use in the Térraba-Sierpe wetland. Through

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

consultation with experts and key stakeholders,

scenarios were constructed.

Aguilar & Moulaert (2011) conducted a

study to estimate the loss in ecosystem services

due to the conflict in Isla Portillos in the Nor-

theast Caribbean Wetland. This study identifies

ecosystem services such as food, drinking water,

fuel, plant fiber, among others. In another study,

Reyes et al. (2013) identified and valued ecosys-

tem services associated with Las Baulas Marine

National Park (PNMLB). This valuation was

based on the local tourism cluster: lodging,

food, transportation, and tour operation. Pro-

yecto Humedales SINAC-PNUD-GEF (2017)

pursued an economic valuation of ecosystem

services of Ramsar wetlands for Costa Rica

using the method known as transfer value per

unit. This method uses secondary information

from literature review to identify ecosystem

services. The authors complement this method

with expert interviews and remote sensing to

assign land use categories. This study has been

the most relevant found in the literature about

identification and valuation of ecosystem servi-

ces in Costa Rica. However, as Proyecto Hume-

dales SINAC-PNUD-GEF (2017) underlines,

each wetland exhibits unique characteristics

distinct from others; hence, the optimal ecosys-

tem services assessment process involves deve-

loping methodologies that can adjust to this

heterogeneity. From this perspective, we pro-

pose to approach the communities that interact

directly with Ramsar wetlands; thus, we hope

to capture the heterogeneity of socio-ecological

relations and related ecosystem services in the

Ramsar wetlands of Costa Rica.

MATERIALS AND METHODS

We propose a methodological framework

to inventory Ramsar wetlands’ ecosystem ser-

vices and to understand them in light of social

actors. This is important since within the Ram-

sar Secretary, social actors analysis and ecosys-

tem services inventory are two steps that must

be carried out to provide a robust context for

policymakers and for further economic valua-

tion exercises (De Groot et al., 2006).

Thus, we propose an approach that con-

siders the activities of the communities that

interact directly with Ramsar wetlands to iden-

tify ecosystem services. For this, in-depth inter-

views were conducted with communal leaders.

The research instrument focused on questions

that helped to identify the usages and relation-

ships that the local population establishes with

the wetlands. To identify social actors related

to the wetlands, the results of a workshop were

used, which took place with the National Sys-

tem of Conservation Areas (SINAC) officials

in charge of each wetland. The identification

of ecosystem services was conducted in seven

Ramsar wetlands in Costa Rica: Las Baulas,

Palo Verde, Caño Negro, Maquenque, Caribe-

Noreste, Gandoca-Manzanillo, and the Térra-

ba-Sierpe. These wetlands are shown in Fig. 1.

The analysis is limited to these seven wetlands

because these are the ones where there is direct

interaction between the communities and the

wetland. In other Ramsar wetlands, such as Isla

del Coco and Turberas, there are no communi-

ties directly interacting with the wetlands.

Once the ecosystem services were identi-

fied, an analysis was conducted to determine

and quantify which ecosystem services were

most relevant. This part used information from

a survey that took place between 2015 and

2016 as part of the Wetland Project (Conser-

vación, Uso Sostenible de la Biodiversidad y

Mantenimiento de los Servicios de los Ecosiste-

mas de Humedales Protegidos de Importancia

Internacional), which was coordinated bet-

ween SINAC-PNUD-GEF. Besides the empiri-

cal research, a secondary literature review was

also conducted. The objective of this review

was to identify all ecosystem services reported

in previous studies. Generally, this methodolo-

gical approach integrates fieldwork using social

research methods such as interviews, a survey,

and a participatory workshop, with network

analysis. It aims to provide a nuanced unders-

tanding of the socio-ecological interactions

surrounding wetlands that are evidenced in the

ecosystem services.

Mapping social actors: The objective

of this part is to determine the interactions

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

between social actors and wetlands. For this,

we aim to identify stakeholders and socio-eco-

logical interactions. The final analysis provides

valuable insights into the dynamics between

social actors and wetlands, aiding in the deve-

lopment of targeted management strategies and

fostering a greater understanding of the shared

responsibilities and benefits associated with

wetland conservation. To analyze interactions,

network analysis proves to be a valuable tool

for determining the influence and degree of

relationships among actors (Bödin & Prell,

2011). In addition, network analysis is crucial

for interpreting how natural resources are co-

managed, as the actor network reveals commu-

nication patterns regarding a resource (Crona

& Bödin, 2006).

The analysis of results was based on infor-

mation gathered in the workshop: “Planning

the Wetlands Project at the Conservation Area

Level.” This workshop took place from 17th to

19th March 2015 and included the participa-

tion of the officials in charge of each Ramsar

wetland. There are nine Ramsar wetlands of

Costa Rica included in this actors’ map; howe-

ver, our analysis focuses on only seven out of

the nine wetlands. Isla del Coco and Turberas

wetlands were excluded from this study for the

reasons previously mentioned.

The result of this part is a network where

wetlands interact with social actors. A rela-

tion is registered if the rangers expressed that

there is collaboration and positive interaction

between the social actor and the wetland. To

present the results, two network statistics were

used: indegree centrality and outdegree centra-

lity. Indegree network centrality measures the

number of incoming relations, indicating the

prominence of an actor as a collaborator with

the wetland management processes. The inde-

gree centrality is estimated as follows: , where i

is any actor and represents incoming relation-

ships of those linked to a wetland, as they are

nominated as contacts (Brandes & Erlebach,

2005). Outdegree, on the other hand, quantifies

the number of outgoing relations, highlighting

Fig. 1. Costa Rica’s Ramsar wetlands included in the study. Source: self-elaborated based on SINAC (2023).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

a wetland’s significance in creating intercon-

nections. The outdegree centrality is estimated

as follows: , where i is any actor and represents

those linked to i, namely, his neighborhood

when, i, wetland authorities collaborated with

a social actor, their outgoing relations (Brandes

& Erlebach, 2005).

By considering these two network sta-

tistics, we gain insights into the centrality of

actors within the socio-ecological network,

shedding light on their relevance and influen-

ce in the overall structure. The use of these

metrics allows for a comprehensive analysis of

the network’s central actors and their respective

roles. Additionally, we present the Ego-network

of each wetland. An Ego-network, as defined

by Crossley et al. (2015), is the network rela-

ted to any node, i. The actors related directly

to this node are known as alters. The ego-net

can be limited to the representation of an ego

and its alters or can be expanded to include the

relations between alters (Crossley et al., 2015).

Here, we limit the ego-net to the representation

of ego and its alters. This network technique

aims to better represent the individual wetland

immediate social actor’s circle.

The statistical analysis and network mode-

ling were conducted using the R statistical

software (R Core Team, 2023). Specifically, the

library igraph (Csardi & Nepusz, 2006). For

network visualization, the specialized software

Gephi (Bastian et al., 2009) and Cytoscape

(Shannon et al., 2003) were used.

Interviews with communal leaders: We

interviewed communal leaders because these

local actors exhibit many characteristics of an

opinion leader, as outlined by Rogers (2003).

Opinion leaders have a positive influence on

others and enhance the diffusion of ideas,

information, and innovations within the net-

work (Burt, 1999; Rogers, 2003; Valente &

Davis, 1999). Therefore, interviews with com-

munity leaders provided an initial appreciation

and helped to list ecosystem services perceived

by influential actors of communities in each

wetland. The communities were chosen using

the criteria to prioritize the most diverse and

strongest possible relations with the wetland.

To pursue this, we based our selection on the

workshop results, personal conversations with

the SINAC officials, and the literature review.

The experience of the SINAC rangers, the most

cited communities, and wetland uses were cen-

tral. Thus, the goal of the community selection

was to identify as many ecosystem services as

possible. The key respondents were selected

based on initial approximations in the field,

recommendations of the SINAC officials, and

other local institutions. These actors must be

active within the community and have lived

long enough to have perceived any change in

the interactions between the community and

the wetland over time.

Although each wetland has a different

number of villages in the surroundings, not all

have the same relation with it. As observed in

Table 1, there are 143 communities within the

influence area of the wetlands. Thus, we wor-

ked with 10 % of all the communities, but with

those who were identified as most important

in a two-way relationship between the wetland

and the population, as explained.

Finally, two communal leaders of each

community were interviewed, except for the

case of Las Cubas, in Caño Negro wetland,

where only one interview was conducted due

Table 1

Number of communities within the wetland influence area.

Wetland Number

of communities

Selected

communities

Las Baulas 21 -Mata Palo

-Villareal

Palo Verde 17 -El Rosario

-Bagatsí

Caño Negro 15 -Caño Negro

-Las Cubas

Maquenque 30 -Golfito

-Boca San Carlos

Caribe Noreste 12 -Tortuguero

-San Francisco

Gandoca-

Manzanillo 10 -Manzanillo

-Gandoca

Térraba-Sierpe 38 -Ajuntaderas

-Coronado

Total 143 14

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

to the rejection of a communal leader. In gene-

ral, four communal leaders per wetland were

interviewed, and three in the Caño Negro wet-

land, resulting in a total of 27 interviews. The

interviews were conducted between March and

May 2015.

The survey: The survey was conducted

with the aim of capturing detailed informa-

tion on the interactions between the local

population and the wetland, which is revealed

in the provisioning ecosystem services. The

results are a detailed inventory of this type of

ecosystem services, quantified for each Ramsar

wetland. More specifically, the provisioning

ecosystem services are the most dominant and

diverse ecosystem services identified by the

communal leaders. In this type of services, the

socio-ecological relationships are more evident

in those activities related to the extraction of

a biological wetland product. Thus, our aim is

to describe the socio-ecological network that

results from this ecosystem service. The objec-

tive is mainly to observe the relations between

each wetland and the different types of wetland

products collected by households, for instance

fish, mollusks, and plants.

Therefore, through a survey, it was possible

to determine the significant extent to which

community residents are connected to wetlands

through ecosystem services. The survey was

conducted to determine the quantities in which

different ecosystem services contribute directly

or indirectly to the household. The survey was

conducted from September to October 2015,

in communities that inhabit and interact with

the wetlands.

Based on the previous results of the work-

shop, literature review, and key informant

interviews, along with time and financial limi-

tations, we decided to apply the survey in the

two communities that were selected for the

interviews with communal leaders in each

Ramsar wetland. These communities have a

key and close role in the interaction with the

wetland, which was our main criterion for the

identification of ecosystem services. To guaran-

tee a full mapping of the ecosystem services and

the socio-ecological interactions in the com-

munities, the research instrument was applied

to all household heads. Thus, we interviewed

all (census) household heads within all the

selected communities; we did not sample the

community (Frank in Carrington et al., 2005).

The total population found in the villages diffe-

red from the total that the National Institute of

Statistics and Censuses (INEC) had from the

2011 census (Instituto Nacional de Estadísti-

ca y Censos [INEC], 2011). The population

dynamics show an increase in some cases and a

decrease in others, as shown in Table 2.

A total of 14 communities were surveyed,

representing approximately 10 % of the 143

communities in the wetland influence zone.

The total number of surveys conducted was

744, with 140 rejections. Table 2 shows the com-

munities that were surveyed. The total surveys

applied per wetland are as follows: Las Baulas

with 159, Palo Verde with 113, Caño Negro

with 70, Maquenque with 56, Caribe Noreste

with 163, Gandoca-Manzanillo with 77, and

Térraba-Sierpe with 106. Finally, 85 % of the

households were surveyed, and the remaining

15 % rejected to participate in the study.

The information was collected using recall

questions about the type of products, species,

quantity, and prices (if sold) of the products

collected from the wetland over the last 12

months. The use of recall questions has some

inconveniences, as the respondents were not

always able to recall the exact details of the pro-

ducts. For example, the respondents were able

to identify the type of fish they caught and pro-

vide an approximate number of fishes, but the

details were not enough to establish the species.

