Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Coral reefs restoration initiatives in Costa Rica: ten years building hope

Juan José Alvarado1*; https://orcid.org/0000-0002-2620-9115

Katharine Evans2; https://orcid.org/0009-0004-2116-4714

Joan A. Kleypas3,4; https://orcid.org/0000-0003-4851-7124

José Andrés Marín-Moraga4; https://orcid.org/0000-0001-8499-5620

Mauricio Mendez-Venegas5; https://orcid.org/0000-0003-4855-3731

Carlos Pérez-Reyes6; https://orcid.org/0000-0003-0663-9019

Marylaura Sandoval4; https://orcid.org/0009-0002-8120-412X

María José Solano1; https://orcid.org/0009-0002-4624-2913

Tatiana Villalobos-Cubero4; https://orcid.org/0000-0002-2259-5514

1. Centro de Investigación en Ciencias del Mar y Limnología (CIMAR), Escuela de Biología, Centro de Investigación en

Biodiversidad y Ecología Tropical (CIBET), Universidad de Costa Rica, San Pedro, 11501-2060 San José, Costa Rica;

juan.alvarado@ucr.ac.cr (*Correspondance); majosogo98@gmail.com

2. Marine Conservation Costa Rica, Quepos, Costa Rica; kat@marineconservationcostarica.org

3. National Center of Atmospheric Research (NCAR), Boulder, Colorado, Estados Unidos de América; kleypas@ucar.edu

4. Raising Coral Costa Rica (RCCR), San José, Costa Rica; jamm@raisingcoral.org; marylaura@raisingcoral.org:

tatiana14.villalobos@gmail.com

5. Ministerio de Ambiente y Energía, Sistema Nacional de Áreas de Conservación, Costa Rica;

mauricio.mendez@sinac.go.cr

6. Instituto Nacional de Aprendizaje, Puntarenas, Costa Rica; cperezreyes@ina.ac.cr

Received 19-VII-2024. Corrected 15-X-2024. Accepted 27-I-2025.

ABSTRACT

Introduction: Costa Rica has been recognized worldwide for its high biodiversity and the conservation actions

it has implemented. One of the most iconic ecosystems are coral reefs, which have experienced strong anthropo-

genic and natural pressures in recent years. To ensure these ecosystems’ preservation and services, a series of coral

restoration initiatives have emerged in the last ten years along both Pacific and Caribbean coasts.

Objective: To document the different advances of the various coral restoration initiatives Costa Rica’s Pacific

and Caribbean coasts.

Methods: This review focuses on the implementation of the different reef restoration efforts, indicating the

restoration techniques used, the coral species used in the nurseries, as well as the general results of survival and

growth.

Results: The first coral restoration project in Costa Rica occurred in the 1990s and was the only such effort until

the 2010s. In 2013, a pilot project began in the Golfo Dulce area, which was later replicated in other areas of the

country, such as Manuel Antonio, Sámara, and Bahía Culebra on the Pacific coast, and more recently in Punta

Cahuita in the Caribbean. Various artificial structures have been used as nurseries in the water column such as

trees and ropes, and benthic structures such like A-frames, tables, and spiders, the former being very effective

for branching species (Pocillopora spp.), while the rest have worked successfully both for branched and massive

species (Pavona spp. and Porites spp.). The results shows a growth rates have been between 6 and 9 cm/year, with

survival of 60–90 % of the branching and massive colonies. All sites were seriously affected by the El Niño 2023

phenomenon, with high bleaching values and loss of colonies in the nurseries and on the reef.

https://doi.org/10.15517/rev.biol.trop..v73iS1.63695

SUPPLEMENT

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

INTRODUCTION

Ecological restoration is a series of ongoing

activities that seek to conserve biodiversity and

human well-being. Within this continuum of

activities, ecological restoration is a process that

assists in the recovery of an ecosystem that has

been degraded, damaged or destroyed (Gann et

al., 2019). Coral reef restoration garners atten-

tion as human activities and anthropogenic

climate change result in an unprecedented scale

of reef degradation, compromising ecosystem

resilience and survival (Hoegh-Guldberg et al.,

2023; Hughes et al., 2018; Suggett et al., 2024).

Restoration projects dating back to the early

1980’s focused on research and understanding

the basics of coral gardening (Bowden-Kerby,

1997; Guzmán, 1991; Guzmán & Cortés, 1989;

Harriott & Harrison, 1984; Rinkevich, 1995).

In the 2000’s, more projects were established

as restoration practices were increasingly rec-

ognized as a positive tool for reef management

and to prevent the extinction of certain coral

species such as acroporids in the Caribbean

(Lirman & Schopmeyer, 2016; Rodriguez-Mar-

tinez et al., 2014; Young et al., 2012). A recent

emergence of initiatives occurred after 2016

following El Niño’s global scale impact on coral

Conclusion: Despite geographic and oceanographic distinctions, these projects have emphasized local engage-

ment and perception of coral reefs, fostered intersectoral public-private collaborations for financial and human

resources, and operated within established governmental regulatory frameworks. All projects face vulnerabilities

such as El Niño events and Harmful Algal Blooms.

Key words: Acropora; Caribbean; El Niño; growth rate; nurseries; Pacific; Pavona; Pocillopora; Porites; survival.

RESUMEN

Iniciativas de restauración de arrecifes de coral en Costa Rica: diez años construyendo esperanza

Introducción: Costa Rica ha sido reconocido mundialmente por su alta biodiversidad y las acciones de conserva-

ción que han implementado. Uno de los ecosistemas más icónicos son los arrecifes de coral, los cuales han expe-

rimentado fuertes presiones antrópicas y naturales en los últimos años. Para asegurar la preservación de dichos

ecosistemas y de sus servicios ecosistémicos, una serie de iniciativas de restauración coralinas han surgido en los

últimos diez años a lo largo de ambas costas, Pacífica y Caribe.

Objetivo: Documentar los diferentes avances de las variadas iniciativas de restauración coralinas tanto en la costa

Pacífica y Caribe de Costa Rica.

Métodos: Esta revisión bibliográfica se centra en los avances de los diferentes grupos, indicando las técnicas de

restauración utilizadas, las especies de coral utilizadas en los viveros, así como los resultados generales de sobre-

vivencia y crecimiento.

Resultados: El primer proyecto de restauración coralina en Costa Rica se da en la década de los 1990, y fue el

único esfuerzo hasta la década de 2010. En el 2013 comienza un proyecto piloto en la zona de Golfo Dulce, que

luego se va replicando en otras zonas del país como Manuel Antonio, Sámara, Bahía Culebra, en la costa Pacífica,

y más recientemente en Punta Cahuita en el Caribe. Se han utilizado varias estructuras como viveros en la

columna de agua como árboles y cuerdas, y bentónicos como marcos en A, mesas, y arañas, siendo estas últimas

muy efectivas para especies ramificadas (Pocillopora spp.), mientras que el resto han funcionado exitosamente

tanto para especies ramificadas como masivas (Pavona spp. y Porites spp.). En términos generales las tasas de

crecimiento han rondado entre 6 y 9 cm/año, con supervivencias entre 60–90 % de las colonias. Todos los sitios

fueron gravemente afectados por el fenómeno de El Niño 2023, con altos valores de blanqueamiento y pérdida

de colonias en los viveros.

Conclusión: A pesar de las distinciones geográficas y oceanográficas, estos proyectos han enfatizado el compromi-

so y la percepción local de los arrecifes de coral, han fomentado colaboraciones público-privadas intersectoriales

para recursos financieros y humanos y han operado dentro de marcos regulatorios gubernamentales establecidos.

Todos los proyectos enfrentan vulnerabilidades como eventos de El Niño y floraciones de algas nocivas.

Palabras clave: Acropora; Caribe; El Niño; Pacífico; Pavona; Pocillopora; Porites; sobrevivencia; tasa de crecimien-

to; viveros.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

reefs, with special urgency in the Great Barrier

Reef (Alvarado et al., 2020; McLeod et al., 2022;

Stuart-Smith et al., 2018; Suggett et al., 2019).

Restoration goals and techniques have

evolved over the last 20 years. As changing

environments modified ecosystem health and

composition, new objectives and technologies

were required to improve efficiency and holis-

tically tackle the interventions (Mumby et al.,

2021; Vardi et al., 2021). Initially, coral restora-

tion was centered on direct coral transplanta-

tion and coral gardening, and often neglected

the maintenance of species and genetic diversi-

ty (Baums et al., 2019; Boström-Einarsson et al.,

2020; Rinkevich, 2005). Over time, scientists

and coral reef managers suggested a combina-

tion of techniques and even, the implementa-

tion of higher risk practices, going from genetic

and sexual advances (e.g. assisted evolution and

coral migration) to socio-ecological interven-

tions, as citizens developing key restoration

efforts (Anthony et al., 2017; Hagedorn et al.,

2021; Kleypas et al., 2021a; Shaver et al., 2022;

Villalobos-Cubero et al., 2023).

In Latin America, reef restoration has been

widely practiced in the Caribbean, with less

incidence in the Eastern Tropical Pacific (ETP)

(Bayraktarov et al., 2020; Bowden-Kerby et al.,

2005; Rodriguez-Martinez et al., 2014; Young

et al., 2012). Concern for coral reefs’ health

and the lack of information have motivated

more projects in ETP, some with a research

purpose, while some others in fact, to assist

natural recovery as possible (Bayraktarov et

al., 2020; Combillet et al., 2022; Liñán-Cabello

et al., 2010; Martínez-Castillo et al., 2023). In

recent years, the number of coral restoration

efforts in the ETP has increased dramatically

(Bayraktarov et al., 2019). However, experience

is still limited compared to other regions, such

as the Caribbean or the Indo-Pacific (Boström-

Einarsson et al., 2020) and few projects use

coral gardening (Ishida-Castañeda et al., 2020).

Certainly, the Coral Restoration Consortium’s

lead role has been key to spreading scientific

progress and facilitating experience exchange,

which is helping new restoration projects stay

at the forefront (Vardi et al., 2021). Technology

implementation and restoration scale are still

a challenge but understanding and coordi-

nating efforts at national and regional lev-

els means an opportunity to connect funds,

reefs and challenges (Anthony et al., 2017;

Bayraktarov et al., 2020).

With this literature review we wish to

recapitulate the advances in the last ten years

in the restoration of coral reefs in Costa Rica,

both on its Pacific and Caribbean coasts. Our

interest is to present a brief account of various

artificial reef establishment activities from the

1980s to the present, government support for

coral restoration initiatives, and then present

the different restoration projects that have been

carried out in the last years. We conclude with

a series of recommendations and future lines of

work at the national and regional level.

Previous restoration initiatives

(80 & 90’s) in Costa Rica

The first initiatives to restore marine envi-

ronments in Costa Rica come from the 80’s,

when an attempt to deploy artificial reefs in

the Gulf of Nicoya (Playa Mantas) to provide

alternatives for fisheries management (Campos

& Guzmán, 1986; Guzman et al., 1988). Using

approximately 1 900 tires tied with propylene

rope and galvanized wire, an artificial reef was

built at 8–10 m depth in modules of 5, 10 and

15 units. Monthly data was collected to com-

pare the fish population versus a natural rocky

reef for a year. Throughout this period, the pro-

gram managers determined that the biomass

and number of species of commercial interest

were greater than in rocky reefs, being ten times

more productive (Campos & Guzman 1986;

Guzman et al., 1988). Likewise, they indicated

that the artificial reef served as a breeding area

for species of commercial value (Campos &

Gamboa, 1989; Guzman et al., 1988). By 1987,

the reef had 5 000 tires, and they observed a

change in the structure of the fish community

composition, with a change in dominance from

lutjanide to haemulidae. Species of commercial

interest represented 47 % of the total species

(Campos & Gamboa, 1989). By April 1987,

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Thorne et al. (1989) investigated the same site

to determine the potential use of hydroacous-

tics for studies of abundance and behavior in

artificial reefs. It was possible to detect a strong

association of fish with the artificial reef during

daylight hours, but a high dispersion during

the night.

