Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

Water-based tourism dynamics: spatial and temporal characterization

in three coral reefs from Bahía Culebra, North Pacific of Costa Rica

Gabriela Mercedes López Romero1*; https://orcid.org/0009-0001-4763-9536

Juan José Alvarado2,3,4; https://orcid.org/0000-0002-2620-9115

1. Posgrado en Gestión Integrada de Áreas Costeras Tropicales, Universidad de Costa Rica, San Pedro, San José 11501-

2060, Costa Rica; gamelopez1995@gmail.com (*Correspondence)

2. Escuela de Biología, Universidad de Costa Rica, San Pedro, San José 11501-2060, Costa Rica; juan.alvarado@ucr.ac.cr

3. Centro de Investigación en Biodiversidad y Ecología Tropical (CIBET), Escuela de Biología, Universidad de Costa Rica,

San Pedro, San José 11501-2060, Costa Rica.

4. Centro de Investigación en Ciencias del Mar y Limnología (CIMAR), Universidad de Costa Rica, San Pedro, San José

11501-2060, Costa Rica.

Received 30-X-2025. Corrected 21-IV-2026. Accepted 23-IV-2026.

ABSTRACT

Introduction: Coral reefs are essential ecosystems for tourism-based economies worldwide, offering ecological,

social and economic benefits. However, increasing on-water activities represent a risk to reef health. In Culebra

Bay, there is limited understanding of how on-water tourism interacts with the coral reefs of the area.

Objective: To characterize the spatial and temporal distribution of on-water tourism interactions around coral

reefs sites in Culebra Bay.

Methods: Direct observations were conducted twice a month from April 2021 to April 2022, at three coastal

coral reefs to document the number of interactions and users, the types of activities and watercraft involved, and

the duration of each interaction. A Kruskal-Wallis test was applied to evaluate differences across sites, and the U

Mann-Withney test for compare tourist seasons (peak and off-season). Additionally, a Correspondence Analysis

(CA) was used to explore the relationship between activities and sites.

Results: We recorded 2,437 interactions involving 6 728 users, and identified ten tourism-related activities. The

spatial distribution of tourism interactions was heterogeneous among sites, with the highest volume of interac-

tions, activities, and users concentrated at Jícaro reef. The activities involving watercraft-based activities were

dominant, representing 93% of all interactions. Also, the recorded activities were different by site: Blanca reef was

mainly linked to non-motorized tours, while Virador reef was primarily used for transit. We found no statistical

differences in the volume of users, the temporal distribution of interactions and its duration between seasons.

Conclusion: Aquatic tourism in Bahía Culebra was concentrated at Jícaro reef and dominated by motorized

activities. This pattern may pose potential environmental risks for reef resilience.

Key words: coral reefs; tourism; marine-based activities; coastal management.

RESUMEN

Dinámica del turismo acuático: caracterización espacial y temporal

en tres arrecifes de coral de Bahía Culebra, Pacífico Norte de Costa Rica

Introducción: Los arrecifes de coral son ecosistemas esenciales para las economías basadas en el turismo en

todo el mundo, ya que ofrecen beneficios ecológicos, sociales y económicos. Sin embargo, el aumento de las

https://doi.org/10.15517/szqjfj14

SUPPLEMENT

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

INTRODUCTION

Coral reefs are the most diverse marine

ecosystems in the world (Gaston, 2013), where

important biological processes and complex

ecological relationships occur (Karlson, 1999).

For such attributes, these ecosystems are able

to provide local communities with a variety

of ecosystem services, such as food provision,

coastal protection, recreational opportunities,

and, in many coastal zones, economic oppor-

tunities through the use of reefs as a tourism

resource (Pabel & Prideaux, 2018). It is estimat-

ed that the tourism industry on a global scale

contributes approximately 10% to gross domes-

tic product (GDP) and generates direct and

indirect employment (World Tourism Organi-

zation [UNWTO], 2017). Coral reef tourism

alone generates an estimated USD 35.8 billion

annually worldwide (Spalding et al., 2017).

However, all that glitters is not gold. Coral

reefs are currently facing profound transforma-

tions caused by the tourism industry and its

associated coastal development (Dong, 2025).

While tourism may sustain the economy of sev-

eral coastal areas, it also brings significant chal-

lenges to vulnerable ecosystems such as coral

reefs (Chakraborty, 2021; Wolf et al., 2019).

Major significant concerns include environ-

mental degradation and pollution, ecosystem

destruction, and the loss of marine and coastal

resources (Hall, 2001). Such effects may depend

on tourism and coastal development depend on

a country’s ecological conditions, local gover-

nance structures, and economic priorities.

Culebra Bay, located in the Gulf of

Papagayo on the North Pacific coast of Costa

Rica, is an area historically characterized by

the development of the most extensive coral

reefs (up to 2 km long) in that region of the

country (Cortés, 1996-1997; Cortés & Jiménez,

2003), primarily dominated by the branching

coral Pocillopora and the presence of abundant

associated marine fauna (Cortés, 2012; Cortés

& Jiménez, 2003). This area is influenced by

the intensification of the Easterly trade winds

between December and April, generating a

seasonal upwelling phenomenon characterized

by an increase in nutrients and a decrease in

seawater temperature (Cortés, 1997). Due to

these natural characteristics, its scenic beauty,

and cultural richness, the Gulf of Papagayo was

deemed suitable in the 1970s for the creation

of a mass tourist destination and to promote

actividades realizadas sobre el agua representa un riesgo para la salud de los arrecifes. En Bahía Culebra existe un

conocimiento limitado sobre cómo el turismo acuático interactúa con los arrecifes de coral del área.