Hence, we only determined the family of the

fish based on the common name provided by

the respondent. For any future economic valua-

tion, several factors must be considered; the-

refore, it is necessary to explore the valuation

method that fits best. Products from the sea,

such as fish, are mostly sold locally. Conversely,

freshwater fish are mostly consumed by house-

holds, and if sold, they are also sold locally to

neighbors, small supermarkets, or local bars.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

To analyze the data, the socio-ecological

network was examined as defined by Sayles et

al. (2019) and Barnes et al. (2019). The nodes

represent either a wetland or a wetland pro-

duct. The wetland represents the aggregation

of the household head’s responses. Since it was

possible to estimate the quantity collected from

each product by the households, the network

is a weighted graph, where the tie represents

the quantity of the products (Kleinberg, 1999).

Therefore, to identify the most central nodes,

network weighted centrality measures must be

used. To estimate the degree centrality in the

weighted network, two statistics were applied:

the hub score and the authority score developed

by Kleinberg (1999).

The authority score represents how valua-

ble the wetland product is for the households

as they collect (demand) from the wetland.

The hub score indicates how crucial a Ramsar

wetland is by providing products to many hou-

seholds, playing a key role in offering valuable

natural resources for the local population. In

the graph, , the non-negative authority weight

is , and the non-negative hub weight is . Thus,

the value for the authority statistic is: , and for

the hub statistic is: , where is a network node.

Therefore, a node with value of 1 will have the

highest value possible (Kleinberg, 1999). These

metrics were calculated using the random sur-

fer model, where relations with larger weights

influence the probability of being selected by

the surfer (Csardi & Nepusz, 2006). In addition,

the Ego-network of each wetland was visualized

to better explain the results, using the defini-

tion of Crossley et al. (2015). The statistical

analysis and network modeling were conducted

using the R statistical software (R Core Team,

2023), specifically, the library igraph (Csardi

& Nepusz, 2006). For network visualization,

the specialized software Gephi (Bastian et al.,

2009) and Cytoscape (Shannon et al., 2003)

were used.

Finally, we estimated network clusters

based on the survey results. The objective

of this method is to identify network sub-

groups that belong to the same category based

on the ecosystem products collected by their

Table 2

The surveyed communities.

Wetland Communities Applied surveys Rejections Total households

Las Baulas Mata Palo 71 15 86

Villareal 88 34 122

Palo Verde El Rosario 80 8 88

Bagatsí 33 4 37

Caño Negro Caño Negro 62 4 66

Las Cubas 8 1 9*

Maquenque Golfito 25 4 29

Boca San Carlos 31 1 32

Caribe Noreste Tortuguero 105 25 130

San Francisco 58 9 67

Gandoca-Manzanillo Manzanillo 33 15 48

Gandoca 44 7 51

Térraba-Sierpe Ajuntaderas 25 0 25

Coronado 81 15 96

Total 744 142 877**

*According to the INEC population census of 2011, there were 35 households living in Las Cubas. However, at the moment of

the field research, we found that many had left the community. **Based on the INEC population census of 2011, we expected

a total of 955 households; nonetheless, we found a dynamic where many communities have reduced their population due to

many factors such as unemployment. In other places, like in Coronado, in Térraba-Sierpe wetland, the population was 16

households more than expected, thus, it gained population.

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

households. For instance, two wetlands may

belong to the same cluster if the products they

collect are similar, i.e. such as fish from the

same families. To achieve this, we applied the

Walktrap method (Pons & Latapy, 2006), which

utilizes a random walk technique to construct

clusters based on the neighborhood structure of

nodes. The results of this analysis are important

because wetlands within the same cluster tend

to share similar collected or demanded pro-

ducts; therefore, management strategies could

be considered in a similar manner.

RESULTS

Literature review: Before delving into the

empirical results, we present the main ecosys-

tem services in Costa Rica’s Ramsar wetlands

that we found through the literature review.

They are detailed presented in Appendix 1-4.

The types of ecosystem services with the hig-

hest number of identified cases in the literature

are provisioning and regulating. Functions such

as providing society with raw materials and

food are among the most commonly found.

Functions like regulating climate and hydro-

logical related functions are also frequently

mentioned. Cultural and supporting functions

are the ones with the fewest ecosystem services.

This may be due to the difficulty in measuring

such services and, at times, the lack of resour-

ces for research in areas such as ethnology and

biological species. Although there is literature

about the wetlands, very little is focused on

ecosystem services or on the economic valua-

tion of wetlands in Costa Rica.

In summary, Table 3 presents the number

of ecosystem services that the literature review

led to identifying. The Proyecto Humedales

SINAC-PNUD-GEF (2017) is the most com-

plete source because it identified ecosystem

services for all seven wetlands of interest. The

Térraba-Sierpe wetland has the highest number

of services with 23; it is also the one with the

most identified provisioning services and regu-

lating services. The wetlands of Las Baulas and

Caño Negro have the least number of provisio-

ning services identified. Besides that, the rest of

the services are equally distributed.

Mapping social actors around Ramsar

wetlands: Our findings indicate that wetlands

interact with three primary types of social

actors: communal, public, and private indivi-

duals or enterprises. Fig. 2 displays the resul-

ting network, composed of 61 actors with 139

connections. The number of mentions that

each actor receives is used to estimate which

actors are the most central within the network.

In this network, 41 actors (the majority) are

public institutions, 8 are communal, and 5

are private-individual actors, such as producer

organizations. Fig. 2 illustrates the four types of

actors in the network: wetlands (yellow), com-

munal actors (red), public actors (green), and

individual actors (blue). The two main actors

interacting with the wetlands are the Munici-

palities (9 relations) and the Ministry of Public

Security (9 relations).

Table 3

Total mentions of ecosystem services found according to the literature review.

Wetland Type Total

Provisioning Cultural Supporting Regulating

Térraba-Sierpe 5 5 5 8 23

Las Baulas 2 4 4 4 14

Caño Negro 2 4 4 4 14

Maquenque 3 4 4 4 15

Caribe Noreste 3 4 4 4 15

Gandoca Manzanillo 3 4 4 4 15

Palo Verde 3 4 4 5 16

Source: self-elaborated based on literature review.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

We can visualize the network node size

according to the number of incoming links

(network indegree centrality). Fig. 3 shows this

statistic, with Municipalities and the Ministry

of Public Security depicted as the two larger red

nodes. Following closely in size is a third actor,

Public Universities (8 relations), positioned as

the third red node, indicating its significance

within the network. It is important to clarify

that the position of the nodes in the network

is related to their shared relations with others.

Thus, we observe larger node sizes for social

actors that interact with many wetlands and are

positioned at the center of the network. This

results in actors with fewer ties being situated

at the periphery of the network, forming a core-

periphery structure. Additionally, two other

Fig. 2. Ramsar wetland’s social actors network.

important actors, the Costa Rican Institute of

Electricity (ICE, 7 mentions), also depicted in

red, and Communal Associations (7 mentions),

shown in green, are noteworthy.

If we determine the size of the node based

on the wetlands with the most relations in

the network (outdegree centrality), we obtain

the network visualization shown in Fig. 4.

The wetlands that reported the most ties with

other actors were Turberas (22 mentions), Palo

Verde (21 mentions), Gandoca-Manzanillo (20

mentions), and Caribe Noreste (20 mentions).

Additionally, we find Isla del Coco with 18

mentions, Caño Negro with 11 mentions, and

Las Baulas, Maquenque, and Térraba-Sier-

pe with 9 mentions each. In our study case,

Palo Verde, Gandoca-Manzanillo, and Caribe

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Noreste are those with a denser network. Con-

versely, Las Baulas, Maquenque, and Térraba-

Sierpe are the wetlands with fewer connections

to social actors.

In terms of differences in the composition

of the ego-networks, we can differentiate three

groups:

Group 1: Wetlands with 20 and 21 or more

actors that have a presence of public, com-

munal, and private actors, although there

is a majority of public actors. Here we find

Caribe Noreste, Gandoca-Manzanillo, and

Palo Verde wetlands.

Group 2: Wetlands where the combination of

two types of actors is balanced, while the

third type is missing. In this category, we

find Caño Negro with public and commu-

nal actors (11 in total), but without any

private actor. Additionally, Las Baulas wet-

land has only 9 actors, with participation

from three private and six public actors,

but lacks communal participation in the

ego-network.

Group 3: Maquenque and Térraba-Sierpe wet-

lands, each with only nine actors in the

ego-network. In these cases, the network is

predominantly composed of public actors,

with only one communal actor interacting

with them.

The ego-network and the wetland actor

composition can be observed in Fig. 5. This

figure presents the number of actors associated

Fig. 3. Ramsar wetland’s social actors network: size of nodes based on indegree centrality.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

with each wetland. Both the map and the

ego network represent this relational infor-

mation in different ways, contributing to a

better understanding of the socio-ecological

interconnections.

Identifying ecosystem services through

communal leaders: Based on interviews with

communal leaders, numerous ecosystem servi-

ces were identified, with provisioning emerging

as the most prominent service type. Within

this category, the provision of timber and non-

timber products exhibited the highest diversity

and number of ecosystem services. Examples

include fishing, freshwater, and drinking water,

shrimp, lobster, and mollusk harvesting, all of

which are non-timber resources extracted from

wetlands for human consumption. Additiona-

lly, a variety of agricultural crops were cited

frequently by interviewees, such as beans, rice,

maize, oil palm, tubers, banana, pineapple,

sugar cane, plantain, watermelon, and papaya.

Furthermore, pastures and cattle ranging were

highlighted as activities benefiting from wet-

lands, primarily due to water provisioning.

Artisanal fishing was mentioned in six

out of the seven assessed wetlands, with only

Palo Verde wetland not mentioned by the

interviewees. Commercial fishing was speci-

fically noted in Caño Negro Wetland. Various

products were reported to be consumed or

extracted in several wetlands, including wood

(Maquenque and Gandoca-Manzanillo), turtle

eggs (Las Baulas), land animals, and lobster

Fig. 4. Ramsar wetland’s social actors network: size of nodes based on outdegree centrality.

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

(Gandoca-Manzanillo), fresh-water shrimp

(Maquenque), and piangua (Térraba-Sierpe).

Additionally, the importance of wetland use

for transportation was highlighted, particularly

in Maquenque. These provisioning ecosystem

services identified by communal leaders are

summarized in Table 4.

In agriculture, the predominant products

across most wetlands include rice and beans

production, observed in Wetlands such as

Térraba-Sierpe, Palo Verde, and Caño Negro.

Other monoculture crops are also cultivated in

at least two wetlands: palm oil (Térraba-Sierpe

and Northeast Caribbean), banana (Northeast

Caribbean and Gandoca-Manzanillo), and

pineapple (Northeast Caribbean and Maquen-

que). These crops hold significance due to their

extensive production, which necessitates the

use of substantial quantities of agrochemicals.

Additionally, economically vital monocultu-

re crops encompass tubers such as cassava

(Caño Negro and Maquenque), sugarcane (Palo

Verde), plantain (Maquenque), and watermelon

(Palo Verde).

Among the wetlands, Palo Verde stands

out with seven mentioned agricultural pro-

ducts, followed by Térraba-Sierpe with four,

Caño Negro with three, Northeast Caribbean

with three, Maquenque with three, and Gan-

doca-Manzanillo with one. Notably, Las Baulas

wetland did not receive any mentions regar-

ding this ecosystem service. It is essential to

highlight that interviewees emphasized these

products due to their perceived importance to

the local economy. The identified agricultural

products are listed in Table 5.

Livestock farming emerges as a significant

economic activity in five out of the seven wet-

lands, underscoring its importance to the local

communities. Notably, Las Baulas and Gando-

ca-Manzanillo wetlands do not feature promi-

nent livestock farming. Among the wetlands

Fig. 5. Number of social actors per wetland.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Table 4

Provisioning ecosystem services identified by communal

leaders.