After this initiative in Playas Mantas, other

attempts were made in Bahía Culebra in the

1990s with PVC pipes (J. Campos, personal

communication, March 2024), or in various

areas of the Gulf of Nicoya with tires (Inter

Press Service, 1999), “ReefBalls” (Infobae, 2021;

La Nación, 2005; Soto-Méndez, 2021) or ship-

wrecks (La Nación, 2000; La Nación, 2004).

Likewise, the Costa Rican Institute of Electric-

ity (ICE) established an artificial reef with por-

celain insulators in Playa Hermosa Guanacaste

(Garza, 2019). The most recent initiative comes

from the National Learning Institute-Institu-

to Nacional de Aprendizaje (INA) in Playa

Blanca (Punta Leona) using “ReefBalls (Mar-

tínez, 2021). All initiatives involved civil soci-

ety through fishermen’s organizations, primary

schools or groups of divers, but to date there

are no publications or data that quantify their

follow-up. Due to this, in 2021, the “Technical

guide for the establishment of artificial reefs in

Costa Rica” was established, however this guide

has not been ratified and made official by the

Government. This manual is intended to be a

regulatory instrument of the technical process-

es to be considered in the planning, installation

and management projects of artificial reefs in

Costa Rica, where the possible implications and

responsibilities they entail are clear. Likewise,

for 2018, a specific guide was generated for the

area of Paquera, in the Gulf of Nicoya (Sánchez

& Azofeifa, 2018).

The only coral reef restoration initiative

established in that period was performed on

the Biological Reserve Isla del Caño, in the

southern Pacific of Costa Rica (Guzman, 1991).

In response to the impacts of the 1982–1983

El Niño phenomenon and red tides on mas-

sive mortalities of the reefs of this island and

arguing the low reproductive potential of the

corals in the area, Guzman (1991) set out to

demonstrate the feasibility of reef restoration.

Fifty-two coral fragments of Pocillopora were

placed on the reef frame by attaching them with

wire to 30-cm-long steel stakes and were moni-

tored for almost three years (1987–1989). The

results indicated that live coral cover increased

from 20 % to nearly 60 %, with a mortality rate

of only 20 % on the experimental plot. Like-

wise, the natural fragmentation produced the

appearance of asexual recruits, with an increase

of 117 % of new colonies. In this way, Guzman

(1991) demonstrated that this type of initiative

is feasible in the region under various conser-

vation schemes such as the implementation of

the same activity within a marine protected

area, and that under the impacts of El Niño,

restoration can be a tool to help the recovery of

these ecosystems.

The role of the Government

in the restoration initiatives

Ecological restoration activities should be

conceptualized as a part of governance, which

integrates collaborative processes of multiple

actors, including the State, the productive sec-

tor and society (Richardson & Lefroy, 2016). In

these processes, the State can adopt different

roles, although often the main contribution

is that it is expected to provide the political

and legal conditions for restoration initiatives

to take place in an articulated and harmoni-

ous manner, avoiding conflicts between actors,

whether the ecosystem in question is used for

consumptive purposes. Flores-Aguilar et al.

(2018), when studying cases of environmental

governance and payments for environmental

services in Latin America, concluded that the

State still has an important role in arbitra-

tion and allocation of roles among the actors

involved. In the case of the policy for “Payment

for Environmental Services” (PES) that Costa

Rica implemented in the 1990s, this consisted

of a system of incentives to landowners for the

protection of forests and promotion of refores-

tation for productive purposes, with results that

have been considered very successful. The par-

ticipating families stated that the PES allowed

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

improvements in the quality of life of the family

nucleus, improvement in the environment at

the farm and community levels, as well as inter-

personal and emotional relationships at the

family and personal levels, beyond the thou-

sands of hectares dedicated to the protection

of the forest, carbon sequestration and water

recharge of these sites (Ortiz-Malavasi, 2004).

In August 2024, the Costa Rican gov-

ernment established Law No. 23555, called

“Incentive for the protection of marine-coast-

al biodiversity.” This law establishes the fund

for payment for environmental services for

marine-coastal ecosystem services, to promote

the conservation, sustainable use and restora-

tion of ecosystems located within the territorial

sea, as well as the exclusive economic zone of

Costa Rica. The groups that can benefit from

this fund will be artisanal fishermen, associa-

tions, fishing or tourism cooperatives, as well

as organized groups within coastal communi-

ties that carry out restoration, conservation or

sustainable use activities of marine resources.

The fund has not yet entered into force and

is awaiting the development of the regulations

that will put it into operation.

In 2018, an interesting case arose in the

implementation of the triangular cooperation

project “Development of an innovative financial

mechanism for coral reef conservation in the

Dominican Republic”, financed by the Region-

al Fund for Triangular Cooperation in Latin

America and the Caribbean of the German

Cooperation Agency (GIZ). In this project, the

Dominican Republic, as both the requesting

and receiving country, expected to develop a

PES mechanism for the conservation of coral

reefs, based on the experience in PES for forest

protection developed by Costa Rica. During

the proposal implementation process, it was

identified that there was an opportunity for

Costa Rica to receive knowledge transfer and

experience in coral reef restoration from the

Dominican Republic as part of the same proj-

ect. Although Costa Rica was already develop-

ing some experiences in coral reef restoration,

the Dominican Republic’s experience in coral

gardening became a pivot for Costa Rica for the

definition of a country strategy on the subject

(Larghi, 2022).

The implementation of this project led

to the enactment of Executive Decree No.

41774-MINAE “Promotion of restoration and

conservation initiatives for the recovery of coral

ecosystems”, which aims to promote the protec-

tion and conservation of reef ecosystems and

their associated species throughout the national

territory. Under this legal framework and with

the implementation of the Costa Rica-Domin-

ican Republic-Germany triangular proposal,

implusing the development of coral restoration

initiatives in Culebra Bay, Golfo Dulce, Sámara

Beach, Manuel Antonio and more recently

Southern Caribbean. Under this initiative,

knowledge was shared between projects being

developed in Costa Rica and coral reef restora-

tion projects in Bayahibe, Dominican Republic,

executed by FUNDEMAR and the Center for

Marine Innovation of the Punta Cana Group

Foundation, both in the Dominican Republic.

Another topic that was considered nec-

essary and developed in this context was a

“Protocol for the restoration of reefs and coral

communities in Costa Rica” (SINAC-GIZ. 2020).

This document aims to provide a legal technical

guidance framework for coral reef restoration

projects in Costa Rica. This guide also arises

from the experience of the Dominican Republic

in the establishment of coral gardening proj-

ects, where the appearance and disappearance

of restoration projects sometimes responded to

economic rather than environmental interests,

which in some cases caused the loss of colonies

and the affectation of natural reefs. Ensuring

that all restoration projects use a technical and

legal basis reduces this risk. This protocol was

implemented through a project in which the

University of Costa Rica, the NGO Raising

Coral Costa Rica, the company Ecodesarrollo

Papagayo, the National System of Conservation

Areas (SINAC) and GIZ participated. This pilot

project consisted of the development of a coral

gardening project in Culebra Bay, North Pacific

of Costa Rica (see section below).

Finally, following the example of the

Dominican Republic, in 2023 the first Coral

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Restoration Consortium was established in

Costa Rica. This consortium seeks to be a

meeting point to discuss the different coral

reef restoration initiatives in Costa Rica. The

consortium is currently being led by the NGO

Raising Coral Costa Rica, and is free adhesion,

promoting that to-date is constituted by differ-

ent national organizations.

Study cases

The studies are presented from north to

south on the Pacific side (Fig. 1), without a

chronological order, only geographically; while

in the Caribbean they only occur in the south-

ern area of said coast. Each geographic area

presents its origin, advances and challenges.

Bahía Culebra

The coral reef restoration project in Cul-

ebra Bay started in 2019, after a cooperation

of more than 20 years between the company

Ecodesarrollo Papagayo and CIMAR, and the

interaction between the GIZ through its Bio-

diversity and Business program (DaBio) and

the National System of Conservation Areas

(SINAC). In addition, the non-profit asso-

ciation Raising Coral Costa Rica joined the

initiative to support all the technical knowledge

developed in Golfo Dulce. It was decided to

work in Culebra Bay due to the great loss of

coral cover that the area had suffered in the last

decade (Alvarado et al., 2018; Fabregat-Malé &

Alvarado, this supplement; Sánchez-Noguera et

al., 2018).

The interaction of the academic, private,

state, international cooperation, and civil soci-

ety sectors established the Culebra Reefs Gar-

dens alliance, where the parties cooperate to

recover the coral reefs in Culebra Bay and

their ecosystem services through ecological

restoration. To achieve this objective, a series

of activities were proposed, from training of

personnel, adequate selection of sites and spe-

cies, installation of structures that serve as a

pilot, continuous monitoring, and evaluating

Fig. 1. Location of coral restoration projects in Costa Rica. A. Culebra Bay; B. Sámara Beach; C. Manuel Antonio; D.

Golfo Dulce; E. Punta Cahuita. Orange rectangle: Current nurseries; Yellow triangle: restoration areas; Green ellipse: Pilot

nurseries; Red polygon: permanent structures.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

how this could be integrated into local respon-

sible tourism activities. Each party has a series

of obligations that are related to the activities

mentioned above.

A five-year work schedule was established

that includes: 1) project planning, 2) the devel-

opment of a theoretical and practical coral reef

restoration workshop with different stakehold-

ers, 3) the installation of nurseries and place-

ment of coral fragments, 4) monitoring and

implementation of a coral gardening pilot,

and 5) scaling-up the number of nurseries

and corals, together with an environmental

education program, in parallel with other

research such as the analysis of coral reproduc-

tion in the nurseries.

Stage one of the project started in April

2019 and stage 2 in August of the same year,

and stage 3 started in September with the

installation of a ropeline-type nursery and two

tree-type nurseries (Fig. 2), both floating, with

585 coral fragments in Playa Jicaro (Fig. 1). In

the ropeline the branching coral Pocillopora

spp. was used, while in the trees the same

branching species was used together with the

massive corals Pavona gigantea, Pavona clavus

Fig. 2. Nursery types and coral species used in the Culebra Reefs Gardens coral restoration project. A. Rope line; B. Tree; C.

A-frame; D. Spider; E. Pocillopora spp.; F. Pavona gigantea; G. Pavona clavus; H. Porites lobata.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

and Porites lobata (Fig. 2) (Fabregat-Malé et

al., 2023). The fragments, from healthy adult

colonies, were collected from different points in

the bay and in the Gulf of Papagayo. In January

2020, 4 spider-like nurseries were set up with

42 fragments each (Combillet et al., 2022), and

in August 2020 two A-frame nurseries were

established on the substrate with approximately

400 coral fragments each (Fabregat-Malé, 2022;

Fabregat-Malé et al., 2024). Only Pocillopora

fragments were used in these nurseries. All

nurseries were monitored for at least one year,

and growth and mortality rates were measured

monthly. At the end of this experimental stage

in the nurseries, it was determined that main-

tenance costs were higher in the latter despite

the higher growth in the floating nurseries.

Likewise, it was quantified that the branched

Pocillopora species showed greater growth and

survival rates than the massive ones. Therefore,

it was decided to advance to the next stage of

the project used only Pocillopora corals in the

substrate nurseries. However, it is established

as a priority to look for mechanisms to improve

the conditions of the massive species.