Service Wetland

Traditional-small scale fishing Térraba-Sierpe

Las Baulas

Caño Negro

Maquenque

Caribe Noreste

Gandoca Manzanillo

Timber wood extraction Maquenque1

Gandoca-Manzanillo

Marine species eggs Las Baulas

Firewood Caño Negro

Hunting for self-consume Gandoca-Manzanillo

Commercial fishing Caño Negro

Lobster extraction Gandoca-Manzanillo

Freshwater shrimp Maquenque

Las Baulas

Caño Negro

Caribe Noreste

Gandoca-Manzanillo

Tree plantations Maquenque

Honey Palo Verde

Shrimps Caribe-Noreste

Gandoca-Manzanillo

Térraba-Sierpe

Mollusk extraction Térraba-Sierpe

Las Baulas

Transport Maquenque

Térraba-Sierpe

Plants Palo Verde2

Térraba-Sierpe

Fruits 3Las Baulas

Caño Negro

Caribe Noreste

Gandoca-Manzanillo

Palo Verde

Térraba-Sierpe

1. Legal. 2. For handicrafts. 3. No plantations, diverse wild

and traditional species. Source: self-elaborated based on

field data.

services identified. These services primarily

revolve around tourism, sport fishing, environ-

mental education, and recreational activities.

Tourism was reported for all wetlands except

Palo Verde. Sport fishing, on the other hand,

was noted in the wetlands of Térraba-Sierpe,

Table 5

Provisioning ecosystem services related to agriculture

identified by communal leaders.

Service Wetland

Beans production Térraba-Sierpe

Palo Verde

Caño Negro

Rice production Térraba-Sierpe

Palo Verde

Caño Negro

Maiz production Térraba-Sierpe

Palo Verde

Oil palm production Térraba-Sierpe

Caribe Noreste

Tuber production Caño Negro

Maquenque

Banana production Caribe Noreste

Gandoca-Manzanillo

Pineapple production Caribe Noreste

Maquenque

Sugar row production Palo Verde

Plantain production Maquenque

Watermelon production Palo Verde

Papaya production Palo Verde

Grasslands1Palo Verde

1. Refers to give services to maintain the grassland for

livestock. Source: self-elaborated based on field data.

with notable livestock activity, Palo Verde and

Caño Negro stand out, where the provision of

watering services by the wetlands is particularly

valued. Further details can be found in Table 6.

The category of Cultural ecosystem ser-

vices ranks second in terms of the number of

Table 6

Provisioning ecosystem services related to livestock

identified by communal leaders.

Service Wetland

Cattle production Térraba-Sierpe

Palo Verde

Caño Negro

Maquenque

Caribe Noreste

Goat production Gandoca-Manzanillo

Source: self-elaborated based on field data.

16 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Caribe Noreste, and Gandoca-Manzanillo.

Recreational use, distinct from tourism, was

specifically mentioned only in Las Baulas wet-

land. In this context, recreation encompasses

the activities undertaken by the local popula-

tion within the wetland, while tourism involves

visitors from outside the area, be they national

or international. For a detailed breakdown of

cultural ecosystem services by wetland, please

refer to Table 7.

Table 7

Cultural ecosystem services identified by communal

leaders.

Service Wetland

Tourism Térraba-Sierpe

Las Baulas

Caño Negro

Maquenque

Caribe Noreste

Gandoca Manzanillo

Recreation1Las Baulas

Sport fishing Térraba-Sierpe

Caribe Noreste

Gandoca Manzanillo

1. Related to recreational use by community members.

Source: self-elaborated based on field data.

The ecosystem services related to regu-

lation and support appear to be largely unk-

nown to community leaders. Only in Las

Baulas was the wetland’s capacity to mitigate

floods (flood reduction) mentioned, while in

Gandoca-Manzanillo, the significance of the

wetland’s role in providing habitat and shelter

to flora and fauna was emphasized.

In summary, Table 8 illustrates the number

of ecosystem services identified by communal

leaders. It’s important to note that we are not

assigning importance to one ecosystem service

over another; rather, we are highlighting a clear

trend wherein key informants were able to iden-

tify a range of benefits from wetlands. The total

count of ecosystem services ranges between 7

and 13, with Térraba-Sierpe wetland having

the highest count and Las Baulas wetland the

lowest. This disparity underscores the varia-

tions in the relationships between wetlands and

the local population. For instance, in Las Baulas

wetland, provisioning services are fewer com-

pared to other wetlands. Across all wetlands,

provisioning services are more prominent than

cultural services, especially notable in the case

of Térraba-Sierpe and Palo Verde.

Additionally, during the interviews, com-

munal leaders were asked about which ecosys-

tem services they believed could be promoted to

increase benefits to the local population, as out-

lined in Table 9. Communal leaders highlighted

two types of tourism that could be further deve-

loped, which were either not well-established or

only in the initial stages at the time of the field

research. Hence, the local leaders made two

main distinctions. Socially responsible tourism

was mentioned in the wetlands of Térraba-

Sierpe and Las Baulas, while community-based

rural tourism was identified in Palo Verde,

Table 8

Total ecosystem services found according to communal leaders.

Wetland Type Total

Provisioning Cultural Supporting Regulating

Térraba-Sierpe 11 2 0 0 13

Las Baulas 42017

Caño Negro 9 1 0 0 10

Maquenque 9 1 0 0 10

Caribe Noreste 8 2 0 0 10

Gandoca Manzanillo 9 2 1 0 12

Palo Verde 11 0 0 0 11

Source: self-elaborated based on field data.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Maquenque, and Caribe Noreste. Concerning

environmental education, communal leaders

expressed interest in this ecosystem service,

particularly in the case of Gandoca-Manzanillo

and Maquenque wetlands, although their expe-

rience with it has been limited. Generally, the

provisioning ecosystem services to be impro-

ved are related to the promotion of productive

projects aimed at reducing human impact on

the wetland while simultaneously increasing

economic benefits for the local population.

Identification and quantification of main

provisioning services: These results primarily

focus on services that are consumed or directly

collected from the wetland by households. For

fish, we utilized Spanish common names along

with their respective families. In Appendix 5, a

table containing the Spanish common names of

fish, their English equivalents, and their fami-

lies is provided for reference. Table 10 presents

the types of products, their common names,

and the total quantities reported annually by

household heads in the case of the Térraba-

Sierpe wetland. Whenever feasible, we included

the family names of the products, particularly

for fish species.

The results reveal that in the Térraba-

Sierpe wetland, Pianguas are the most extracted

product, totaling 307 440 units. However, fish

exhibit a wide variety of families being captu-

red. Among these, the families Lutjanidae and

Centropomidae stand out, with the highest

number of individuals captured, with 2 174 and

2 562 units respectively.

Table 11 provides information on the

collected products for Maquenque wetland. In

contrast to Térraba-Sierpe wetland, the range of

products in Maquenque wetland is less diverse.

The primary product reported is fish, particu-

larly freshwater fish, with Guapote (Cichlidae),

Barbudo (Pimelodidae), Machaca (Bryconi-

dae), and Mojarra (Cichlidae) being the most

commonly caught species.

Table 12 presents the products collected by

households in the Gandoca-Manzanillo wet-

land. The primary product collected in this

wetland is fish, predominantly comprising sea

fish families. Among these, Jurel (Carangidae)

stands out as the most important.

Table 13 displays the products collected by

households in the Caribe Noreste wetland. This

wetland offers a wide variety of products, inclu-

ding mollusks, crustaceans, and fish. However,

Table 9

Ecosystem services that must be incentivized by authorities: identified by communal leaders.

Type of service Service Wetland

Provisioning Aquaculture Caño Negro 1

Maquenque 2

Organic agriculture Caño Negro

Palo Verde

Water consumption Térraba-Sierpe

Gandoca Manzanillo

Firewood Térraba-Sierpe

Gandoca Manzanillo

Cultural Social-responsible tourism Térraba-Sierpe

Las Baulas

Community-based rural tourism Palo Verde

Maquenque

Caribe Noreste

Environmental education Gandoca Manzanillo

Maquenque

1. Fishing, including the tropical gar fish. 2. River shrimp. Source: self-elaborated based on field data.

18 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

fish have the highest recorded quantity. Among

the fish families, Robalo (Centropomidae),

Jurel (Carangidae), Pargo (Lutjanidae), And

Mojarra (Cichlidae) are the most captured.

Table 14 presents the products collected

by households in the Caño Negro wetland.

Fish is the primary provisioning service in this

wetland, particularly freshwater fish such as

Mojarra (Cichlidae) and Guapote (Cichlidae).

These two account for 77 % of the total repor-

ted products collected.

Table 10

Provisioning ecosystem services: products collected by

households in Térraba-Sierpe wetland.

Type of

Product Product 1

Total

quantity/

year

Mollusk Piangua (Arcidae) 307 440

Fruits Coconut 120

Fruits Mango 25

Fruits Unripe coconut (pipa) 120

Plants Zorrillo 120

Shrimp no id. 26

Fish Bagre (Ariidae) 12

Fish Corvina (Sciaenidae) 12

Fish Macarela (Carangidae) 12

Fish Mero (Serranidae) 48

Fish Jurel (Carangidae) 84

Fish Lisa (Mugilidae) 90

Fish Pargo negro (Lutjanidae) 90

Fish Pargo rojo (Lutjanidae) 90

Fish no id. 240

Fish Roncador (Haemulidae) 300

Fish Robalo Gualaje (Centropomidae) 540

Fish Cuminate (Ariidae) 612

Fish Pargo (Lutjanidae) 1 994

Fish Gualaje (Centropomidae) 2 562

Mollusk Oyster (Osteridae) 24

Total 314 561

1. For the case of fish, we use common local names, with

their respective family names provided in parentheses,

based on Bussing & Lopez (2011), Angulo et al. (2021),

and MarViva (2015). It is important to note that this

information is solely based on the responses from the

respondents, and no biological field work was conducted

to verify or confirm any scientific names.Source: Self-

elaborated based on field data.

Table 11

Provisioning ecosystem services: products collected by

households in Maquenque wetland.

Type of

Product Product 1Total quantity/

year

Fresh water

shrimp

no id 12

Fish Robalo (Centropomidae) 24

Fish no id. 39

Fish Sabalo (Megalopidae) 96

Fish Tilapia (Cichlidae) 96

Fish Roncador (Haemulidae) 120

Fish Guabina (Eleotridae) 684

Fish Bagre (Heptapteridae) 720

Fish Guapote (Cichlidae) 2 424

Fish Barbudo (Pimelodidae) 2 586

Fish Machaca (Bryconidae) 2 676

Fish Mojarra (Cichlidae) 5 622

Total 15 099

1. For the case of fish, we utilize common local names,

with their respective family names provided in parentheses,

based on Ángulo (2013), Ángulo et al. (2021), Instituto

Costarricense de Electricidad (2007), and CRAFF (2023).

It is important to note that this information is solely based

on the responses from the respondents, and no biological

field work was conducted to verify or confirm any scientific

names. Source: Self-elaborated based on field data.

Table 12

Provisioning ecosystem services: products collected by

households in Gandoca-Manzanillo wetland.

Type Of

Product Product 1Total Quantity/

Year

Mollusk Cambute (Strombidae) 48

Fruits Coconut 12

Crustacean Shrimp 12

Crustacean Lobster 36

Fish Pargo Rojo (Lutjanidae) 12

Fish Sabalo (Megalopidae) 12

Fish Tilapia (Cichlidae) 48

Fish Roncador (Haemulidae) 72

Fish Robalo (Centropomidae) 84

Fish Bobo (Mugilidae) 96

Fish Macarela (Carngidae) 204

Fish Pargo (Lutjanidae) 384

Fish Jurel (Carangidae) 9 072

Total 10 092

1. For the case of fish, we utilize common local names,

with their respective family names provided in parentheses,

based on Ángulo (2013) and Ángulo et al. (2021), as we

rely solely on the information provided by the respondents,

and no biological field work was conducted to verify any

possible scientific names. Source: Self-elaborated based

on field data.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Table 15 presents the products collected

by households in the Las Baulas wetland. Fish

is the main provisioning service reported by

households in this wetland, particularly Pargo

(Lutjanidae) and Jurel (Carangidae). These

two species represent 80 % of the total units

collected.

Table 16 illustrates the products collected

by households in Palo Verde wetland. While

wild birds are important, with 328 units regis-

tered by households, fish remains the primary

source of products. Most of the fish are caught

in agricultural irrigation channels and rivers.

Tilapia (Cichlidae) is the most commonly cau-

ght, with 1 162 units, representing 33.7 % of the

total product units.

Table 17 provides a summary of the total

number of units collected by product type and

by wetland. Of the total products collected,

21.5 % are fish and 77. 5% are mollusks. Howe-

ver, these numbers are skewed by the significant

number of Pianguas collected in the Térraba-

Sierpe wetland. Excluding Térraba-Sierpe, fish

captures represent between 78 % and 99 % of

all products across the other wetlands. The

dominance of fish collection is evident, with

percentages ranging from 99 % in Maquenque

to 77.8 % in Palo Verde.

Table 13

Provisioning ecosystem services: products collected by

households in Caribe Noreste wetland.

Type of

Product Product 1Total quantity/

year

Mollusk Clams (no id) 200

Crustacean Jaiba (Potunidae) 240

Mollusk Cambute (Strombidae) 12

Fruits Noni 120

Fruits Uva de playa (Sea Grape) 1

Crustacean Freshwater Shrimp 12

Crustacean Shrimp 372

Fish Tiburon Bolillo (Subclasss:

Elasmobranchii) 12

Fish Mero (Serranidae) 12

Fish Pargo rojo (Lutjanidae) 12

Fish Macarela (Carngidae) 24

Fish Cara Seca-gualaje

(Centropomidae) 26

Fish Calva (Centropomidae) 100

Fish Pampano (Carangidae) 144

Fish Corvina (Sciaenidae) 156

Fish no id. 388

Fish Sabalo (Megalopidae) 216

Fish Dorado (Coryphaenidae) 348

Fish Guapote (Cichlidae) 576

Fish Roncador (Haemulidae) 582

Fish Robalo (Centropomidae) 1 272

Fish Jurel (Carangidae) 1 362

Fish Pargo (Lutjanidae) 1 418

Fish Mojarra (Cichlidae) 1 518

Total 9 222

1. For the case of fish, we employ common local names,

accompanied by their respective family names in

parentheses, as referenced from Ángulo (2013), Ángulo

et al. (2021), and MarViva (2015), given that we solely rely

on the information provided by the respondents, and no

biological field work was conducted to verify any possible

scientific names. In the case of Tiburon Bolillo, it may

refer to at least 24 species of shark that are sold under the

term “bolillo” in Costa Rica (MarViva, 2012). Source: Self-

elaborated based on field data.

Table 14

Provisioning ecosystem services: products collected by

households in Caño Negro wetland.

Type of

Product Product 1Total

quantity/year

Fruits Oranges 120

Wood no id. 30

Crustacean Freshwater shrimp 666

Fish Roncador (Haemulidae) 1

Fish Machaca (Bryconidae) 13

Fish Guapote lagunero (Cichlidae) 72

Fish Barbudo (Pimelodidae) 90

Fish No Id. 96

Fish Sabalo (Megalopidae) 96

Fish Tilapia (Cichlidae) 96

Fish Guabina (Eleotridae) 288

Fish Gaspar (Lepisosteidae) 370

Fish Guapote pinto (Cichlidae) 888

Fish Mojarra (Cichlidae) 1 708

Fish Guapote (Cichlidae) 7 972

Total 12 506

1. For the case of fish, we employ common local names,

with their respective family names in parentheses, based on

Ángulo (2013) and Ángulo et al. (2021), as we solely rely on

information provided by the respondents, and no biological

fieldwork was conducted to confirm any possible scientific

names. Source: Self-elaborated based on field data.

20 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Térraba-Sierpe stands out as the wetland with

the highest quantity, followed by Las Baulas,

Caño Negro, and Maquenque in the second

category. Caribe Noreste and Gandoca-Manza-

nillo wetlands fall into the third category, while

Palo Verde has the least number of products in

the last category.

Table 15

Provisioning ecosystem services: products collected by

households in Las Baulas wetland.

Type of

Product Product 1Total quantity/

year

Fruits Coco 720

Fruits Pipas 24

Crustacean Freshwater shrimp 12

Mollusk Piangua (no id) 144

Fish Dorado (Coryphaenidae) 12

Fish Robalo (Centropomidae) 36

Fish no id. 204

Fish Pargo (Lutjanidae) 5 208

Fish Jurel (Carangidae) 25 824

Total 32 184

1. For the case of fish, we utilize common local names, with

their respective family names in parentheses, sourced from

Ángulo (2013), Ángulo et al. (2021), and MarViva (2015).

Since we solely rely on information from the respondents,

and no biological fieldwork was conducted to confirm any

possible scientific names. Source: Self-elaborated based on

field data.

Table 16

Provisioning ecosystem services: products collected by

households in Palo Verde wetland.

Type of

Product Product 1Total quantity/

year

Wild birds Zarceta (Anatidae) 72

Wild birds Piche (Anatidae) 256

Fruits no id. 300

Crustacean Shrimp 111

Fish Robalo (Centropomidae) 12

Fish Carpa (Cyprinidae) 24

Fish Colossoma-Tambaqui

(Serrasalmidae) 24

Fish Cuminate (Ariidae) 68

Fish Guapote (Cichlidae) 408

Fish Bagre (Heptapteridae) 413

Fish no id. 485

Fish Tilapia (Cichlidae) 1 162

Total 3 450

1. For the case of fish, we utilize common local names,

with their respective family names in parentheses, based

on Ángulo (2013), Ángulo et al. (2021), and van Anrooy

et al. (1996). Since we solely rely on information from the

respondents, and no biological fieldwork was conducted

to ensure any possible scientific names. Source: Self-

elaborated based on field data.

Table 17

Total number of products collected by households.

Wetland Fish Mollusk Crustacean Plants Wild birds Fruits Wood Total

Térraba-Sierpe 6 686 307 464 26 120 0 265 0 314 561

Maquenque 15 087 0 12 0 0 0 0 15 099

Gandoca-Manzanillo 9 984 48 48 0 0 12 0 10 092

Caribe Noreste 8 166 212 624 0 0 220 0 9 222

Las Baulas 31 284 144 12 0 0 744 0 32 184

Palo Verde 2 596 0 111 0 328 300 0 3 335

Caño Negro 11 690 0 666 0 0 120 30 12 506

Total 85 493 307 868 1 499 120 328 1 661 30 396 999

Source: self-elaborated based on field data.

Table 18 shows a summary of some impor-

tant species in the wetlands that match the

field data and the literature. Although the com-

mon names are the same of our documented

fish, without formal identification research, we

cannot ensure that they are the same species.

Therefore, we stay at the fish family identifi-

cation level.

Finally, Fig. 6 depicts a map of the Ramsar

wetlands, with the area colored according to the

total number of units of all collected products.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Table 18

Fish families important for the Ramsar wetlands.

Caribe Noreste Gandoca

Manzanillo

Caño Negro Maquenque

Important fish

families for

provisioning

services**

Cichlidae*

Characidae

Pimelodidae

Carcharinidae

Centroponidae*

Lutjanidae*

Serranidae

Centropomidae*

Carangidae*

Lepisosteidae*

Megalopidae

Centropomidae

Cichlidae*

Poeciliidae

Characidae*

Cichlidae*

Lepisosteidae

Species of

particular

economic interest

- Centropomus

pardelles (Robalo)

- Megalops atlanticus

(Sábalo)

- Atractosteus tropicus

(Gaspar fish)

- Atractosteus tropicus

(Gaspar fish)

- Megalops sp. (Sábalo

real)

- Centropomus sp.

(Róbalo)

- Parachromis sp.

(Guapote)

- Atractosteus tropicus

(Gaspar fish)

- Ciclasoma doviiy (Guapote)

- Ciclasoma loisellei

- Ciclasoma managuense

(Guapote tigre)

- Ciclasoma nicaragüense

(La vieja)

* Families also identified by our study in the respective wetland. ** No fish families or species were detailed in Térraba-Sierpe,

Las Baulas and Palo Verde wetlands. Source: Proyecto Humedales SINAC-PNUD-GEF (2017).

Fig. 6. Total units of products collected by households in the studied wetlands. Source: self-elaborated based on field data.

22 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Socio-ecological network of the provi-

sioning services: Fig. 7 illustrates the net-

work representation of the products collected

by households in the research communities,

the number in the lines refers to the num-

ber of units collected. In the center of the

network, we observe the products common

to many wetlands. For instance, fish such as

pargo (Lutjanidae), macarella (Carangidae),

jurel (Carangidae), robalo (Centropomidae),

and roncador (Haemulidae) are located in the

denser area of the network. Other products

like shrimp and coconut also hold relevance for

households across different wetlands. Conver-

sely, some products are exclusively associated

with specific wetlands, evident in the periphery

of the network. For example, we find wild birds

in Palo Verde, jaibas in Caribe-Noreste, wood

and gaspar fish in Caño Negro, and lobster

in Gandoca-Manzanillo. The dominant tie is

observed in the case of piangua collection in the

Térraba-Sierpe wetland.

To estimate the most central products in

the network, centrality statistics are employed.

Fig. 8 illustrates the network where the size

of the nodes represents the authority score

value. Among the top ten products, eight are

fish types, and one is a crustacean, listed in

descending order: shrimps (authority -score

= 1), robalo (Centropomidae, authority -score

= 0.88), roncador (Haemulidae, authority

-score = 0.81), pargo (Lutjanidae, authority

-score = 0.66), jurel (Carangidae, authority

-score = 0.65), sabalo (Megalopidae, authority

-score = 0.63), guapote (Cichlidae, authority

-score = 0.56), macarella (Carangidae, autho-

rity -score = 0.55), tilapia (Cichlidae, authority

-score = 0.50), mojarra (Cichlidae, authority

-score = 0.48), and coconut (authority -score =

0.43). The likelihood of being ranked highly is

influenced not only by the quantity collected by

households but also by the number of wetlands

where the product is recorded.

The network hub score indicates the wet-

lands where households collect a more diverse

and larger quantity of products, as depicted

in Fig. 9. The highest value is attributed to

Caribe Noreste (hub -score = 1), followed by

Fig. 7. Socio-ecological network representation of products collected by households in Ramsar wetlands. Source: self-

elaborated based on field data.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Térraba-Sierpe (hub -score = 0.81), and Gan-

doca Manzanillo (hub-score = 0.70), with

Maquenque (hub -score = 0.62), and Caño

Negro (hub -score = 0.52) out the top five wet-

lands. Las Baulas (hub -score = 0.45) and Palo

Verde (hub -score = 0.40) wetlands occupy the

last two places in the ranking.

From the presented network, we can extract

the respective Ego-network for each wetland, as

depicted in Fig. 10. Visually, the Ego-networks

of Caribe Noreste and Térraba-Sierpe confirm

that these wetlands are those where households

collect a more diverse number of products.

Meanwhile, Palo Verde and Las Baulas are

where households collect the least.

Finally, Fig. 11 illustrates the network clus-

ters identified in the analysis. Three clusters

were found: the first cluster (green) includes

four wetlands—Gandoca-Manzanillo, Térra-

ba-Sierpe, Baulas, and Caribe-Noreste. The

second cluster (orange) groups Caño Negro

and Maquenque, while the third cluster consists

solely of the Palo Verde wetland. The type of

products and fish families shared among the

wetlands within each cluster determined their

membership.

DISCUSSION

Costa Rica’s wetlands ecosystem services:

Parting from the literature review, regulating

ecosystem services are found to be the most

prevalent in Ramsar wetlands in Costa Rica,

followed by supporting and cultural services.