In this third stage of the project, an experi-

mental planting of 30 colonies of Pocillopora

spp. and 29 fragments of massive corals (P.

gigantea, P. c l av u s and P. lobata) in a pilot area in

Güiri-Güiri (Fig. 1) which had been cultivated

in the nurseries at Playa Jícaro in the previous

months, was also carried out. For the branching

colonies of Pocillopora spp., metal rods were

driven into the calcareous substrate (former

coral framework) to which the transplants were

tied employing plastic gauze (Fig. 3). Each of

the transplanted colonies was labeled with its

own number for later identification during the

following months of monitoring. For the mas-

sive coral transplants, an underwater drill was

used to create holes in the substrate into which

the fragments were placed with marine epoxy

glue. Over the course of a year, where coral

growth, contribution to substrate cover and

Fig. 3. Transplantation of nursery-grown colonies of Pocillopora spp. in A. Güiri-Güiri and B. Pelonas Islands, Culebra Bay,

Gulf of Papagayo. C. Monitoring of corals fragments by phototransects.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

survival rate were measured (Fabregat-Malé,

2022; Fabregat-Malé et al., 2023). After one year

of monitoring, a 100 % survival for Pocillopora

spp. colonies and 58.62 % for massive corals

was determined. The lower survival for the

massive growth species probably corresponds

to the technique used to perform the trans-

plants, since no dead fragments were observed,

but rather a loss of them and of the ceramic disc

on which they were glued (Fabregat-Malé et al.,

2023). It is important to mention that through-

out these three years of research (2019–2021),

sea temperature, salinity, and nutrients were

constantly monitored.

The fourth stage of the project, scaling

up, starts in August 2021 with the increase of

sites and nurseries. In Playa Blanca (Fig. 1) 28

spiders and 5 A-frames were placed, in Güiri-

Güiri nine spiders were established and in Islas

Pelonas five spiders were placed. At the end

of that year, in December, the first regional

CORALMANIA, a joint activity with Hondu-

ras and the Dominican Republic, was carried

out contributing to the transplantation of 300

Pocillopora colonies of to the Güiri-Güiri reef

and 100 to the Pelonas Islands reef. These

colonies have been monitored monthly, and

an increase in coral cover has been determined

in Güiri-Güiri from 4 % to 20 %, and in Pelo-

nas from 5 % to 16 %, as of February 2004 in

both sites (Alvarado et al., in prep.). The El

Niño 2023 phenomenon strongly affected the

colonies, presenting a strong bleaching but rap-

idly recovering (Alvarado et al., in prep.). Both

transplant sites are monitored monthly for fish

diversity and abundance, as well as sea urchin

(Diadema mexicanum), and a gradual increase

in the number of fish species has been observed

from the beginning of monitoring (November

2021: 20 species) to March 2024 (70 species)

(Alvarado et al., in prep.). Currently the project

has 134 spiders with at least 7 000 coral frag-

ments growing in the area and is working on

the reproduction (histological analysis) results

of Pocillopora and on the genetic diversity of the

coral and their endosymbionts.

As a branch derived from Bahía Culebra’s

coral restoration project, the Playa Ocotal’s

restoration program was born, which has a

strong social community component as a char-

acteristic feature and can be classified as a col-

laborative project (Bonney et al., 2009). This

initiative has been carried out between the

CIMAR and the non-profit association Alianza

Mar y Tierra. Alianza Mar y Tierra (AMT)

was founded in 2022 to serve as a platform for

conservation and restoration projects of terres-

trial and marine environments; it’s conformed

by six members of the board of directors and

32 volunteers who collaborate their programs.

Besides Ocotal’s coral reef restoration project,

the AMT organizes monthly activities such as

seabed and beach cleaning, involving dozens of

Ocotal and Playas del Coco residents.

Ocotal’s coral reef has a common history

with others in the North Pacific, in which local

and global stressors acted in synergy and led to

a devastating reduction in live coral cover (Bezy

et al., 2006; Cortés, 2012; Jimenez et al., 2001;

Morales-Ramírez et al., 2001; Navarro-Cerdas,

2013; Sánchez-Noguera, 2012) Specifically, in

Ocotal a significant increase in the coverage of

Caulerpa sertularioides and an overgrowth of

this alga on the colonies of Pocillopora spp. was

reported (Fernández-García et al., 2012). Prior

to the first intervention on the reef, a visit was

carried out where a degraded coral framework

was evident, with isolated colonies of massive

corals and Pocillopora spp. healthy (Fig. 4).

For the intervention on this reef, a citi-

zen science approach has been used; this can

be defined as processes in which community

members who don’t have a formal science edu-

cation engage in scientific research (Cigliano et

al., 2015). In coral reefs, favorable results have

been obtained from incorporating of citizen

scientists in biological monitoring (Forrester et

al., 2015) and ecosystem restoration (Hesley et

al., 2017). Ocotal’s coral reef restoration project

officially began in February 2023 with a com-

munity workshop open to the public, resulting

in the recruitment of volunteers from outside

the AMT. Starting in March 2023, training in

biological monitoring of substrate, fish and

mobile invertebrates began, as well as the place-

ment and cleaning of structures (Fig. 5).

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

In April 2023, the first 15 structures were

placed with fragments from the Jícaro reef and

the Playa Blanca restoration point. Thanks to

funds from the UCR and private donations,

the total number of structures reached 45 by

August 2023. In September 2023, a partnership

was established with the Costa Rican company

Garnier & Garnier Desarrollos Inmobiliarios,

which allowed an increase to 65 structures in

Ocotal and the intervention of another degrad-

ed reef in Punta Cacique. By May 2024, there

are 113 structures distributed at the two resto-

ration points, which represents approximately

2 260 living fragments of Pocillopora spp.

The inclusion of the human component

in the restoration of Ocotal represents a valu-

able opportunity to include social dimensions,

which have been highlighted as essential to

ensure the future of coral reefs (Lamont et

al., 2022). On the other hand, it serves as an

environmental education platform that seeks to

improve the relationship that community mem-

bers have with the marine ecosystems of the

region (Dean et al., 2018) and promotes stabil-

ity over time in this type of projects since there’s

a reduction in the economic costs associated

with restoration process (Hesley et al., 2017).

Samara

Coral reefs in Playa Sámara (Fig. 1) were

confirmed to exist through research conducted

by Cortés & Guzmán (1998), identifying coral

species such as P. lobata, Pocillopora capitata,

Pocillopora damicornis, Porites panamensis,

and Porites (Synarea) rus, this last one being

observed only in 1982 and not since. Recently,

Fig. 4. A. Degraded reef framework prior to the intervention of Ocotal’s reef, Gulf of Papagayo (January 2023). B. Spider

structures used for the restoration of the Ocotal’s coral reef (September 2023).

Fig. 5. A. Training carried out on land on biological monitoring methodologies (March, 2023) and B. at the restoration point

(May, 2023). C. Citizen scientist cleaning the structures (May, 2024).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Psammocora spp. was added to this list (Cortés

et al., 2010).

The coral reefs in Playa Sámara face sig-

nificant anthropogenic pressures from sedi-

mentation and pollution (Pereira-Pérez &

Mairena-Rodríguez, 2011; Pizzimenti et al.,

2011). In response, the Tempisque Conserva-

tion Area (ACT) under the Ministry of Envi-

ronment and Energy (MINAE) enlisted the

Instituto Nacional de Aprendizaje (INA) to

support a coral reef restoration plan at Playa

Sámara, drawing on experience from a similar

project initiated in Golfo Dulce in 2013 (Vargas

et al., 2020). The goal was to restore ecosystem

services crucial for the area’s productive tour-

ism and fishing industries.

The project from September 2017 to

December 2022 involved monthly coral growth

measurements. Work was concentrated on

Chora Island, significantly altered by a 7.6

Mw earthquake in 2012, raising its coastline

by 0.45 m (Linkimer et al., 2013) and modi-

fying coral community structures vertically

and horizontally.

The methodology followed protocols out-

lined by Edwards & Gómez (2007), Rodrí-

guez et al. (2022), Shafir et al. (2006) and

SINAC-GIZ (2020). Initially, the Coral Project

Association trained personnel according to the

SINAC-GIZ Coral Gardening Protocol, (2020).

Exploratory dives off Samara Beach identified

coral colonies suitable for restoration efforts,

including species from Pocillopora spp., Porites

spp., Psammocora spp., and P. g i g a n t e a (Fig. 6).

Different nursery types—platforms, trees, and

ropelines (Fig. 7)—were developed based on

the Coral Gardening protocol for Costa Rica

(SINAC-GIZ, 2020).

Maintenance, monitoring, and cleaning of

structures occurred every fifteen days under

suitable sea conditions. Growth data, including

height, diameter, and colony volume (cm3) were

collected (Robles-Payan et al., 2021). Colonies

that doubled in size were transplanted to the

Cangrejal area (Fig. 1) within the Bay of Playa

Sámara, where ongoing monitoring included

assessments of fish diversity and live colony

counts, following the PRONAMEC protocol

Fig. 6. Coral species found in Samara beach for the coral restoration project: A. Pocillopora spp., B. Porites spp., C. Pavona

gigantea and D. Psammocora spp.

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

for coral reefs (Sistema Nacional de Áreas de

Conservación, [SINAC] 2016).

From 2018 to 2022, 1 404 coral colonies

were harvested, totaling 86 610.7 cm3, with an

average width of 2.80 ± 0.07 cm and length of

5.63 ± 0.28 cm. The growth rate over five years

was 6.72 cm/year. By species, contributions

were 49.7 cm3 (N = 32) for P. g i g a n t e a , 372.3

cm3 (N = 45) for Psammocora spp., 1 210 cm3

(N = 108) for Porites spp., and 84 977 cm3 (N =

1 221) for Pocillopora spp. (Fig. 7). Colony loss

during harvesting was 15 % due to mortality,

detachment, or predation.

Cultivation data indicated rope line struc-

tures were most effective, yielding 8 590.2

g/year, followed by trees at 155.64 g/year.

Pocillopora spp. demonstrated the highest

performance, while platforms showed higher

mortality rates across all coral species cultured

(Fig. 8).

Regarding ichthyofauna in the planting

area, 1 131 individuals representing 23 fish spe-

cies were recorded. Halichoeres dispilus, Thalas-

soma lucasanum, and Abudefduf troschelii were

the most abundant, alongside various inver-

tebrates and vertebrates. In 2023, during the

El Niño phenomenon, 77 coral colonies were

surveyed in the transplant area, with 21 affected

by bleaching.

Fig. 7. Gardening coral structures used in Isla Chora of

Playa Sámara to carry out the harvest of coral reefs in the

Cangrejal sector.

Fig. 8. Coral species grow in three types of gardening in the middle of the water column on Chora Island of Samara Beach.

The trend is that branched or sub-branched species grow better on antenna and clothesline, while massive or sub-massive

species grow better on the platform or antenna.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

The results of coral restoration efforts in

Sámara align with findings from Vargas et al.

(2020) and Rodríguez et al. (2022), emphasiz-

ing the success of branched or sub-branched

species. The total volume produced from 2018

to 2022, 86 610.7 cm3, is comparable to results

from Robles-Payan et al. (2021) over 396 days,

highlighting the need for improved production

efficiency. Hanging structures like ropelines

and trees proved effective for branched species

such as Pocillopora spp., whereas platforms were

suitable for massive species like Porites spp.

Manuel Antonio

Marine Conservation Costa Rica (MCCR),

a non-profit organization, was founded in 2019

by biologists and dive professionals from the

Oceans Unlimited Dive Center in Quepos (Fig.

1), on Costa Rica’s central Pacific coast. In the

same year, the organization received permis-

sion from the SINAC to initiate a pilot study on

coral reef restoration in Manuel Antonio, which

houses the renowned Manuel Antonio National

Park (MANP). As the largest marine protected

area on mainland Costa Rica, this region was

selected due to its historical focus on terrestrial

conservation, while its coral reefs have suffered

degradation over decades from various local

stressors such as river runoff, sedimentation,

and agricultural and domestic contamination

(Sistema Nacional de Áreas de Conservación,

2013). The project involves contributions from

the founding members, core staff biologists,

trained volunteers, international interns, and

local community divers.

Following a planning period that included

surveys for reefs and nursery sites, the pilot

study started in April 2019. Initially, four nurs-

ery sites were chosen: two on the inshore sides

of islands, which offered greater wave protec-

tion, and two more on exposed sites (Fig. 1).