Provisioning services, on the other hand, are

identified as the least prominent. However, our

findings suggest a different perspective when

considering the local population, as provisio-

ning services emerge as the most identified.

This result aligns with the notion that sup-

porting and regulating services are often less

perceptible, as emphasized by De Groot et al.

(2006).

Fig. 8. Network authority score of products collected by households in Ramsar wetlands. Source: self-elaborated based on

field data.

24 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

This discrepancy is significant, considering

that local communities directly experience and

benefit from wetland ecosystems on a daily

basis. Similar results were reported by Gouwa-

kinnou et al. (2019), who found that provisio-

ning services are predominantly recognized by

villages, particularly those with low socioecono-

mic conditions. In contrast, supporting services

were generally unknown to the local communi-

ties. These findings suggest that socioeconomic

conditions play a significant role in shaping

the identification and perception of different

types of ecosystem services. Examples of factors

that influence local population perceptions of

ecosystem services include educational level,

poverty, and proximity of the community to the

ecosystem (Gouwakinnou et al., 2019). Further

research on this issue in Costa Rica will contri-

bute to and enhance the discussion on wetland

sustainable management.

This underscores the importance of invol-

ving local populations in any policy or manage-

ment efforts aimed at ensuring the sustainable

use of wetlands, as advocated by the Ramsar

Convention (Ramsar Convention Secretariat,

2010). In this context, our work provides valua-

ble insights and information to support this

crucial task. This idea is supported by Evan-

gelista et al. (2024), who explore how Local

Ecological Knowledge (LEK) and perceptions

of ecosystem services are shaped by socio-

ecological variables. For example, participa-

tion in conservation and restoration initiatives

significantly influences the LEK levels of local

populations (Evangelista et al., 2024). Therefo-

re, understanding the ecosystem services most

relevant to communities is essential for desig-

ning and implementing effective initiatives to

maintain and ensure the flow and accessibility

of ecosystem services.

Fig. 9. Network hub score of products collected by households in Ramsar wetlands. Source: self-elaborated based on field

data.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Fig. 10. EGO-Network of Ramsar wetlands. Source: self-elaborated based on field data.

26 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Overall, there exists a notable gap in empi-

rical research on the analysis of ecosystem ser-

vices. Among all the Ramsar wetlands in Costa

Rica, the literature on the Térraba-Sierpe wet-

land contains the most comprehensive infor-

mation about ecosystem services. Conversely,

wetlands such as Las Baulas and Caño Negro

have fewer ecosystem services identified in

the literature.

Social actors and wetland ecosystem ser-

vices: The results of the social mapping reveal

that Palo Verde wetland, Caribe Noreste, and

Gandoca Manzanillo exhibit the highest num-

ber of social actors in their ego-networks.

While public actors predominate, communal

actors are also represented by several entities.

This is significant because wetlands with grea-

ter connectivity to diverse social actors, inclu-

ding communal, public, or individual entities,

are expected to foster greater social cooperation

and control over wetland uses. In this regard,

Bodin et al. (2011) emphasize that involving all

relevant social actors in the planning of wet-

lands is fundamental for achieving robust and

sustainable management.

Therefore, collaboration among national

entities, local governments, non-governmental

organizations, and the local population is cru-

cial for resolving conflicts that may arise over

the use of wetland natural resources. Howe-

ver, achieving this collaboration requires a

mutual understanding and trust among the

actors involved (Bodin et al., 2011). Thus,

merely having more actors in the network

does not necessarily imply greater cooperation.

However, an existing network may hold a com-

parative advantage over wetlands that lack a

social network structure for their management,

as the latter would need to build such a network

Fig. 11. Network clusters of Ramsar wetlands. Source: self-elaborated based on field data.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

from scratch. In our case, this advantage arises

because the social actor network is founded

on prior positive experiences of cooperation

and problem-solving in management. Conse-

quently, there is already a foundation of trust

and shared experience to build upon, as Bodin

et al. (2011) and Kininmonth et al. (2015)

suggest in the context of integrating social

actors into wetland management.

Thus, based on past wetland management

experiences reflected in the social actor net-

work, Maquenque and Térraba-Sierpe wetlands

have fewer social actors in their networks, with

communal groups mentioned only once. Con-

sequently, these wetlands may encounter grea-

ter management challenges when integrating

communal groups into planned actions.

For future wetland management conside-

rations, it is essential to acknowledge that each

situation and management problem must be

evaluated individually for each wetland. This

is because private and public interests do not

always align, and each scenario may require a

tailored approach, as noted by Cárcamo et al.

(2014) in the case of artisanal and industrial

fishers. Clare et al. (2013) highlights that public

policy decisions often create winners and losers

in the use of natural resources, thereby intensi-

fying management tensions.

It is crucial to acknowledge the diverse

ways in which local communities interact with

wetlands, with each wetland exhibiting varying

intensities of these interactions. For example,

Las Baulas offers significant cultural ecosystem

services compared to Maquenque. However,

Maquenque provides more provisioning servi-

ces than Las Baulas. The local socio-economic

dynamics influence the demand for products

and services from the wetland by households.

According to communal leaders, in Palo Verde,

there are minimal benefits to the communities

from tourism. Nevertheless, studies by Proyec-

to Humedales SINAC-PNUD-GEF (2017) and

Moreno et al. (2010) have identified recreatio-

nal and educational benefits from the wetland.

Moreno et al. (2010) conducted a study

that identified cultural ecosystem services such

as tourism and scientific research from Palo

Verde. Local perceived benefits reported by

communal leaders did not match those repor-

ted by Moreno et al. (2010). This difference

may stem from Moreno et al. (2010) including

a broader range of localities, overlapping only

with one of the communities studied. Our fin-

dings also reveal that Palo Verde has the highest

number of social actors involved in its network.

Conversely, local leaders in Las Baulas

wetland recognize that communities benefit

from tourism. This finding aligns with the

conclusions of Reyes et al. (2013) and Proyecto

Humedales SINAC-PNUD-GEF (2017). The

recognition of their use of wetland resources

influences the way local populations interact

with the ecosystem and may affect their willing-

ness to protect the natural resources of the wet-

land. For instance, Reyes et al. (2013) identified

and valued ecosystem services associated with

Las Baulas wetland, with cultural ecosystem

services such as lodging, food, transportation,

and tour operation being prominent. Despite

the involvement of numerous actors in utilizing

wetland benefits in Las Baulas, our findings

indicate that only nine actors are involved in

the wetland’s network.

Communal leaders’ interviews reveal that

for wetlands Caribe Noreste and Gandoca Man-

zanillo, small-scale fishing and tourism are the

main ecosystem services. Proyecto Humeda-

les SINAC-PNUD-GEF (2017) supports these

findings and underlines that food and water

provisioning are central services in both wet-

lands, while genetic material is also an impor-

tant provisioning service in Caribe Noreste

wetland (Proyecto Humedales SINAC-PNUD-

GEF, 2017). We found that Caribe Noreste and

Gandoca Manzanillo are two of the wetlands

with more involved social actors with 20 (Pro-

yecto Humedales SINAC-PNUD-GEF, 2017).

Therefore, potential benefits such as educa-

tional uses, identified by Proyecto Humedales

SINAC-PNUD-GEF (2017) of the wetland, can

be enhanced in both cases.

In a similar case, in Caño Negro wet-

land, small-scale fishing and tourism are

the main ecosystem services underlined by

communal leaders. Proyecto Humedales

28 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

SINAC-PNUD-GEF (2017) identified food

and water provisioning, spiritual, recreational,

aesthetic, and educational services as central

ecosystem services. Nonetheless, there are only

11 social actors identified on the wetland net-

work; a low value compared to Caribe Noreste

wetland (Proyecto Humedales SINAC-PNUD-

GEF, 2017). In Maquenque wetland, communal

leaders also underline the presence of tourism,

however, it was not strongly perceived as an

important activity as it was for the leaders of

Caño Negro. In this wetland, fishing is also

an important activity, especially for household

consumption. Proyecto Humedales SINAC-

PNUD-GEF (2017) also found that food and

water provisioning is a central ecosystem ser-

vice. This author also identified recreational,

aesthetic, and educational services as with high

potential in the wetland.

Térraba-Sierpe is the wetland with more

research done, the main identified ecosystem

services in the literature are flood protection,

natural protection against droughts, nutrient

cycling, biodiversity, spiritual inspiration, pro-

viding “piangua”, shrimp extraction, fishing,

water for agriculture and tourism, capture and

fixation of carbon (Barton, 1995; BIOMARCC-

SINAC-GIZ, 2012; Kocian et al., 2010; Proyecto

Humedales SINAC-PNUD-GEF, 2017; Reyes et

al., 2004; Sanchez et al., 2013) As the authors

mention, most of these services are non-exclu-

sive public goods, benefiting everyone. Our

results indicate that communal leaders mention

a wide range of ecosystem services, where pro-

visioning services such as “piangua” extraction

and small-scale fishing dominate, besides cul-

tural services such as tourism. Other authors

such as Barton (1995), Kocian et al. (2010),

Reyes et al. (2004) and Sanchez et al. (2013) also

found that piangua extraction was one of the

main provisioning services in Térraba-Sierpe

wetland.

In general, Térraba-Sierpe wetland has a

great variety of ecosystem services that crea-

te an economic value. However, our results

show that it is one of the wetlands with the

lowest number of social actors involved on its

network (9 actors). Thereby, the documented

high socioeconomic dynamic of the wetland

demands a high contact with social actors; thus,

wetland authorities confront a challenge ensu-

ring sustainability.

Communal leaders mentioned the cha-

llenge for the local communities to use wetland

natural resources without endangering the sus-

tainability of ecosystem services. To enhan-

ce the coexistence between communities and

the wetland ecosystem, these key informants

propose the necessity to develop sustainable

productive projects with the participation of

different social actors. For instance, tourism

was not mentioned as an ecosystem service in

Palo Verde, although indeed there is tourism

attending to the wetland, locals do not feel that

tourism gives any benefit to the local communi-

ties. Meanwhile, activities such as aquaculture,

organic, and ecological production are propo-

sed for Palo Verde and Caño Negro communal

leaders. Taking advantage of legally framed

resources for consumption such as water and

firewood also seems important to communities.

Environmental education would be a service

desired in wetlands such as Gandoca-Manzani-

llo and Maquenque.

In the case of Gandoca Manzanillo wet-

land, early research of Alvarado (2008) con-

cluded that, at the interaction level, activities

such as organic farming, conservation incenti-

ves, low-impact tourism, agriculture, livestock,

and fishing can provide social and technical

solutions to negative interactions between the

environment and society. These practices could

be crucial and should be encouraged in the

area. However, we aggregate to the last that to

encourage these activities diverse social actors

must be included in the initiatives.

Therefore, identified potential uses such

as communal rural tourism, other uses such as

fishing and traditional tourism require greater

involvement from the Costa Rican Tourism

Institute (ICT) and the Ministry of Environ-

ment and Energy (MINAE) to regulate and

enhance these activities in wetlands. Based on

the wetlands’ social actors network, ICT only

has three mentions. Greater contact with wet-

lands is required by the Ministry of Agriculture

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

and Livestock (MAG), for example, to enhance

the use of forages from invasive species in wet-

lands and by the Ministry of Public Education

(MEP) to implement environmental education

programs in communities geographically coe-

xisting with wetlands. So far, MAG only has

five mentions, and MEP has five mentions in

the network. Much is still to be enhanced in

terms of incorporating social actors to coope-

rative work to improve wetland relations with

social actors.

Furthermore, Municipalities and the

Ministry of Public Security (MPS) are the

actors with the highest number of contacts

in the network (high indegree centrality),

followed by public universities. This highlights

positive relations in terms of controlling illegal

activities within the wetland, developing acade-

mic research in the wetlands, and connecting

the ecosystem to local productive activities.

These ongoing connections should be further

enhanced. Municipalities are the local govern-

ments managing the local territory; thus, they

play a central role in developing sustainable

relations between the local population and

the wetland ecosystems. Collaboration among

social actors in wetland management is crucial,

especially considering the budget reductions

faced by SINAC-MINAE (Programa Estado de

la Nación, 2023). Therefore, incorporating local

communities is essential as they are the direct

beneficiaries of ecosystem services due to their

geographical proximity.

As mentioned, our results position the

local government in a central role within the

wetlands network. This indicates that positive

relationships between wetlands and their mana-

gement are crucial for problem-solving and

cooperation. Another network-based approach,

applied by Kininmonth et al. (2015), focused

on the role of municipalities in the governan-

ce and local management of wetlands. Their

findings suggest that higher interconnections

between local governments are desirable for

effective cooperation in wetland management.

This is because limited resources and in-house

expertise often necessitate collaboration among

neighboring municipalities to achieve goals and

improve operational efficiency (Kininmonth

et al., 2015). However, wetland management

in Costa Rica is centralized under SINAC.

Therefore, further studies should analyze the

relationships between this institution and

municipalities, as well as among local govern-

ments, in managing Ramsar wetlands from a

broader geographical perspective.

Cárcamo et al. (2014) also utilized social

actors to identify ecosystem services. They

found differences in the prioritizations of diffe-

rent stakeholder groups, particularly between

other actors and fishermen. These differences

stem from components related to their activi-

ties and livelihoods. This example underscores

the challenges of cooperative work between

network actors. In the case presented by Cár-

camo et al. (2014), private interests such as

industrial and illegal fishing are perceived as

high threats, while artisanal fishing is seen as

a low threat. Hence, we concur with Cárcamo

et al. (2014) that achieving common goals

and agreements among stakeholders is cru-

cial to improve decision-making and wetland

management. Their opinions and perceptions

must be incorporated into a science-policy

framework. Thus, securing the sustainability of

wetlands and maintaining ecosystem services

requires the involvement of all stakeholders

in the development and implementation of a

management planning process (Ramsar Con-

vention Secretariat, 2010).

Differences in wetlands’ provisio-

ning services: Communal leaders identified

14 provisioning services associated with the

extraction and use of timber and non-timber

products from wetlands, one service related to

transport, 12 services benefiting agricultural

production, and two related to animal produc-

tion. Only three cultural services were reported

by communal leaders. Therefore, the collection

of products from the wetlands, pertaining to

provisioning services, is the most evident for

the informants. The diversity of these products

was quantified using the survey. This techni-

que allows us to delve into the differences in

30 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

products and the demand for wetland products

among households.

Fish is the main product collected by hou-

seholds from the wetlands. There are diffe-

rences between wetlands that do not have a

coast and those that are coastal wetlands. It is

important to mention that most studies iden-

tifying fishing as a relevant ecosystem service

did not provide detailed information about the

fish types, families, or species collected within

the wetland. The only study that reported

families and some species of particular interest

is Proyecto Humedales SINAC-PNUD-GEF

(2017). Table 18 shows the fish families repor-

ted by Proyecto Humedales SINAC-PNUD-

GEF (2017), with the underlined fish families

also registered in the applied survey. We concur

with Proyecto Humedales SINAC-PNUD-GEF

(2017) on these families:

• Caribe Noreste: Cichlidae, Centropomi-

dae, and Lutjanidae.

• Gandoca Manzanillo: Lutjanidae, Centro-

pomidae, Carangidae.

• Caño Negro: Lepisosteidae, Cichlidae.

• Maquenque: Characidae, Cichlidae.

Our results show that in coastal wetlands,

most of the fish families are sea fish, while in

wetlands without a coast, the main fish families

captured are freshwater fishes. Here are the

wetlands and the main fish families captured by

households, with the quantity in units.

• Térraba-Sierpe: Pargo (Lutjanidae, 1 994),

Gualaje (Centropomidae, 2 562).

• Gandoca Manzanillo: Pargo (Lutjanidae,

384), Jurel (Carangidae, 9 072).

• Caribe Noreste: Robalo (Centropomidae,

1 272), Jurel (Carangidae, 1 362), Pargo

(Lutjanidae, 1 418), Mojarra (Cichlidae,

1 518).

• Las Baulas: Pargo (Lutjanidae, 5 208), Jurel

(Carangidae, 25 824).

• Palo Verde: Tilapia (Cichlidae, 1 162).

• Caño Negro: Mojarra (Cichlidae, 1 708),

Guapote (Cichlidae, 7 972).

• Maquenque: Guapote (Cichlidae, 2 424),

Barbudo (Pimelodidae, 2 586), Machaca

(Bryconidae, 2 676), Mojarra (Cichlidae,

5 622).

The wetland where one fish family stands

out over all the wetlands is the case of Jurel

(Carangidae, 25 764) in Las Baulas. The higher

demand for fish in this wetland was somewhat

expected since Las Baulas resides in a region

where tourism and economic development

have increased in the last decade (Programa

Estado de la Nación, 2017). Therefore, there

must be an increase in the wetland products

demanded by the households.

Most of the households reported that they

fish for self-consumption and to pursue addi-

tional income. However, the case of freshwater

fish was mentioned to have fewer market pos-

sibilities than sea fish. The case of Palo Verde is

also a special case because due to the existence

of open irrigation channels for agriculture,

these have become habitat for local and exotic

species brought by the people, for example, the

case of tilapia (Cichlidae, 1 162). The case of

piangua (Mollusk, 307 464) is a special case in

Térraba-Sierpe wetland, where its extraction is

the most important provisioning service identi-

fied and quantified.

In general, the socio-ecological network

authority score shows that shrimps (crusta-

cean), robalo (Centropomidae), roncador

(Haemulidae), pargo (Lutjanidae), and jurel

(Carangidae) are those products that are most

central in the network, determined not only by

the quantity of those but also by the number

of wetlands where they are simultaneously

demanded. The three wetlands that provide

more and more diverse products are Caribe-

Noreste, Térraba-Sierpe, and Gandoca Manza-

nillo. Although, as mentioned, fish is the most

dominant product, each wetland has specific

products demanded by local households, these

products are found on the periphery of the

network. For instance, wild birds in Palo Verde,

lobster (crustacean) in Gandoca Manzanillo,

Gaspar fish in Caño Negro, and jaibas (crusta-

cean) in Caribe Noreste.

The results of the network cluster analy-

sis provide valuable insights for integrating

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

management criteria across different wetlands.

The interconnection patterns reflected in the

network can serve as a structural framework

to understand household demands and their

similar relationships betweenn wetlands. The

seven wetlands are grouped into three network

subclusters. The first cluster includes Caribe

Noreste, Las Baulas, Gandoca Manzanillo, and

Térraba Sierpe. These four wetlands belong to

the same cluster as they share several products

demanded by the households, such as jurel

(Carangidae) and pargo (Lutjanidae). From

a sustainable management perspective, it can

be inferred that certain management criteria

applicable to these products could be similarly

applied to these wetlands.

The second cluster consists of the Maquen-

que and Caño Negro wetlands. Both are fres-

hwater ecosystems located in the northern

region of the country, and their products are

comparable—freshwater fish families domi-

nate the products demanded by households.

This similarity suggests that their management

could integrate similar criteria.

Lastly, Palo Verde exhibits distinct con-

ditions and products compared to the other

wetlands. As a result, it occupies a unique

cluster within the network and warrants a

management approach tailored to its specific

characteristics.

Since fisheries are so important for the

local communities, it is recommended to inte-

grate and motivate local fishers into wetland

governance. For instance, identification of cen-

tral leaders in this activity is a strategy for

improving fisheries management (Ramírez-

Sánchez, 2011), a statement aligned with our

research results, since it highlights how com-

munal leaders emphasize their relationships

with fisheries.

Integrating social actors such as fishers

will have a higher impact since they are socially

and emotionally engaged rather than following

institutions as-rules (Ramírez-Sánchez, 2011).

Moreover, it must be considered the differences

in fish species and levels of engagement of local

leaders and communities. Therefore, different

wetlands may pursue different strategies for

effective management (Marambanyika & Bec-

kedahl, 2017). We concur with Marambanyi-

ka & Beckedahl (2017) and propose that the

results of the network cluster analysis can be

utilized to develop integrated management cri-

teria. Thus, strategies may operate at different

levels: those tailored to the unique characte-

ristics of each wetland, and those addressing

comparable conditions among wetlands within

the same network cluster.

Finally, in the case of fisheries, it is recom-

mended to follow an adaptive management

approach (Sandström, 2011). In this type of

management, social actors continuously con-

sider information on the fisheries conditions

and movements. Therefore, management rules

must be constantly revised based on ecological

knowledge (Sandström, 2011). However, in

the case of Costa Rica, this last point could be

a difficult task since fisheries conditions and

health are not continuously monitored.

Wetland management tensions: If we

integrate the results of the social actors’ net-

work and the results of the provisioning ser-

vices socio-ecological network, we have two

main dimensions of wetland management.

First, collaboration with social actors is key to

addressing tensions with local communities.

Second, attending to the demand in terms of

quantity and diversity of products that hou-

seholds have from the wetlands is crucial to

ensuring sustainability, not only for the ecosys-

tem but also for the local socio-economy of the

population. Therefore, the first dimension is

captured by the network outdegree centrality of

the social actors’ network. The second dimen-

sion is reflected by the provisioning services

socio-ecological network hub-score.

Table 19 presents the recategorization of

the mentioned variables and estimates a total

value based on the combination of both. The

conceptual value behind the first point is that

the higher the network hub score, the higher

the pressure on the natural resources of the

wetland. Therefore, much care should be taken

by the authorities at the moment to coordinate

and control human activities. Meanwhile, the

32 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

number of social actors reveals the social tools

that the wetland authorities may have to coo-

perate and create alliances to solve problems.

Hence, the higher the number of actors positi-

vely linked to the wetland, the higher the possi-

bilities to achieve agreements and solutions by

integrating the opinions of actors.

Here, we assume that the social actors are

willing to cooperate and build trust, as our

network of social actors is based on positive

relationships and prior cooperation experien-

ces. Therefore, following Bodin et al. (2011)

and Kininmonth et al. (2015), a greater number

of relationships in the network, founded on

cooperation and trust, may enhance the social

capital of wetlands, thereby improving the con-

ditions for solving management problems com-

pared to a less interconnected wetland.

The identification shows a high propensity

for wetland management tensions in Térraba-

Sierpe and Maquenque wetlands. These two

cases have a high demand for products and a

low number of social actors on the wetland’s

network. Moderately exposed to management

tensions are Caribe Noreste and Gandoca Man-

zanillo, as they have both a high demand for

products and a high number of actors on their

actors’ network. Caño Negro and Las Baulas

wetlands also have moderate expectations for

tensions, as they have both a low demand for

products and a low number of social actors.

Finally, Palo Verde is the only wetland that has

a low expected propensity for tensions, since

the wetland has a low demand for products and

a high number of actors on the social network.

As observed, 29 % (two out of seven: Térra-

ba-Sierpe and Maquenque) of the wetlands

have conditions conducive to high tension in

managing the wetland ecosystem, while 57 %

(five out of seven: Caribe Noreste, Gandoca

manzanillo, Caño Negro and Las Baulas) have

a moderate propensity and 14 % (one out of

seven: Palo Verde) have a more favorable sce-

nario. These aspects should be addressed by

policymakers, as decisions regarding sustai-

nability and wetlands must be integrated into

a comprehensive planning context to enhance

well-being and improve environmental quality

(Turner et al., 2000). In this regard, Bodin et

al. (2011) emphasize that collaborative mana-

gement, which incorporates strong relations

between social actors, is necessary for suc-

cessful management. This integrated planning

approach is only feasible if the relationships

between social actors and wetland ecological

units are underscored and considered when

assigning responsibilities.