Due to the high-water movement in the area,

fixed table structures were selected for the nurs-

eries. Initially, large nursery tables made from

PVC tubing were used (Fig. 9A). However,

after several months in high-energy environ-

ments, these were scaled down to smaller, more

robust table structures (Fig. 9B). Nurseries were

installed at each of the four sites, at depths of

10–12 m to avoid the shallow surge.

In June 2019, the next phase of the pilot

study involved adding coral fragments to the

nurseries, starting with Pocillopora spp. and P.

gigantea, followed shortly by P. lobata, with 15

samples of each coral species at each site. By

early 2020, the focus shifted to expanding the

Fig. 9. A. Original table nursery, B. Small table nursery, C-D. Coral fusion techniques, E-F. Coral fragmenting techniques.

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

more protected nursery sites, and in July 2020,

the first coral colonies were outplanted to the

restoration site north of Isla Larga (Fig. 1).

The restoration project has expanded sig-

nificantly from 2011 to 2024. As of April 2024,

there are 36 nursery structures. Following the

success of Pavona gigantea, restoration efforts

have included Pavona frondifera and Pavona

duerdeni. In 2022, the project adopted the coral

fusion method (Fig. 9C, Fig. 9D), where frag-

ments of the same genotype coral are adhered

to a plate to fuse and reform larger colonies,

facilitating faster growth and maturity before

outplanting. This method has been tested suc-

cessfully with Pocillopora spp., P. frondifera, and

P. g i g a nt e a , except for P. lobata, which did not

compete well against algal growth.

The project has faced various challenges,

necessitating adaptations in methodology, often

with advice from other restoration groups in

Costa Rica. Key issues and responses include:

1) Algal Growth: Coral fragments are cut to 2-3

cm to better compete against algal growth (Fig.

9E, Fig. 9F); 2) Sustainability and Plastic Use:

Initially, coral fragments were attached using

plastic wall plugs and zip ties (Fig. 10A). In

2021, the project began producing custom discs

with posts made from natural sand and cement,

minimizing plastic use and eliminating plastic

at outplanting sites (Fig. 10B). The current

nursery structures utilize repurposed durable

plastic crates, originally used for delivering

produce in the food industry. These crates,

which would otherwise be discarded and end

up in landfills, are reused in our project, there-

by extending their lifecycle. Additionally, the

crates are economical, easy to install and relo-

cate as needed; 3) Water Movement: Custom-

made coral posts snugly fit into nursery meshes

(Fig. 10C) and are secured to the reef using

marine epoxy (Fig. 10D). Nursery structures

are weighed with cement blocks and anchored

with metal pins; 4) Red Tides: In 2022, reduced

growth was observed due to prolonged red tide

presence. Harvesting and fragmentation of P.

lobata was halted for one year due to its suscep-

tibility to reduced water quality; 5) High Water

Temperatures: During the 2023 El Niño event,

high water temperatures caused bleaching in

shallow outplanted corals (< 8 m depth) and

partial bleaching in nurseries (10–12m depth).

Fragmenting and outplanting were paused due

to concerns abbout additional stress to the cor-

als, and resistant wild colonies were identified

for future harvesting.

As of April 2024, there are 700 fragments in

the two nursery sites. Coral fragments typically

remain in nurseries for 6–12 months before

Fig. 10. A-C. Nursery attachment techniques, D-E. Coral outplants, F. Coral tank.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

outplanting. Due to refined methodologies,

survival rates have improved from 70–80 % to

over 90 %. Post-outplanting survival is approxi-

mately 95 % (Fig. 10E) with most losses due to

algal overgrowth. Over the past four years, 1

750 corals have been outplanted. Monitoring

from January 2019 to January 2024 has shown

an increase in fish species from 18 to 31 and an

increase in coral cover from 1 % to over 3 %.

The project’s future goals include signifi-

cantly increasing outplanting numbers, aiming

to add thousands of new colonies annually. As

part of our restoration project, we are establish-

ing a coral tank in May 2024 (Fig. 10F) to hold

corals before transferring them to the nurseries

and to educate the community on coral reefs

and restoration. Additionally, this provides an

opportunity to explore techniques for poten-

tially utilizing ex-situ tanks to enhance the

production of coral colonies.

MCCR has received permission from

SINAC to install permanent reef structures

in Manuel Antonio Bay to alleviate pressure

on popular snorkel sites and serve as educa-

tional tools for coral conservation. This project,

delayed due to funding issues, is set to begin in

late 2024.

Golfo Dulce

Golfo Dulce (Fig. 1) has been a site of coral

reef restoration for nearly a decade, as part of

an effort to (1) rehabilitate coral reefs that had

been severely impacted by sedimentation and

(2) to do so in ways that promote resilience of

the reefs to climate change.

Beginning in 1990, Cortés (1990a), Cor-

tés (1990b) and Cortés (1992) documented

how sedimentation associated with logging

of the rainforest and road construction had

likely caused severe degradation of most of

the coral reefs in the gulf, and the rarity of

Pocillopora, particularly in comparison with

other Pacific coast reefs of Costa Rica. Fol-

lowing government actions to halt the logging

and other causes of sedimentation, water qual-

ity improved and coral reefs showed signs of

recovery, and Cortés (1992) suggested that

Golfo Dulce would benefit from coral propaga-

tion and reef restoration.

The first attempt to propagate corals in

Golfo Dulce was initiated in 2014 by the INA.

They placed a mid-water platform structure for

coral propagation at about 8 m depth near Playa

Nicuesa (Fig. 1). They deployed 70 fragments

of Porites lobata, Pavona varians and Psam-

mocora stellata and demonstrated good growth

of the fragments over the following 17 months

(Vargas et al., 2020), although the P. lobata and

P. varians fragments died after eight months,

which the authors correlate with the presence

of a red tide. INA has continued the deploy-

ment of coral propagation structures in sev-

eral other locations in the Gulf, and over time

propagated and outplanted coral fragments at

multiple sites.

In 2016, once corals recovered following

a bleaching event that caused significant mor-

tality in Golfo Dulce (Alvarado et al., 2020),

researchers and students from the University of

Costa Rica installed two mid-water tree struc-

tures, also near Playa Nicuesa, to pilot test the

efficacy of coral propagation and outplanting

methods in the gulf, as well as the knowledge

and perceptions of residents about coral reefs

(Villalobos-Cubero et al., 2023). The latter was

a major part of the restoration effort, follow-

ing the philosophy of Suding et al. (2015) hat

outlined the four main components of success-

ful restoration programs, including not only 1)

assessment of the sites environmental past and

future, 2) consideration of the ecological prin-

ciples, and 3) commitment to sustained effort,

but also 4) strong engagement of local com-

munities. Raising Coral has invested in local

communities of Golfo Dulce primarily through

a coral gardener training and employment

program that includes 20 local people, but also

through active engagement of local government

officials and the public, with a network of over

60 national and international allies.

Initially, fragments of Porites evermanni/

lobata, P. g i g a n t e a , and Pocillopora spp. were

propagated, outplanted and monitored for

several years, testing species responses to a

variety of techniques (Kleypas et al., 2021b;

16 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Villalobos-Cubero, 2019). In 2019 the project

evolved into a formal effort with the establish-

ment of the NGO Raising Coral Costa Rica,

and operations gradually expanded to include

more structures (currently 11 trees, five rope

nurseries, Fig. 11) with a capacity of nearly

2000 coral fragments. In 2021, Raising Coral

became an official user of the Mars reef res-

toration system (MARRS; Smith et al., 2021)

and has so far deployed 95 reef stars with a

total capacity of about 1 400 fragments. Since

the start of the project, all donor coral colonies

have been marked in the field and all fragments

and outplanted colonies have been tracked with

a donor ID, except when using fragments of

opportunity or when sampling Psammocora

stellata, which at some sites exhibits a growth

habit of loosely interlocking benthic cover rath-

er than distinct colonies.

Raising Coral works with the 9 reef-build-

ing coral species in Golfo Dulce, but so far

has focused on rebuilding the populations of

Fig. 11. Propagation and outplanting techniques at the Golfo Dulce site. A. donor colony of Pocillopora spp.; B. midwater

trees; C. vertical rope (experimental); D. MARRS reef stars; E. rope nurseries; F. monitoring of Pocillopora spp. on reef stars;

G. propagated colony of Pavona frondifera; H. direct outplants of Porites spp. to “reskin” older Porites surface; I. cluster of

fused Pocillopora spp. Outplants; J. fused outplants of Pavona gigantea; K. fused outplants of Porites spp.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Pocillopora species in Golfo Dulce. Pocillopora

colonies were rare in Golfo Dulce in 2016, and

in fact, over the course of several years fewer

than 20 natural colonies were encountered that

were large enough to sample for propagation.

Most of these were found in waters between

7–12 m deep, except for several large Pocillopora

colonies and recruits found at much shallower

depths. Fragments of Pocillopora (1–2 cm in

length) grew well in every type of structure to

outplantable colonies of 10–14 cm diameter

in 8–12 months. Before outplanting, several

fragments were obtained from each colony to

continue the propagation in the nursery.

Over time, about 3000 thousand Pocillo-

pora colonies were propagated and outplanted

in multiple areas, with the majority at the

northernmost reef sites of Golfo Dulce. The

outplanting design included placing 8–15 clon-

al colonies in patches to increase the probability

of colony fusion, and then arranging patches

to maximize cross-fertilization amongst dif-

ferent donors. The colonization of some of

the first outplanted colonies of Pocillopora by

cryptofauna was investigated by Chomitz et al.

(2023a) and Chomitz et al. (2023b).

The number of propagated and outplanted

Pocillopora colonies grew progressively through

early 2023, with interruptions caused by harm-

ful algal blooms (“red tides”) and bleaching.

Red tides became increasingly common over

that period and particularly in late 2020, caus-

ing the mortality of 11 % of the Pocillopora

fragments in the nursery; despite this event

by 2022, the average survival percentage for

nursery fragments was 98 %, and outplants sur-

vival was 94 %. Later, in early 2023, a red tide

followed by a record marine heat wave caused

the mortality of 90–95 % of the Pocillopora

colonies, particularly outplants at shallower

depths. Coral fragments in the MARRS struc-

tures survived best, likely due to multiple fac-

tors (Sandoval et al., 2022).

The timeline of the restoration effort in

Golfo Dulce provides an example for sustain-

ing corals and reefs in a warming world (Fig.

12), i.e., planning for expected setbacks due to

environmental threats such as harmful algal

Fig. 12. Changes in the Raising Coral nursery capacity (not including fragments deployed using MARRS), number of

Pocillopora fragments in propagation, and total number of Pocillopora outplants between 2016 and 2024. Projected values are

shown through June 2026, using modest rates of Pocillopora propagation.

18 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

blooms and climate change (as well as coral

disease). Unlike in other Pacific case studies in

Costa Rica, the slow start to Pocillopora propa-

gation was hindered by the rarity of colonies

large enough for fragmenting and a small team

and nursery capacity. Over time, the capacity

and efficiency of propagation and outplant-

ing steadily increased, and despite the severe

mortality of 2023, the ability to recover over

the next two years is now greatly enhanced by

(1) the fact that there are still more colonies of

Pocillopora than when the project began, (2)

an existing large nursery capacity, and (3) the

experience of the community members work-

ing as coral gardeners, which greatly increases

the efficiency of the work. Our team also found

that the survival and growth of some spe-

cies improve when fragments are propagated

directly on the reef rather than in a nursery

(Sandoval et al., 2022). Tracking fragments by

donor has also yielded useful information, such

as identifying genotypes that are more resistant

to bleaching or more likely to recover following

bleaching. In anticipation of the 2023 bleach-

ing event, Raising Coral implemented a Coral

Bleaching Contingency Plan that included sev-

eral actions to minimize coral loss: lowering

structures in the nursery, shading experiments,

and mainly distributing fragments from each of

the Pocillopora genotypes across a wide array

of sites. Such in situ coral banking is a good

alternative when ex situ coral banks are not

available, and at the same time helps identify

locations that can serve as coral refugia during

future marine heat waves.