Early literature has already documented

that conflicts in wetlands management are not

uncommon, as wetlands management autho-

rities must contend with both land and water

ecosystems. Therefore, various types of actors

exert pressure on management interests (Han-

sen, 1982). In this context, Clare et al. (2013)

emphasize that winners and losers emerge

depending on how management policies are

designed. They argue that a better approach

Table 19

Identification of wetland management tensions.

Térraba-Sierpe Caribe-Noreste Gandoca Manzanillo Maquenque Caño Negro Las Baulas Palo Verde

Household demand from the

wetland-Network hub score1

Middle High High Middle High Middle High Middle Low Middle Low Low

Social actors interrelated

with the wetland-Network

outdegree2

Low Middle High Middle High Low Middle Low Low High

Total value37 6 5 7 5 6 2

1. Network hub score from the socio-ecological network: Low = 1 (1–0.81), Middle-Low = 2 (0.82–0.53), Middle-High =

3 (0.54–0.4), High = 4 (less than 0.4). 2. Actors interrelated with the wetland (network outdegree) from the social actors’

network: Low = 4 (less than 9), Middle-Low = 3 (9–11), Middle-High = 2 (11–20), High = 1 (more than 20). 3. High = 8–7,

Middle-High = 6–5, Middle-Low = 4–3, Low = 2–1. Source: self-elaborated based on field data.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

to mitigating tensions in wetland manage-

ment involves incorporating diverse opinions

of social actors and their perspectives into

wetland policy discussions. Similarly, to reduce

potential tensions in wetland management,

Cárcamo et al. (2014) stress the importance,

of integrating ecosystem services and com-

mon social and environmental goals in the

early planning stages of marine protected areas.

However, in some cases, the integration of

social and environmental goals has not been

prioritized from the outset. In such instances,

mitigating conflicts is beneficial for protec-

ting wetlands, maintaining ecosystem services,

and coordinating local socioeconomic develop-

ment, as noted by Sun et al. (2021).

As we have demonstrated, combining the

wetland social actor network with the pro-

visioning services network provides valuable

insights into the potential challenges faced by

Ramsar wetlands in Costa Rica. Integrating the

local population into future conservation and

restoration initiatives could enhance the aware-

ness of local inhabitants regarding a broader

range of ecosystem services. For example, the

results show that the Ministry of Education is

not a central actor in the network, as indicated

by its low indegree centrality. Thus, fostering

collaboration between SINAC and the Ministry

of Education together with the communities

could potentially enhance the perspective that

future generations have regarding the benefits

of wetlands. This, in turn, may foster more

positive relationships between communities,

the ecosystem, and management authorities, as

suggested by Evangelista et al. (2024). Conse-

quently, one could expect the social networks of

these wetlands to become better interconnected

in the future.

We also agree with Marambanyika &

Beckedahl (2017) who found that the local

population plays a central role in both bene-

fiting from and conserving Ramsar wetlands.

As our results also suggest, Marambanyika

& Beckedahl (2017) results found that poor

institutional actors participation is associated

with challenges in wetland governance, lea-

ding to management tensions, confusion, and

conflict among social actors. In this context

the engagement of communal leaders and local

committees indicates a commitment to wet-

land conservation, given that their livelihoods

depend on the health of the ecosystem. We fully

endorse Marambanyika & Beckedahl’s (2017)

conclusion that local communities should be

at the core of institutional governance structu-

res, particularly in developing countries where

resources for wetland management and protec-

tion are limited.

Wetland conflicts typically arise from

land use, human activities within wetlands,

and, more recently, urban expansion (Sun et

al., 2021). Similarly, Veas-Ayala et al. (2022)

demonstrated that wetlands in Costa Rica

face various vulnerability drivers associated

with human activities in their vicinity. Spe-

cifically, Térraba-Sierpe wetland was identi-

fied as highly vulnerable, along with Caño

Negro and Palo Verde wetlands. In our study,

Térraba-Sierpe wetland exhibited conditions

conducive to high management tensions, while

Caño Negro had moderately high conditions

and Palo Verde had low conditions, primarily

based on social actors and household demand

for products. Therefore, future research could

benefit from combining approaches to gain a

broader understanding of the challenges in wet-

land management. For example, integrating our

approach, which considers community-based

ecosystem services and wetland social actors’

network, with that of Veas-Ayala et al. (2022),

which focuses on ecosystem health, would pro-

vide valuable insights.

Our findings suggest that sustaining

ecosystem services is achievable through sus-

tainable management that involves social actors

in the process. By focusing on socio-ecological

relationships, our study elucidates that societal

and environmental interdependencies are criti-

cal to understanding wetland socio-ecological

systems (Sayles et al., 2019). Consequently,

actors within socio-ecological networks can

learn from interactions, facilitating the deve-

lopment of updated management strategies

based on socio-ecological changes (Barnes et

al., 2019). It is also noteworthy to emphasize

34 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

that the effectiveness of cooperative decision-

making processes depends on various factors,

including external policies and regulatory

mandates, internal institutional administrative

and budget structures, as well as institutio-

nal cultures and approaches to science and

decision-making criteria (Behnken et al., 2016).

Therefore, identifying constraints on coopera-

tive action early in the wetland management

planning process can help prevent future con-

flicts and management tensions.

Finally, we have shown that local inha-

bitants are aware of the benefits they obtain

from wetlands. However, these benefits are

primarily centered on provisioning and cul-

tural services. While these services vary from

wetland to wetland, fish provisioning is the

most common across all Ramsar wetlands in

Costa Rica. Given that the social actors network

within wetlands highlights clear necessities for

involving these actors, wetland management

authorities should consider these factors to

enhance ecosystem management and mitigate

potential tensions among ecosystem service

beneficiaries. Although these tensions differ

between wetlands, almost all wetlands expe-

rience moderate to high conditions of mana-

gement tensions due to their social network

structures and the demand for wetland pro-

ducts by local 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.

See supplementary material

a05v73s1-suppl1

ACKNOWLEDGEMENTS

All the information here was gathered

within the framework of the project entitled

“Conservation, Sustainable Use of Biodiversity,

and Maintenance of Ecosystem Services of Pro-

tected Wetlands of International Importance”

(Conservación, Uso Sostenible de la Biodi-

versidad y Mantenimiento de los Servicios de

los Ecosistemas de Humedales Protegidos de

Importancia Internacional), which was coor-

dinated between SINAC-PNUD-GEF, project

PIMS 4966 ID 00088054. Special thanks go

to all the members of the project, the coordi-

nators, and all the households and communal

leaders who took part in the research. Thanks

also go to Lucia Vargas for her guidance in

identifying fish families.

REFERENCES

Aguilar, B., & Moulaert, A. (2011). A preliminary ecolo-

gical-economic estimation of the environmental ser-

vice loss due to current conflict in the Isla Portillos

Region in the Caribe Noreste Wetland in Northeastern

Costa Rica [Technical report]. Fundación Neotrópica,

Costa Rica.

Alvarado, H. (2008). Para valorar lo que valoramos. Valora-

ción multicriterio y valor económico total de Gandoca-

Manzanillo (Costa Rica) y San Pond Sak (Panamá)

[Informe técnico]. Unión Internacional para la Con-

servación de la Naturaleza.

Ángulo, A. (2013). Nombres comunes y técnicos de los

peces de agua dulce de Costa Rica. Filología y Lingüís-

tica, 39(2), 77–103.

Ángulo, A., Ramirez-Coghi, A. R., & López, M. (2021).

Claves para la identificación de los peces de las aguas

continentales e insulares de Costa Rica. Parte I: Fami-

lias. UNED Research Journal, 13(1), e3303. http://

dx.doi.org/10.22458/urj.v13i1.3145

Barbier, E. (2011). Wetlands as natural assets. Hydrological

Sciences Journal, 56(8), 1360–1372. https://doi.org/10.

1080/02626667.2011.629787

Barnes, M., Bodin, Ö., McClanahan, T., Kittinger, J., Hoey,

A., Gaoue, O., & Graham, N. (2019). Social-ecolo-

gical alignment and ecological conditions in coral

reefs. Nature Communications, 10, 2039. https://doi.

org/10.1038/s41467-019-09994-1

Barton, D. (1995). Valoración económica parcial de alterna-

tivas de manejo para los humedales de Térraba-Sierpe

Costa Rica [Master’s Thesis]. Universidad Nacional,

Costa Rica.

Bastian, M., Heyman, S., & Jacomy, M. (2009, May 17–20).

Gephi: an open source software for exploring and

manipulating networks [Demonstration Paper]. Third

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

international AAAI conference on weblogs and social

media, 3(1), 361–362. https://doi.org/10.1609/icwsm.

v3i1.13937

Behnken, J., Groningen, J., & Akamani, K. (2016).

Institutional constraints to collaborative

ecosystem management within a wetlands conser-

vation partnership. Journal of Contemporary Water

Research & Education, 158, 19–33. https://doi.

org/10.1111/j.1936-704X.2016.03216.x

BIOMARCC-SINAC-GIZ. (2012). Evaluación de Carbono

en el Humedal Nacional Térraba Sierpe [Informe téc-

nico]. Sistema Nacional de Áreas de Conservación y

Agencia de cooperación Alemana para el Desarrollo.

San José, Costa Rica.

Bodin, Ö., & Prell, C. (2011). Social Networks and Natural

Resource Management. Cambridge University Press.

Bodin, Ö., Ramirez-Sanchez, S., Ernstson, H., & Prell,

C. (2011). A social relational approach to natural

resource governance. In Ö. Bodin & C. Prell (Eds.),

Social Networks and Natural Resource Management

(pp. 3–28). Cambridge University Press. https://doi.

org/9780521146234

Brandes, U., & Erlebach, T. (2005). Network Analysis: Fun-

damental Foundations. Springer-Verlag.

Burkhard, B., Crossman, N., Nedkov, N., Petz, K., &

Alkemade, R. (2013). Mapping and modelling

ecosystem services for science, policy and practice.

Ecosystem Services, 4, 1–3. https://doi.org/10.1016/j.

ecoser.2013.04.005

Burt, R. (1999). The social capital of opinion leaders.

The Annals of the American Academy of Poli-

tical and Social Science, 566, 37. http://dx.doi.

org/10.1177/000271629956600104

Bussing, W., & López, M. (2011). Peces: Demersales y pelági-

cos costeros del Pacífico de Centroamérica Meridional.

Editorial UCR.

Cárcamo, F., Garay-Flühmann, R., Squeo, F., & Gay-

mer, C. (2014). Using stakeholders’ perspective of

ecosystem services and biodiversity features to plan

a marine protected area. Environmental Science &

Policy, 40, 116–131. http://dx.doi.org/10.1016/j.

envsci.2014.03.003

Castro, A. J., García-Llorente, M., Martín-López, B., &

Iniesta-Arandia, I. (2013). Multidimensional Approa-

ches in Ecosystem Services Assessment. In D. Alca-

raz-Segura, M. Di Bella & L. Straschnoy (Eds.), Earth

Observation of Ecosystem Services, (pp. 445–461).

CRC Press.

Csardi, G., & Nepusz, T. (2006). The igraph software

package for complex network research. InterJournal

Complex Systems, 1695(5), 1–9.

Chopra, K., & Adhikari, S. K. (2004). Environment deve-

lopment linkages: modelling a wetland system for

ecological and economic value. Environment and

Development Economics, 9, 19–45. https://doi.

org/10.1017/S1355770X03001037

Clare, S., Krogman, N., & Caine, K. (2013). The “balan-

ce discourse”: A case study of power and wetland

management. Geoforum, 49, 40–49. http://dx.doi.

org/10.1016/j.geoforum.2013.05.007

Constanza, R., d’Arge, R., De Groot, R., Farber, S., Grasso,

M., Hannon, B., Limburg, K., Naeem, S., O’Neill, R.,

Paruelo, J., Raskin, R., Sutton, P., & van den Belt, M.