Southern Caribbean

Coral reef restoration in Costa Rica’s

Southern Caribbean began in 2021, driven by

the interest of Cahuita’s National Park marine

management program in assessing the viability

of coral restoration techniques as a manage-

ment tool to achieve their conservation goals.

These goals include (1) conserving an ecologi-

cally representative sample of Caribbean reef

and seagrass systems, along with their associ-

ated fauna and flora, and (2) contributing to

adaptation and mitigation measures for marine

and coastal biodiversity in response to the

impacts of climate change in the Costa Rican

Caribbean. Consequently, Raising Coral Costa

Rica and ACLAC-SINAC established a joint

restoration project.

The reef system at Cahuita National Park

(PNC) (Fig. 1) is the most developed and

extensive (600 ha) reef habitat in the country,

yet it is also one of the most deteriorated and

threatened. Since the early 1980s, Jorge Cortés

and colleagues have documented a significant

decline in live coral cover, from a baseline of

40 % in the early 1980s to just 10 % in the early

1990s (Cortés, 1994; Cortés et al., 2010). This

decline is primarily attributed to sustained

nutrient runoff, high sedimentation stress, and

the deleterious effects of bleaching events over

the years, notably in 1983, 1992, 1995, 1998,

2005 (Cortés, 2016; Cortés & Jimenez, 2003a;

Cortés et al., 2010; Jiménez, 2001), and 2020,

2023 (Quezada-Perez et al., 2023).

More recently, Quezada-Perez et al. (2023)

reported around 12–25 % reef cover, which is

significantly lower than the original baseline

but somewhat higher than in the 1990s. The

same study found an increasing trend in algae

cover, rising from 37 % in 2003 to 70–73 %

in 2023. These findings suggest a potential

phase shift from a coral-dominated reef to an

algae-dominated reef. This shift results in low

levels of key ecological indicators, such as fish

diversity and reef rugosity, leading to the loss of

critical ecosystem services.

These circumstances underscore the neces-

sity for sustained implementation of resilience-

based coral restoration efforts, which aim to

reduce stressors to reefs and enhance ecological

and social processes that improve Costa Rica’s

Caribbean reef health, as Cortés (2016) and

Cortés et al. (2010) stated.

As of April 2021, three types of nurseries

were established and tested for the growth and

survivorship of reef-building species: symmet-

rical brain coral (Pseudodiploria strigosa), fused

staghorn (Acropora prolifera), and endangered

elk horn (Acropora palmata). The nurseries

included tree-like and rope-type structures for

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

the branching A. prolifera and a mid-water

floating platform for culturing the massive

P. strigosa. Acropora palmata fragments were

tested in both the tree and platform nurser-

ies (Fig. 12). These nurseries were installed at

depths of 2–4 m, with fragments sourced from

donor colonies selected from different reef

habitats. Sea surface temperature was recorded

using autonomous HOBO data loggers and the

Aqualink Sofar Buoy. Trained SINAC officials

and volunteers played a critical role, assisting

with nursery maintenance and fragment moni-

toring. Survival rates varied between species

and cultivation techniques. For A. palmata,

fragments placed on tree structures showed

survival rates ranging from 56 % to 100 %,

depending on the donor colony. Notably, frag-

ments from colony D1 exhibited 100% survival

on the tree structure and 38.5 % when placed

on “cement cookies” in the platform, likely due

to sedimentation stress based on observations

of sediment accumulation on the exposed area

of the cement cookies. In contrast, P. strigosa

fragments achieved 100 % survival in the plat-

form structure. Acropora prolifera fragments

had survival rates of 90 % on tree structures

and 100 % on rope nurseries. On platforms, A.

palmata fragments grew from 38 cm² to 46.9

cm² (Δ8.9 cm²/0.06 cm² day-1), while on the

tree nursery they grew from 48.9 cm² to 72.8

cm² (Δ23.8 cm² /0.18 cm² day-1). Pseudodiplo-

ria strigosa grew from 20.07 cm² to 21.29 cm²

(Δ1.02 cm²/0.007 cm² day-1), reflecting their

slower growth rates. In the case of A. prolifera

fragments on ropes grew from 9.6 cm to 25.4

cm (Δ15.8 cm/0.12 cm day-1) in linear tissue

extension, while those on tree branches grew

from 7.2 cm to 26.8 cm (Δ19.6/0.15 cm day-1),

confirming their rapid growth potential (Lir-

man et al., 2010) (Fig. 13).

Building upon these initial results, dur-

ing the rest of 2022 and early 2023, more coral

trees were placed, the size and material of the

cookies were reduced, and rope nurseries were

installed among the remaining pilings of an

old pier near the Perezoso area (Fig. 1), with a

capacity to produce between 600–1 200 frag-

ments. However, despite the positive response

of the fragments to the cultivation techniques,

modifications had to be made to the cultivation

strategy and project management. Given Costa

Rica’s high wave conditions in the Caribbean

(Lizano, 2007), mid-water floating structures

suffered structural damage. Therefore, fixed

structures to the bottom, such as reef-stars

frames, were tested. Concurrently, an adaptive

management model was adopted, enabling a

flexible decision-making strategy incorporat-

ing experimentation, monitoring, and iteration,

allowing the opportunity to adjust over time

and deal with uncertainties (Anthony et al.,

2015; McLeod et al., 2019).

Several trials are conducted to determine

the most cost-effective outplanting technique

suited for site conditions before initiating mass

outplanting. In June 2022, a pilot outplanting

of four clonal micro-fragments (~ 5 cm²) of

A. palmata was conducted, utilizing nursery-

grown fragments initially harvested from donor

colony D14 at Kawe (Fig. 1) (Papke et al., 2021).

The outplanting took place at the Eduardo reef

patch (9°44’15.2”N, 82° 48’ 21.4”W). By Decem-

ber 2023, the fused basal area of the fragments

had expanded by over 1 000 %, growing from

19.25 cm² to 213.2 cm² (Δ 193.95 cm2/0.35 cm2

day-1). Notably, both the outplanted fragments

and their parental colony remained unaffected

by the 2023 bleaching event, suggesting that

the bleaching resilience of donor colony D14

is a stable trait that persists through the cul-

ture process and transplantation to a new site

(Anthony et al., 2015; Baums et al., 2019).

In early September 2022, a reef-star cluster

(N = 4) was deployed at the Kawe site, holding

49 fragments of A. palmata and 18 fragments

of A. prolifera. Almost a year later, by August

2023, A. palmata fragments had increased 2.4-

fold in size, growing from 28 cm² to 68 cm² (Δ

40 cm2/0.11 cm2 day-1). Similarly, A. prolifera

fragments grew from 10 cm² to 46 cm² during

the same period (Δ 36 cm2/0.10 cm2 day-1).

Prior to the 2023 bleaching event, the corals

exhibited a 100 % survival rate. However, fol-

lowing the event, survival rates decreased to

46 % for A. palmata and 44 % for A. prolifera

(Fig. 14). These observations highlight the

20 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

initial success of the reef-star method in pro-

moting coral growth and survival, though the

notable reduction in survival rates post-bleach-

ing event underscores the vulnerability of these

coral species to thermal stress.

Later that same month (September 2022),

10 fragments of A. prolifera were outplanted

in the Puerto Vargas sector. Small fragments

(< 4 cm²) were adhered to the substrate using

a nail and a plastic zip tie. By April 2024, the

fragments on average had grown from 2.15 cm²

to 23.26 cm², reflecting an increase of 21.11 cm²

(0.04 cm² day-1). Survival rate of the fragments

after year one was 80 %.

Coral spawning observations were con-

ducted around Cahuita, Punta Uva, and

Fig. 13. Nursery types and growth of different coral species. A. Tree structure with a detailed view of an Acropora palmata

fragment; B. Platform structure with a detailed view of Pseudodiploria strigosa fragments; and C. Section of a clothesline with

a detailed view of an Acropora prolifera fragment growing on the rope.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Fig. 14. Outplanting techniques tested in 2022–2023 at Cahuita site. A. A. palmata microfragments attached to substrate with

marine epoxy, B. same fragments fused and developing protobranches, C. fragments of A. prolifera placed at Kawe reef star

cluster, D. same fragments showing growth over the structure, E. an A. prolifera fragment attached to the substrate using nail

and zip tide, F. the same fragment showing positive growth.

22 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Manzanillo from 2022 to 2023. Pseudodiplo-

ria strigosa was observed to spawn in August

2022 in Cahuita and in September 2023 in

both Punta Uva and Manzanillo. Acropora

palmata was also observed to spawn in Sep-

tember 2023 in both Punta Uva and Manzanillo

(Fig. 15). These observations are significant as

they mark the initial step towards studying the

reproductive activity of reef-building corals

and the potential application of assisted sexual

reproduction techniques in the Costa Rican

Caribbean.

The 2023 bleaching event significantly

impacted both the reef and restoration efforts,

reducing the amount of propagative mate-

rial available for transplantation by approxi-

mately 40 %. Currently, efforts are focused

on replenishing coral nurseries and increasing

the frequency of transplantation. These efforts

incorporate lessons learned during the con-

tingency plan implemented before and during

the bleaching event. For instance, there is now

an emphasis on including coral colonies that

demonstrated resistance to bleaching or exhib-

ited a high recovery rate post-event. Contin-

ued research and community involvement are

essential to sustaining and scaling these efforts

and achieving long-term conservation goals in

the face of ongoing environmental challenges.

Challenges, future needs and

concluding remarks

Over nearly a decade of coral restoration

efforts in Costa Rica, work has spanned virtu-

ally all coral regions across the country (Cortés

& Jiménez, 2003a; Cortés & Jimenez, 2003b).

Initiatives have covered diverse areas from

the coastal upwelling zones in the northern

Pacific to the thermally stable environments

of the South Pacific and Caribbean. Restora-

tion efforts have focused on ten coral species,

seven in the Pacific (Pavona clavus, Pavona

gigantea, Pavona frondifera, Pavona duerdeni,

Porites lobata/evermanni, Psammocora stellata,

and Pocillopora spp.) and three in the Carib-

bean (Acropora palmata, Acropora prolifera,

and Pseudiploria strigosa), with notable success

observed in branched species like Pocillopora.

However, challenges persist with massive coral

species, particularly concerning nursery and

transplant survival rates. Various types of

nurseries, including both water-column and

substrate-attached designs, have been utilized

successfully, tailored to the specific oceano-

graphic conditions of each site. This has result-

ed in managing nearly 19 000 coral fragments

across nurseries and transplant areas through-

out the project’s duration.

Fig. 15. Spawning observations during the project’s lifespan. A. P. strigosa releasing gamete bundles in Cahuita National Park

in August 2021, B. A. palmata releasing gamete bundles in Manzanillo in September 2023.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Despite geographic and oceanographic dis-

tinctions, common factors have characterized

these projects. They have emphasized local

engagement and perception of coral reefs, fos-

tered intersectoral public-private collaborations

for financial and human resources, and operat-

ed within established governmental regulatory

frameworks conducive to restoration efforts

(Palou-Zuñiga et al., 2023). All projects face

vulnerabilities such as extreme temperature

variations from El Niño events and detrimental

impacts of algal blooms, significantly affecting

coral fragment survival in nurseries.

To enhance conditions at coral reefs and

restoration sites, urgent measures include map-

ping and marine spatial planning to delineate

and safeguard these areas from conflicting

activities. Continued monitoring of reef condi-

tions and environmental education involving

local organizations are also crucial. Improving

conditions for cultivating massive coral species

and establishing genetic banks across nurs-

ery sites are essential. Furthermore, advanc-

ing assisted reproduction strategies through

enhanced understanding of local species’ sexual

reproduction and adaptation to warming events

via metagenomics and stable isotopes stud-

ies is imperative. Genetic analyses should be

conducted to identify resilient genotypes for

propagation. Educational outreach should be

enhanced, integrating coral restoration efforts

into local tourism and educational programs

to foster community involvement and support.

Additionally, further research into coral diseas-

es, particularly in the Eastern Tropical Pacific

where studies are limited, remains a priority.

Ethical statement: the authors declare that

they all agree with this publication and made

significant contributions; that there is no con-

flict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

ACKNOWLEDGEMENTS

We are very grateful to all the volunteers

who have worked on the different restoration

projects carried out in Costa Rica, without

their contribution and commitment this type

of initiative could not be carried out. Likewise,

we are very grateful to all the state institutions

that have supported this type of initiative, espe-

cially the National Learning Institute (INA), the

University of Costa Rica (UCR), the National

University (UNA) and the National System

of Conservation Areas (SINAC). International

cooperation was an asset for Costa Rica to grow

in the search for the recovery of coral reefs,

especially the support of the German Develop-

ment Cooperation Agency (GiZ), and with it

all the private tour operator companies and

hotels that have believed in and supported these

projects. All restoration projects presented here

have research permits endorsed by SINAC.

REFERENCES

Alvarado, J. J., Beita-Jiménez, A., Mena, S., Fernández-

García, C., Cortés, J., Sánchez-Noguera, C., Jiménez,

C., & Guzmán, A. G. (2018). Cuando la conservación

no puede seguir el ritmo del desarrollo: Estado de

salud de los ecosistemas coralinos del Pacífico Norte

de Costa Rica. Revista de Biología Tropical, 66(S1),

280–308. https://doi.org/10.15517/rbt.v66i1.33300

Alvarado, J. J., Sánchez-Noguera, C., Arias-Godínez, G.,

Araya, T., Fernández-García, C., & Guzmán, A. G.

(2020). Impact of El Niño 2015-2016 on the coral

reefs of the Pacific of Costa Rica: the potential role

of marine protection. Revista de Biología Tropical,

68(S1), 271–282. https://dx.doi.org/10.15517/rbt.

v68is1.41190

Anthony, K. R., Marshall, P. A., Abdulla, A., Beeden,

R., Bergh, C., Black, R., Eakin, C. M., Game, E. T.,

Gooch, M., Graham, N. A. J., Green, A., Heron, S.

F., van Hooidonk, R., Knowland, C., Mangubhai, S.,

Marshall, N., Maynard, J. A., McGinnity, P., McLeod,

E., … Wear, S. (2015). Operationalizing resilience for

adaptive coral reef management under global envi-

ronmental change. Global Change Biology, 21, 48–61.

https://doi.org/10.1111/gcb.12700

Anthony, K., Bay, L. K., Costanza R., Firn, J., Gunn, J.,

Harrison, P., Heyward, A., Lundgren, P., Mead, D.,

Moore, T., Mumby, P. J., van Oppen, M. J. H., Robert-

son, J., Runge, M. C., Suggett, D. J., Schafelke, B.,

Wachenfeld, D., & Walshe, T. (2017). New interven-

tions are needed to save coral reefs. Nature Ecology

24 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

& Evolution, 1, 1420–1422. https://doi.org/10.1038/

s41559-017-0313-5

Baums, I. B., Baker, A. C., Davies, S. W., Grottoli, A. G.,

Kenkel, C. D., Kitchen, S. A., Kuffner, I. B., LaJeunes-

se, T. C., Matz, M. V., Miller, M. W., Parkinson, J. E., &

Shantz, A. A. (2019). Considerations for maximizing

the adaptive potential of restored coral populations

in the western Atlantic. Ecological Applications, 29(8),

e01978. https://doi.org/10.1002/eap.1978

Bayraktarov, E., Banaszak, A. T., Montoya Maya, P., Kleypas,

J., Arias-González, J. E., Blanco, M., Calle-Triviño J.,

Charuvi, N., Cortés-Useche, C., Galván, V., García-

Salgado, M. A., Gnecco, M., Guendulain-García, S.

D., Hernández-Delgado, E. A., Marín-Moraga, J. A.,

Maya, M. F., Mendoza-Quiroz, S., Mercado-Cervan-

tes, S., Morikawa, M., … Frías-Torres, S. (2020). Coral

reef restoration efforts in Latin American countries

and territories. PLoS ONE, 15(8), e0228477. https://

doi.org/10.1371/journal.pone.0228477

Bayraktarov, E., Stewart-Sinclair, P. J., Brisbane, S., Bos-

tröm-Einarsson, L., Saunders, M. I., Lovelock, C.E.,

Possingham, H. P., Mumby, P. J., & Wilson, K. A.

(2019). Motivation, successs, and cost of coral reef

restoration. Restoration Ecology, 27(5), 981–991.

https://doi.org/10.1111/rec.12977

Bezy, M. B., Jimenez, C., Cortés, J., Segura, A., León, A.,

Alvarado, J. J., & Guillén, C. (2006). Contrasting

Psammocora dominated coral communities in Costa

Rica, tropical eastern Pacific. Proceedings of 10th

International Coral Reef Symposium, 376–381.

Bonney, R., Ballard, H., Jordan, R., McCaliie, E., Phillips,

T., Shirk, J., & Wilderman, C. C. (2009). Public par-

ticipation in scientific research: defining the field and

assessing it’s potential for informal science education. A

CAISE Inquiry Group Report [Technical report]. Cen-

ter for Advancement of Informal Science Education.

Bostrom-Einarsson, L., Bayraktarov, E., Ceccarelli D.,

Cook, N., Ferse, S. C. A., Hancock, B., Harrison, P.,

Hein, M., Shaver, E., Smith, A., Suggett, D., Stewart-

Sinclair, P. J., Vardi, T., & McLeod, I. (2020). Coral

restoration – A systematic review of current methods,

successes, failures and future directions. PLoS One,

15(1), e0226631. https://doi.org/10.1371/journal.

pone.0226631

Bowden-Kerby, A. (1997). Coral transplantation in shelte-

red habitats using unattached fragments and cultured

colonies. Proceedings 8th International Coral Reef

Symposium, 2, 2063–2068.

Bowden-Kerby, A., Quinn, N., Stennet, M., & Mejia, A.

(2005). Acropora cervicornis restoration to support

coral reef conservation in the Caribbean. NOAA

Coastal Zone, 5, 7–10.

Campos, J., & Guzmán, H. (1986). An artificial reef for

artisanal fisheries enhancement in Costa Rica. NAGA,

9(2), 21.

Campos, J., & Gamboa, C. (1989). An artificial tire reef in a

tropical marine system: A management tool. Bulletin

of Marine Science, 44(2), 757–766.

Chomitz, B. R., Kleypas, J. A., Cortés, J., & Alvarado, J. J.

(2023a). Succesion of sessile benthic community at a

coral reef restoration site. Revista de Biología Tropi-

cal, 71(S1), e54881. https://doi.org/10.15517/rev.biol.

trop..v71iS1.54881

Chomitz, B. R., Kleypas, J. A., Cortés, J., & Alvarado, J. J.

(2023b). Change in the composition of fauna asso-

ciated with Pocillopora spp. (Scleractinia, Pocillopo-

ridae) following transplantation. Revista de Biología

Tropical, 71(S1), e54882. https://doi.org/10.15517/rev.

biol.trop..v71iS1.54882

Cigliano, J. A., Meyer, R., Ballard, H. L., Freitag, A., Phi-

llips, T. B., & Wasser, A. (2015). Making marine

and coastal citizen science matter. Ocean & Coastal

Management, 115, 77–87. https://doi.org/10.1016/j.

ocecoaman.2015.06.012

Combillet, L., Fabregat-Malé, S., Mena, S., Marín-Moraga, J.

A., Gutierrez, M., & Alvarado, J. J. (2022). Pocillopora

spp. growth analysis on restoration structures in an

Eastern Tropical Pacific upwelling area. PeerJ, 10,

e13248. https://doi.org/10.7717/peerj.13248

Cortés, J. (1990a). Coral reef decline in Golfo Dulce, Costa

Rica, eastern Pacific: anthropogenic and natural dis-

turbances (Ph.D. Dissertation). University of Miami,

United States of America.

Cortés, J. (1990b). The coral reefs of Golfo Dulce, Costa

Rica: distribution and community structure. Atoll

Reserach Bulletin, 344, 1–37.

Cortés, J. (1992). Los arrecifes coralinos de Golfo Dulce,

Costa Rica: aspectos ecológicos. Revista de Biología

Tropical, 40 (1), 19–26.

Cortés, J. (1994). A reef under siltation stress: a decade

of degradation. In Ginsburg, R. N. (Compiler), Pro-

ceedings of the Colloquium on Global Aspects of

Coral Reefs: Health, Hazards and History, 1993. (pp.

240–246). Miami, Florida, U.S.A.: RSMAS, University

of Miami.

Cortés, J. (2012). Historia de la investigación marino-

costera en Bahía Culebra, Pacífico Norte, Guanacas-

te, Costa Rica. Revista de Biología Tropical, 60(S2),

19–37. https://doi.org/10.15517/rbt.v60i2.19961

Cortés, J. (2016). The Caribbean coastal and marine ecosys-

tems. In M. Kappelle (Ed.), Costa Rican Ecosystems

(pp. 591–617). University of Chicago Press.

Cortés, J., & Guzman, H. (1998). Organismos de los arreci-

fes coralinos de Costa Rica: descripción, distribución

geográfica e historia natural de los corales zooxante-

lados (Anthozoa: Scleractinia) del Pacífico. Revista de

Biología Tropical, 46, 55–92. https://doi.org/10.15517/

rbt.v46i1.19353

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Cortés, J., & Jiménez, C. E. (2003a). Past, present and future

of the coral reefs of the Caribbean coast of Costa Rica.

In J. Cortés (Ed.), Latin American Coral Reefs (pp.

223–239). Elsevier Science B.V.

Cortés, J., & Jiménez, C. E. (2003b). Corals and coral reefs

of the Pacific of Costa Rica: history, research and sta-

tus. In J. Cortés (Ed.), Latin American Coral Reefs (pp.

361–385). Elsevier Science B.V.

Cortés, J., Jiménez, C., Fonseca, A., & Alvarado, J. J. (2010).

Status and conservation of coral reefs in Costa Rica.

Revista de Biología Tropical, 58(S1), 33–50. https://

doi.org/10.15517/rbt.v58i1.20022

Dean, A. J., Church, E. K., Loder, J., Fielding, K. S., & Wilson,

K. A. (2018). How do marine and coastal citizen scien-

ce experiences foster environmental engagement?

Journal of Environmental Management, 213, 409–416.

https://doi.org/10.1016/j.jenvman.2018.02.080

Edwards, A., & Gómez, E. (2007). Reef Restoration Concepts

& Guidelines: making sensible management choices in

the face of uncertainty [Technical report]. Coral Reef

Targeted Research & Capacity Building for Manage-

ment Program.

Fabregat-Malé, S. (2022). Restaurando el arrecife: estrategias

para el crecimiento de corales en viveros in situ en

Bahía Culebra, Pacífico Norte de Costa Rica [Tesis de

Maestría en Biología], Universidad de Costa Rica, San

Pedro, Costa Rica.

Fabregat-Malé, S., Mena, S., & Alvarado, J. J. (2023). Nur-

sery-reared coral outplanting success in an upwelling-

influenced area in Costa Rica. Revista de Biología

Tropical, 71(S1), e54879. https://doi.org/10.15517/rev.

biol.trop..v71iS1.54879

Fabregat-Malé, S., Mena-González, S., Quesada-Perez, F.,

& Alvarado, J. J. (2024). Testing the feasibility of coral

nurseries in an upwelling area in the North Pacific of

Costa Rica. Frontiers in Marine Sciences, 11, 1400026.

https://doi.org/10.3389/fmars.2024.1400026

Fernández-García, C., Cortés, J., Alvarado, J. J., & Nivia-

Ruiz, J. (2012). Physical factors contributing to the

benthic dominance of the alga Caulerpa sertularioides

(Caulerpaceae, Chlorophyta) in the upwelling Bahía

Culebra, north Pacific of Costa Rica. Revista de Biolo-

gía Tropical, 60(S2), 93–107.