(1997). The value of the world’s ecosystem services

and natural capital. Nature, 387, 253–260. https://doi.

org/10.1038/387253a0

CRAFF. (2023). Guía de Especies: Curada por ACPM-

CRAFF. Asociación Costarricense de Pescadores con

Mosca, San José, Costa Rica. https://www.craft-acpm.

org/guia-especies

Crona, B., & Bodin, Ö. (2006). What you know is who

you know? Communication patterns among resource

users as a prerequesite for co-management. Ecology

and Society, 11(2), 7.

Crossley, N., Bellotti, E., Edwards, G., Everett, M., Koski-

nen, J., & Tranmer, M. (2015). Social Network Analysis

for Ego-Nets. SAGE Publications Ltd.

Crossman, N., Burkhard, B., Stoyan, N., Willemen, L.,

Petz, K., Palomo, I., Drakou, E., Martín-Lopez, B.,

McPhearson, T., Boyanova, K., Alkemade, R., Egoh,

B., Dunbar, M., & Maes, J. (2013). A blueprint for

mapping and modelling ecosystem services. Ecosys-

tem Services, 4, 4–14. http://dx.doi.org/10.1016/j.

ecoser.2013.02.001

De Groot, R., Stuip, M., Finlayson. M., & Davidson,

N. (2006). Valuing wetlands: Guidance for valuing

the benefits derived from wetland ecosystem services

[Reporte técnico, N°3]. CBD Technical Series 27.

Ramsar Convention Secretariat, Switzerland.

Evangelista, V., Scariot, A., Mancini-Teixeira, H., Lustosa-

Júnior, I. M. (2024). Local ecological knowledge

and perception as a strategy in the management

of ecosystem services. Journal of Environmental

Management, 368, 122095. https://doi.org/10.1016/j.

jenvman.2024.122095

Frank, O. (2005). Network Sampling and Model Fitting. In

P. Carrington, J. Scott & S. Wasserman (Eds.), Models

and Methods in Social Network Analysis (pp. 31–56).

Cambridge University Press.

Gouwakinnou, G., Biaou, S., Vodouhe, F., Tovihessi, M.,

Awessou, B., & Biaou, H. (2019). Local perceptions

and factors determining ecosystem services identifi-

cation around two forest reserves in Northern Benin.

Journal of Ethnobiology and Ethnomedicine, 15, 61.

https://doi.org/10.1186/s13002-019-0343-y

36 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Gunderson, L., & Holling, C. S. (2002). Panarchy: Unders-

tanding transformations in human and natural sys-

tems. Island Press.

Gutiérrez, I., & Siles, J. (2008). Diagnóstico de medios de

vida y capitales de la comunidad de Humedales de

Medio Queso, Los Chiles, Costa Rica. Unión Mundial

para la Naturaleza.

Hansen, P. (1982). Issues, conflicts and strategies in wet-

land management for waterfowl. Canadian Water

Resources Journal, 7(4), 1–21. https://doi.org/10.4296/

cwrj0704001

Instituto Costarricense de Electricidad. (2007). Catálogo de

peces del río Pirrísv [Reporte técnico]. Instituto Cos-

tarricense de Electricidad, Gobierno de Costa Rica,

San José, Costa Rica.

Instituto Nacional de Estadística y Censos. (2011). XII

Censo Nacional de Población y Vivienda 2011 [Repor-

te técnico]. Instituto Nacional de Estadística y Censos,

Gobierno de Costa Rica, San José, Costa Rica.

Kininmonth, S., Bergsten, A., & Bodin, Ö. (2015). Closing

the collaborative gap: Aligning social and ecological

connectivity for better management of interconnected

wetlands. AMBIO, 44(Suppl. 1), S138–S148. https://

doi.org/10.1007/s13280-014-0605-9

Kleinberg, J. (1999). Authoritative Sources in a Hyper-

linked Environment. Journal of the Association for

Computing Machinery, 46(5), 604–632. https://doi.

org/10.1145/324133.324140

Kocian, M., Batker, D., & Harrison-Cox J. (2010). Nature’s

Value in the Térraba-Sierpe National Wetlands: The

Essential Economics of Ecosystem Services. Earth

Economics.

Limburg, K., O’Neil, R., Constanza, R., & Farber S.

(2002). Complex systems and valuation. Ecologi-

cal Economics, 41, 409–420. https://doi.org/10.1016/

S0921-8009(02)00090-3

Marambanyika, T., & Beckedahl, H. (2017). Institutional

arrangements governing wetland utilization and con-

servation in communal areas of Zimbabwe. Review of

Social Sciences, 2(1), 1–17. https://doi.org/10.18533/

rss.v2i1.71

MarViva. (2012). Guía de identificación de filetes de pescado

y mariscos [Reporte técnico]. Fundación MarViva.

MarViva. (2015). Estándar de Responsabilidad Ambiental

para la Comercialización de Pescado de Mar: Guía de

Especies [Reporte técnico]. Fundación MarViva.

Mitsch, W., Gosselink, J., Anderson, C., & Zhang, L. (2009).

Wetland Ecosystems. John Wiley & Sons, Inc.

Moreno, M., González, S., & Mora, C. (2010). Análisis de

las contribuciones socioeconómicas del Parque Nacio-

nal Palo Verde: Un nido para la investigación y la

educación [Reporte técnico]. El Centro Internacional

de Política Económica para el Desarrollo Sostenible

(CINPE), Universidad Nacional de Costa Rica & Sis-

tema Nacional de Áreas de Conservación, Gobierno

de Costa Rica.

Pagella, T., & Sinclair, F. (2014). Development and use of a

typology of mapping tools to assess their fitness for

supporting management of ecosystem service provi-

sion. Landscape Ecology, 29, 383–399.

Pons, P., & Latapy, M. (2006). Computing communities in

large networks using random walks. Journal of Graph

Algorithms and Applications, 10(2), 191–218. https://

doi.org/10.7155/jgaa.00124

Programa Estado de la Nación. (2017). Capítulo 2: Equidad

e integración social. Informe Estado de la Nación

N°23. PEN CONARE. San José, Costa Rica.

Programa Estado de la Nación. (2023). Capítulo 4: Armonía

con la naturaleza. Informe Estado de la Nación N°29.

PEN CONARE. San José, Costa Rica.

Proyecto Humedales de SINAC-PNUD-GEF. (2017). Valo-

ración de los servicios ecosistémicos que ofrecen siete de

los humedales protegidos de importancia internacional

en Costa Rica: Palo Verde, Caribe Noreste, Caño Negro,

Gandoca-Manzanillo, Maquenque, Térraba-Sierpe y

Las Baulas [Reporte técnico]. SINAC/CINPE-UNA/

PNUD.

R Core Team. (2023). R: A language and environment for

statistical computing [Computer software]. R Foun-

dation for Statistical Computing, Vienna, Austria.

http://www.r-project.org/index.html

Ramírez-Sánchez, S. (2011). Who and how: engaging well-

connected fishers in social networks to improve fis-

heries management and conservation. In Ö. Bodin &

C. Prell (Eds.), Social Networks and Natural Resource

Management (pp. 119–146). Cambridge University

Press.

Ramsar Convention Secretariat. (2010). Wise use of wet-

lands: Concepts and approaches for the wise use of wet-

lands [Technical handbook, 4th ed., vol. 1]. Ramsar

Convention Secretariat, Switzerland.

Ramsar Convention Secretariat. (2016). An Introduction

to the Ramsar Convention on Wetlands [Convention

manual, 7th ed.]. Ramsar Convention Secretariat,

Switzerland.

Ramsar Convention Secretariat. (2023a). The Convention

on Wetlands: Costa Rica country profile [Technical

report]. Ramsar Convention Secretariat, Switzerland.

https://www.ramsar.org/country-profile/costa-rica

Ramsar Convention Secretariat. (2023b). Contracting Par-

ties to the Ramsar Convention [Technical report].

Ramsar Convention Secretariat, Switzerland.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63638, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Reyes, V., Miranda, M., Monge, C., & Salas, F. (2004). Valo-

ración Económica del Ecosistema Humedal Nacional

Térraba-Sierpe y Propuesta de Mecanismos para su

Sostenibilidad [Reporte técnico]. Unión Mundial para

la Naturaleza, Costa Rica.

Reyes, B., Biggs, R., Cumming, G., Elmqvist, T., Hej-

nowicsz, A., & Polasky, S. (2013). Getting the measure

of ecosystem service: a social-ecological approach.

Frontiers in Ecology and the Environment. https://doi.

org/10.1890/120144

Rogers, E. (2003). Diffusion of innovations (5th ed.). Free

Press.

Sanchez, R., Reyes, V., Mora, R., Castro, R., Madrigal, C.,

& Cascante, S. (2013). INFORME FINAL: Valoración

Económica de Usos Alternativos de la Tierra del Área

de Amortiguamiento y del HNTS [Reporte técnico].

Centro de Derecho Ambiental y los Recursos Natura-

les (Cedarena), Costa Rica.

Sandström, A. (2011). Social networks, joint building, and

adaptability: the case of local fishery management. In

Ö. Bodin & C. Prell (Eds.), Social Networks and Natu-

ral Resource Management (pp. 288–321). Cambridge

University Press.

Sayles, J., Mancilla, M., Hamilton, M., Alexander, S. M.,

Baggio, J., Fisher, A. P., Ingold, K., Meredith, G., &

Pittman, J. (2019). Social-ecological network analy-

sis for sustainability sciences: a systematic review

and innovative research agenda for the future. Envi-

ronmental Research Letters, 14, 093003. https://doi.

org/10.1088/1748-9326/ab2619

Shannon, P., Markiel, A., Ozier, O., Baliga, N., Wang, J.,

Ramage, D., Amin, N., Schwikowski, B., & Ideker, T.

(2003). Cytoscape: A software environment for inte-

grated models of biomolecular interaction networks.

Genome Research, 13(11), 2498–2504. https://doi.

org/10.1101/gr.1239303

Sistema Nacional de Áreas de Conservación. (2023). Regis-

tro Nacional de Humedales [Reporte técnico]. Sistema

Nacional de Áreas de Conservación (SINAC), Costa

Rica, San José. https://geos1pne.sirefor.go.cr/wfs?

Straton, A. (2006). A complex system approach to the

value of ecological resources. Ecological Economics,

56, 402–411.

Sun, Y., Ge, X., Liu, J., Chang, Y., Liu, G-J., & Chen,

F. (2021). Mitigating Spatial Conflict of Land use

for Sustainable Wetlands Landscape in Li-Cia-River

Region of Central Jiangsu, China. Sustainability, 13,

11189. https://doi.org/10.3390/su132011189

Turner, K., van den Bergh, J., Söderqvist, T., Barendregt, A.,

van der Straaten, J., Maltbt, E., & Ierland, E. (2000).

Ecological Economic analysis of wetlands: scienti-

fic integration for management and policy. Ecologi-

cal Economics, 35(1), 7–23. https://doi.org/10.1016/

S0921-8009(00)00164-6

Valente, T., & Davis, R. (1999). Accelerating the diffusion

of innovations using opinion leaders. The Annals of

the American Academy of Political and Social Science,

566, 55–67.

Van Anrooy, R., Günter, J., Boza, J., & Gálvez, N. (1996).

A Preliminary Market Research About Tambaqui

(Colossoma macropomum) in Costa Rica. UNICIEN-

CIA, 13, 5–11.

Van der Valk, A. (2012). The biology of freshwater wetlands

(2nd ed.). Oxford University Press Inc.

Veas-Ayala, N., Alfaro-Córdoba, M., & Quesada-Román,

A. (2022). Costa Rican wetlands vulnerability index.

Progress in Physical Geography, 47(4), 1–20. https://

doi.org/10.1177/03091333221134189

Xu, X., Chen, M., Yang, G., Jiang, B., & Zhang, J. (2020).

Wetland ecosystem services research: A critical

review. Global Ecology and Conservation, 22, e10027.

https://doi.org/10.1016/j.gecco.2020.e01027