Flores-Aguilar, A., Aguilar-Robledo, M., Reyes-Hernández,

H., & Guzmán-Chávez, M. G. (2018). Gobernanza

ambiental y pagos por servicios ambientales en Amé-

rica Latina. Sociedad y Ambiente, 16, 7–31.

Forrester, G., Baily, P., Conetta, D., Forrester, L., Kintzing,

E., & Jarecki, L. (2015). Comparing monitoring data

collected by volunteers and professionals shows that

citizen scientists can detect long-term change on

coral reefs. Journal for Nature Conservation, 24, 1–9.

https://doi.org/10.1016/j.jnc.2015.01.002

Gann, G. D., McDonald,T., Walder, B., Aronson, J., Nelson,

C. R., Jonson, J., Hallett, J. G., Eisenberg, C., Guari-

guata, M. R., Liu, J., Hua, F., Echeverría, C., Gonzales,

E., Shaw, N., Decleer, K., & Dixon, K. W. (2019).

International principles and standards for the prac-

ticeof ecological restoration. Second edition. Restora-

tion Ecology, 27(S1), S1–S46. https://doi.org/10.1111/

rec.13035

Garza, J. (2019). Arrecifes artificiales ayudarán a salvar

la ecología marina. La República. https://www.lare-

publica.net/noticia/arrecifes-artificiales-ayudaran-a-

salvar-ecologia-marina

Guzman, H. (1991). Restoration of coral reefs in Pacific

Costa Rica. Conservation Biology, 5, 189–195. https://

doi.org/10.1111/j.1523-1739.1991.tb00123.x

Guzman, H. M., & Cortés, J. (1989). Growth rates of eight

species of scleractinian corals in the Eastern Paci-

fic (Costa Rica). Bulletin of Marine Science, 44(3),

1186–1194.

Guzman, H., Campos, J., Gamboa, C., & Bussing, W. A.

(1988). Un arrecife artificial de llantas: Su potencial

para el manejo de pesquerías. Anales del Instituto de

Ciencias del Mar y Limnología, Universidad Autónoma

de México, 15, 249–254.

Hagedorn, M., Page, C. A., O’Neil, K. L., Flores, D. M.,

Tichy, L., Conn, T., Chamberland, V. F., Lager, C.,

Zuchowicz, N., Lohr, K., Blackburn, H., Vardi, T.,

Moore, J., Moore, T., Baums, I. B., Vermeij, M. J.

A., & Marhaver, K. L. (2021). Assisted gene flow

using cryopreserved sperm in critically endangered

coral. Proceeding of the National Academy of Scien-

ces 118(38), e2110559118. https://doi.org/10.1073/

pnas.2110559118

Harriott, V. J., & Harrison, P. L. (1984). Methods to accelera-

te the recolonization of corals in damaged reef systems

[Technical report]. Great Barrier Reef Marine Park

Authority, Australian Government.

Hesley, D., Burdeno, D., Drury, C., Schopmeyer, S., &

Lirman, D. (2017). Citizen science benefits coral reef

restoration activities. Journal for Nature Conservation,

40, 94–99. https://doi.org/10.1016/j.jnc.2017.09.001

Hoegh-Guldberg, O., Skirving, W., Dove, S. G., Spady, B. L.,

Norrie, A., Geiger, E. F., Liu, G., De La Cour, J. L., &

Manzello, D. P. (2023). Coral reefs in peril in a record-

breaking year: Climate change and its impacts on

coral reefs have reached unchartered territory. Scien-

ce, 382(6676), 1238–1240. https://doi.org/10.1126/

science.adk4532

Hughes, T. P., Anderson, K. D., Connolly, S. R., Heron, S.

F., Kerry, J. T., Lough, J. M., Baird , A. H., Baum, J.

K., Berumen M. L., Bridge, T. C., Claar, D. C., Eakin,

C. M., Gilmour, J. P., Graham, N. A. J., Harrison, H.,

Hobbs, J. P. A., Hoey, A. S., Hoogenboom, M., Lowe,

... Wilson, S. K. (2018). Spatial and temporal patterns

of mass bleaching of corals in the Anthropocene.

26 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Science, 359(6371), 80–83. https://doi.org/10.1126/

science.aan8048

Infobae. (18 de febrero de 2021). Instalan “arrecifes artifi-

ciales” en las aguas de Costa Rica para darle hábitat

a los animales. https://www.infobae.com/america/

carbononews/2021/02/18/instalan-arrecifes-artificia-

les-en-las-aguas-de-costa-rica-para-darle-habitat-a-

los-animales/

Inter Press Service. (1999). Costa Rica: Arrecifes artificia-

les regeneran hábitats marinos. https://ipsnoticias.

net/1999/07/costa-rica-arrecifes-artificiales-regene-

ran-habitats-marinos/

Ishida-Castañeda, J., Pizarro, V., López-Victoria, M., &

Zapata, F. A. (2020). Coral reef restoration in the Eas-

tern Tropical Pacific: feasibility of the coral nursery

approach. Restoration Ecology, 28(1), 22–28. https://

doi.org/10.1111/rec.13047

Jiménez, C. (2001). Bleaching and mortality of reef

organisms during a warming event in 1995 on the

Caribbean coast of Costa Rica. Revista de Biología

Tropical, 49(S2), 233–238.

Jiménez, C., Cortés, J., León, A., & Ruíz, E. (2001). Coral

bleaching and mortality associated with the 1997-98

El Niño in an upwelling environment in the eastern

pacific (Gulf of Papagayo, Costa Rica). Bulletin of

Marine Science, 69, 151–169.

Kleypas, J., Allemand, D., Anthony, K., Baker, A. C., Beck,

M. W., Hale, L. Z., Hilmi, N., Hoegh-Guldberg, O.,

Hughes, T., Kaufman, L., Kayanne, H., Magnan, A.

K., Mcleod, E., Mumby, P., Palumbi, S., Richmond,

R. H., Rinkevich, B., Steneck, R. S., Voolstra, …

Gattuso, J. P. (2021a). Designing a blueprint for coral

reef survival. Biological Conservation, 257. https://doi.

org/10.1016/j.biocon.2021.109107

Kleypas, J. A., Villalobos-Cubero, T., Marin-Moraga, J. A.,

Cortés, J., & Alvarado, J. J. (2021b). Reef restoration

in the eastern tropical Pacific, a case study in Golfo

Dulce, Costa Rica. In D. Vaughan (Ed.), Active Coral

Reef Restoration: Techniques for a Changing Planet

(pp. 417–430). J. Ross Publishing.

La Nación. (3 de abril de 2000). Crean arrecife artifi-

cial. https://www.nacion.com/el-pais/crean-arrecife-

artificial/BXGV6PYB6VBRDA3LLBOMIXSZQY/

story/

La Nación. (30 de julio de 2004). Astronauta partici-

pa en hundimiento de barcos para crear arrecife.

https://www.nacion.com/archivo/astronauta-partici-

pa-en-hundimiento-de-barcos-para-crear-arrecife/

NQF5KFZUJVETNEHGQZ2XXXU4XM/story/

La Nación. (27 de noviembre de 2005). Estudiantes de

Paquera crean arrecifes artificiales.

https://www.nacion.com/ciencia/estudiantes-de-paquera-

crean-arrecifes-artificiales/7U543Z5EF5BEVL42XW

GJGYBMA4/story/

Lamont, T. A. C., Razak, T. B., Djohani, R., Janetski, N.,

Rapi, S., Mars, F., & Smith, D. J. (2022). Multi-dimen-

sional approaches to scaling up coral reef restoration.

Marine Policy, 143, 105199. https://doi.org/10.1016/j.

marpol.2022.105199

Larghi, P. (2022). Proyecto: Desarrollo de un mecanismo

financiero innovador para la conservación de arrecifes

de coral en República Dominicana SINAC: República

Dominicana – Costa Rica – Alemania: informe de

evaluación. Bolivia: Centro de Estudios y Proyectos

S.R.L.

Linkimer, L., Arroyo, I., Mora, M., Vargas, A., Soto, G.,

Barquero, R., Rojas, W., Waldo, T., & Taylor, M.

(2013). El terremoto de Sámara (Costa Rica) del 5

de setiembre del 2012 (Mw 7,6). Revista Geológica de

América Central, 49, 73–82. https://doi.org/10.15517/

rgac.v0i49.13104

Lirman, D., & Schopmeyer, S. (2016). Ecological solutions

to reef degradation: optimizing coral reef restoration

in the Caribbean and Western Atlantic. PeerJ, 4, e2597

https://doi.org/10.7717/peerj.2597

Lirman, D., Thyberg, T., Herlan, J., Hill, C., Young-Lahiff,

C., Schopmeyer, S., Huntington, B., Santos, R., &

Drury, C. (2010). Propagation of the threatened stag-

horn coral Acropora cervicornis: methods to minimize

the impacts of fragment collection and maximize

production. Coral Reefs, 29(3), 729–735. https://doi.

org/10.1007/s00338-010-0621-6

Liñán-Cabello, M., Flore-Ramírez, L., Laurel-Sadoval,

M., García, E., Soriano, O., & Delgadillo-Nuño, M.

(2010). Acclimation in Pocillopora spp. during a coral

restoration program in Carrizales Bay, Colima, Méxi-

co. Marine and Freshwater Behavior and Physiology,

44, 61–72. https://doi.org/10.1080/10236244.2010.5

37440

Lizano, O. G. (2007). Climatología del viento y oleaje fren-

te a las costas de Costa Rica. Ciencia y Tecnología,

25(1–2), 43–56.

Martínez, A. (8 de enero de 2021). Colocan primeros

arrecifes artificiales en Playa Blanca. Delfino. https://

delfino.cr/2021/01/colocan-primeros-arrecifes-artifi-

ciales-en-playa-blanca

Martínez-Castillo, V., Rodríguez-Troncoso, A. P., Tortolero-

Langarica, J. J. A., & Cupul-Magaña, A. (2023). Active

restoration efforts in the Central Mexican Pacific as

a strategy for coral reef recovery. Revista de Biología

Tropical, 71(S1), e54795. https://doi.org/10.15517/rev.

biol.trop..v71iS1.54795

McLeod, E., Anthony, K. R., Mumby, P. J., Maynard, J.,

Beeden, R., Graham, N. A., Heron, S. F., Hoegh-

Guldberg, O., Jupiter, S., MacGowan, P., Mangubhai,

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

S., Marshall, N., Marshall, P. A., McClanahan, T. R.,

Mcleod, K., Nyström, M., Obura, D., Parker, B., Pos-

singham, H. P., … Tamelander, J. (2019). The future of

resilience-based management in coral reef ecosystems.

Journal of Environmental Management, 233, 291–301.

https://doi.org/10.1016/j.jenvman.2018.11.034

McLeod, I. M., Hein, M. Y., Babcock, R., Bay, L., Bourne, D.

G., Cook, N., Doropoulos, C., Gibbs, M., Harrison, P.,

Lockie, S., van Oppen, M. J. H., Mattocks, N., Page,

C.A., Randall, C. J., Smith, A., Smith, H. A., Suggett,

D. J., Taylor, B., Vella, K. J., … Boström-Einarsson, L.

(2022). Coral restoration and adaptation in Australia:

The first five years. PLoS ONE, 17(11), e0273325.

https://doi.org/10.1371/journal.pone.0273325

Morales-Ramírez, A., Víquez, R., Rodríguez, K., & Vargas,

M. (2001). Marea roja producida por Lingulodi-

nium polyedrum (Peridiniales, Dinophyceae) en Bahía

Culebra, Golfo de Papagayo, Costa Rica. Revista de

Biología Tropical, 49(2), 19–23.

Mumby, P. J., Mason, R. A., & Hock, K. (2021). Reconnec-

ting reef recovery in a world of coral bleaching. Lim-

nology and Oceanography: Methods, 19(10), 702–713.

https://doi.org/10.1002/lom3.10455

Navarro-Cerdas, S. (2013). “Costa Rica” como periferia del

placer. Poder, colonialidad y resistencia en torno al

turismo y la inmigración en Playa Matapalo [Tesis de

Licenciatura]. Universidad de Costa Rica, Costa Rica.

Ortiz-Malavasi, E. (2004). Efectividad del programa de

pago de servicios ambientales por protección del

bosque (PSA-Protección) como instrumento para

mejorar la calidad de vida de los propietarios de bos-

que en zonas rurales. Revista Forestal Mesoamericana

Kurú, 1(2), 11–21.

Palou-Zuñiga, N., Madrigal-Ballestero, R., Schlüter, J., &

Alvarado, J. J. (2023). Applying the SES framework to

coral restoration projects on the Pacific coast of Costa

Rica. Revista de Biología Tropical, 71(S1), e54853.

https://doi.org/10.15517/rev.biol.trop..v71iS1.54853

Papke, E., Wallace, B., Hamlyn, S., & Nowicki, R. (2021).

Differential effects of substrate type and genet on

growth of microfragments of Acropora palmata.

Frontiers in Marine Science, 8, 623963. https://doi.

org/10.3389/fmars.2021.623963

Pereira-Pérez, A. I., & Mairena-Rodríguez, N. (2011). La

educación y la conservación de la naturaleza: Una

alianza impostergable. Revista Electrónica Educare,

15, 221–230.

Pizzimenti, C. A., McCarthy, C. T., Powell, J. M., & Ander-

son, N. T. (2011). Mala Noche River estuary: assessing

and raising community awareness (Bachelor’s Thesis).

Worcester Polytechnic Institute. https://digital.wpi.

edu/concern/student_works/nv935346t?locale=es

Quezada-Perez, F., Mena, S., Fernández-García, C., & Alva-

rado, J. J. (2023). Status of coral reef communities on

the Caribbean coast of Costa Rica: Are we talking

about corals or macroalgae reefs? Oceans, 4(3), 315–

332. https://doi.org/10.3390/oceans4030022

Richardson, B. J., & Lefroy, T. (2016). Restoration dialo-

gues: improving the governance of ecological restora-

tion. Restoration Ecology, 24(5), 668–673. https://doi.

org/10.1111/rec.12391 

Rinkevich, B. (1995). Restoration strategies for coral

reefs damaged by recreational activities: the use of

sexual and asexual recruits. Restoration Ecology, 3(4),

241–251. https://doi.org/10.1111/j.1526-100X.1995.

tb00091.x

Rinkevich, B. (2005). Conservation of coral reefs through

active restoration measures: recent approaches and

last decade progress. Environmental Science and Tech-

nology 39(12), 4333–4342. https://doi.org/10.1021/

es0482583

Robles-Payan, A., Reyes-Bonilla, H., & Cáceres-Martínez,

C. (2021). Crecimiento y supervivencia de cora-

les durante la fase inicial de cultivo en La Paz,

Baja California Sur, México. Revista Mexicana de

Biodiversidad, 92, e923594 https://doi.org/10.22201/

ib.20078706e.2021.92.3594

Rodriguez, R., Báez-Taveras, D., Valcárcel-Abud, A., Evan-

gelista-Pérez, D., Bello De Lillo, Y., & Matos Merce-

des, J. (2022). Importancia de los viveros de coral y su

impacto económico. Revista de Humanidades y Cien-

cias Sociales, 68(1), 78–91. https://doi.org/1033413/

aulahcs.2022.68i1.199

Rodriguez-Martinez, R. E., Banaszak, A. T., McField, M.

D., Beltran-Torres, A. U., & Alvarez-Filip, L. (2014).

Assessment of Acropora palmata in the Mesoameri-

can Reef System. PLoS One, 9, 4, e96140. https://doi.

org/10.1371/journal.pone.0096140

Sánchez, F., & Azofeifa, J. (2018). Guía para la planificación

e instalación de arrecifes artificiales en el área marina

de pesca responsable Paquera-Tambor, Puntarenas,

Costa Rica. [Technical report]. Fundación Corco-

vado. https://fundecooperacion.org/wp-content/

uploads/2020/10/Paquera-Guia-instalacion-Arrecife-

sArtificiales_AMPR-Paquera-Tambor.pdf

Sánchez-Noguera, C. (2012). Entre historias y culebras:

más que una bahía (Bahía Culebra, Guanacaste,

Costa Rica). Revista de Biología Tropical, 60(2), 1–17.

https://doi.org/10.15517/rbt.v60i2.19960

Sánchez-Noguera, C., Jiménez, C., & Cortés, J. (2018).

Desarrollo costero y ambientes marino-costeros

en Bahía Culebra, Guanacaste, Costa Rica. Revista

de Biología Tropical, 66(S1), 309–327. https://doi.

org/10.15517/rbt.v66i1.33301

Sandoval, M., Villalobos-Cubero, T., Marín-Moraga, J. A.,

& Kleypas, J. (2022). New insights for a successful cul-

tivation and transplantation of Porites lobata/Porites

evermanni, south Pacific coast of Costa Rica [Congress

28 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63695, enero-diciembre 2025 (Publicado Mar. 03, 2025)

speech, September 26-30, 2022]. Reef Futures 2022,

Key Largo, Florida, United States of America.

Shaver, E. C., McLeod, E., Hein, M. Y., Palumbi, S. R., Qui-

gley, K., Vardi, T., Mumby, P. J., Smith, D., Montoya-

Maya, P., Muller, E. M., Banaszak, A. T., McLeod, I.

M., & Wachenfeld, D. (2022). A roadmap to integra-

ting resilience into the practice of coral reef restora-

tion. Global Change Biology, 28, 4751–4764. https://

doi.org/10.1111/gcb.16212

Shafir, S., Van Rijn, J., & Baruch, R. (2006). Steps in

the construction of underwater coral nursery, an

essential component in reef restoration acts. Mari-

ne Biology, 149, 679–687. https://doi.org/10.1007/

s00227-005-0236-6

Sistema Nacional de Áreas de Conservación. (2013). Plan

de Manejo Parque Nacional Manuel Antonio [Reporte

técnico]. Sistema de Áreas de Conservación, Gobier-

no de Costa Rica. https://www.sinac.go.cr/ES/plan-

manejo/Paginas/pmacopac.aspx

Sistema Nacional de Áreas de Conservación. (2016). Pro-

tocolo PRONAMEC: Protocolo para el monitoreo eco-

lógico de formaciones coralinas [Reporte técnico].

Proyecto Consolidación de las Áreas Marinas Protegi-

das. Programa de Naciones Unidas para el Desarrollo

(PNUD) y El Fondo para el Medio Ambiente Mundial

(GEF), San José, Costa Rica.

SINAC-GIZ. (2020). Protocolo para la restauración de arre-

cifes y comunidades coralinas de Costa Rica. [Reporte

técnico]. Sistema Nacional de Áreas de Conservación

y Agencia de Cooperación Alemana, Costa Rica.

https://www.sinac.go.cr/ES/docu/ASP/Protocolo%20

Restauraci%C3%B3n%20Arrecifes%20y%20Cmuni-

dades%20Coralinas%20CR%202020.pdf

Smith, D., Mars, F., Williams, S., van Oöstrum, J., McArdle,

A., Rapi, S., Jompa, J., & Janetski, N. (2021). Case

study Indonesia: Mars assisted reef restoration sys-

tem (MARRS). In D. E. Vaughan (Ed.), Active coral

restoration: techniques for a changing planet (pp.

463–481). J. Ross Publishing.

Soto-Méndez, M. (enero de 2021). Arrecifes artificiales:

una opción para atraer biodiversidad y quitarle pre-

sión a natura. Ojo al clima. https://ojoalclima.com/

articles/arrecifes-artificiales-una-opcion-para-atraer-

biodiversidad-y-quitarle-presion-a-natura

Stuart-Smith, R. D., Brown, C. J., Ceccarelli, D. M., &

Edgar, G. J. (2018). Ecosystem restructuring along

the Great Barrier Reef following mass coral blea-

ching. Nature, 560, 92–96. https://doi.org/10.1038/

s41586-018-0359-9

Suding, K., Higgs, E., Palmer, M., Callicot, J. B., Anderson,

C. B., Baker, M., Gutrich, J. J., Hondula, K. L., LaFevor,

M. C., Larson, B. M. H., Randall, A., Ruhl, J. B., &

Schwartzfewer, K. Z. S. (2015). Committing to ecolo-

gical restoration: Efforts around the globe need legal

and policy clarification. Science, 348 (6235), 638-640.

https://doi.org/10.1126/science.aaa4216

Suggett, D. J., Camp, E. F., Edmondson, J., Boström-Einars-

son, L., Ramler, V., Lohr, K., & Patterson, J. T. (2019).

Optimizing return-on-effort for coral nursery and

outplanting practices to aid restoration of the Great

Barrier Reef. Restoration Ecology, 27, 683–693. https://

doi.org/10.1111/rec.12916

Suggett, D. J., Guest, J., Camp, E. F., Edwards, A., Goergen,

L., Hein, M., Humanes, A., Levy, J. S., Montoya-Maya,

P. H., Smith, D. J., Vardi, T., Winters, R. S., & Moore,

T. (2024). Restoration as a meaningful aid to ecologi-

cal recovery of coral reefs. Ocean Sustainability, 3, 20.

https://doi.org/10.1038/s44183-024-00056-8

Vardi, T., Hoot, W. C., Levy, J., Shaver, E., Winters, R. S.,

Banaszak, A. T., Baums, I. B., Chamberland, V. F.,

Cook, N., Gulko, D., Hein, M. Y., Kaufman, L., Loewe,

M., Lundgren, P., Lustic, C., MacGowan, P., Matz, M.

V., McGonigle, M., McLeod, I., … Montoya-Maya,

P. H. (2021). Six priorities to advance the science

and practice of coral reef restoration worldwide. Res-

toration Ecology, 29, e13498. https://doi.org/10.1111/

rec.13498

Vargas, R., Gómez, C., Pérez, C., Umaña, E., & Acosta,

M. (2020). “Jardinería” para la restauración coralina

en el Golfo Dulce, Costa Rica: Una prueba práctica.

UNED Research Journal, 12(1), e2809. https://doi.

org/10.22458/urj.v11i3v12i1.2809

Villalobos-Cubero, T. (2019). Manejo integrado y restaura-

ción ecológica de los arrecifes y comunidades coralinas

de Golfo Dulce, Pacífico Sur, Costa Rica [Tesis de

Maestría]. Universidad de Costa Rica, Costa Rica.

Villalobos-Cubero, T., Kleypas, J. A., Alvarado, J. J., &

Cortés-Núñez, J. (2023). Percepción comunitaria

sobre arrecifes coralinos en Golfo Dulce: bases para

integración social en programas de restauración.

Revista de Biología Tropical, 71(S1), e54862. https://

doi.org/10.15517/rev.biol.trop..v71iS1.54862

Thorne, R. E., Hedgepeth, J. B., & Campos, J. (1989).

Hydroacoustic observations of fish abundance and

behavior around an artificial reef in Costa Rica. Bulle-

tin of Marine Science, 44, 1058–1064.

Young, C. N., Schopmeyer, S. A., & Lirman, D. (2012). A

review of reef restoration and coral propagation using

the threatened genus Acropora in the Caribbean

and Western Atlantic. Bulletin of Marine Science, 88,

1075–1098. https://doi.org/10.5343/bms.2011.1143