Objetivo: Caracterizar la distribución espacial y temporal de las interacciones turísticas realizadas sobre el agua

en los arrecifes de coral de Bahía Culebra.

Métodos: Se realizaron observaciones directas dos veces al mes entre abril de 2021 y abril de 2022 en tres arre-

cifes costeros para documentar el número de interacciones y usuarios, el tipo de actividad y de embarcaciones

involucradas, y duración de las interacciones. Para evaluar las diferencias entre sitios se empleó la prueba de

Kruskal-Wallis, mientras que la prueba U de Mann-Whitney se utilizó para comparar las temporadas turísticas

(pico y fuera de pico). Además, se realizó un análisis de correspondencia (CA) para examinar las asociaciones

entre las actividades y los sitios arrecifales.

Resultados: Se registraron 2 437 interacciones que involucraron a 6 728 usuarios, y se identificaron 10 actividades

directamente relacionadas con el turismo. La dinámica espacial del turismo fue heterogénea entre sitios, con el

mayor volumen de interacciones, actividades y usuarios concentrado en el arrecife Jícaro. Las actividades que

involucraron embarcaciones fueron dominantes, representando el 93 % de todas las interacciones. Asimismo, las

actividades variaron según el sitio: el arrecife Blanca se asoció principalmente con tours no motorizados, mientras

que el arrecife Virador se utilizó principalmente para tránsito. No se encontraron diferencias estadísticas en la

dinámica turística entre temporadas.

Conclusión: El turismo acuático en Bahía Culebra se concentra en el arrecife Jícaro y está dominado por activi-

dades motorizadas. Este patrón podría representar riesgos ambientales potenciales.

Palabras clave: arrecifes de coral; turismo; actividades marinas; gestión costera.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

economic development in the area. It was not

until 1991 that the large-scale tourism project

“Polo Turístico Golfo de Papagayo” (PTGP) was

implemented (Cordero-Ulate, 2010; Sánchez-

Noguera, 2012), and since its implementation,

the area has experienced unprecedented chang-

es in the rapid development of hotel infra-

structure, luxury residences, and an increase in

the supply of recreational activities (Sánchez-

Noguera, 2012).

Unfortunately, the synergy between natural

and anthropogenic disturbances, at both global

and local scales, including seasonal upwell-

ing, intense El Niño events, Harmful Algal

Blooms (HABs), eutrophication, and coastal

development, has led to a decline in marine

water quality, structural changes in the ben-

thos community through the development of

macroalgae such as the invasive alga Caulerpa

sertularioides, and an increase in the population

of the bioeroding sea urchin Diadema mexica-

num. These factors have resulted in local coral

bleaching and coral mortality (Alvarado et al.,

2018; Fernández & Cortés, 2005; Jiménez, 2007;

Morales-Ramírez et al., 2001).

Consequently, the average live coral cover

decreased from 44% in some sites in the 1990s

to 1-4% in 2011 (Alvarado et al., 2018; Sán-

chez-Noguera et al., 2018). Despite this, Cul-

ebra Bay remains one of Costa Rica’s most

important tourist destinations, even after the

complete shutdown due to COVID-19 (Costa

Rican Tourism Institute [ICT], 2023). Given the

urgent need for sustainability in tourism activi-

ties and the conservation of marine resources,

the first coral reef restoration project in the

country’s North Pacific was implemented in

2019 (Fabregat-Malé et al., 2024) through a

public-private partnership. However, restora-

tion processes demand an understanding of

current tourist pressure, since the success of a

restoration project depends not only on techni-

cal aspects, but also on the intensity of tourist

use and good tourism practices.

While the effects of tourism development

have been documented in various coral reefs

around the world, and the ecological aspects

of reefs in Culebra Bay have been extensively

studied (Cortés, 2012), much of this research

has focused primarily on the effects of coastal

urbanization expansion, pollution, and waste

management (Sánchez-Noguera et al., 2018).

Meanwhile, the impact of aquatic tourism

activities, including recreational diving, snor-

keling, boating, etc., has been less studied local-

ly. However, in other reefs worldwide, it has

been determined that such activities can cause

harmful disturbances to ecosystems, including

direct damage from anchoring, sediment resus-

pension, changes in marine fauna behavior,

and introduction of nutrients into the water.

These disturbances may affect the health of

coral reefs and consequently affect the human

communities that depend on this resource for

their livelihoods.

Unfortunately for Culebra Bay, which lacks

any status of protection, the precise details

of aquatic activities occurring there, number

and types of watercraft and visitors, and how

the intensity of water tourism varies between

peak and off seasons are unknown. This gap of

information limits the assessment of potential

negative impacts of on-water tourism on the

bay’s reefs. Therefore, the research question of

this study was: What are the spatial and tempo-

ral characteristics of on-water tourism on three

coral reefs undergoing ecosystem restoration in

the North Pacific of Costa Rica?

MATERIALS AND METHODS

Study area: This study was conducted in

Bahía Culebra (10°36’54.38” N, 85°39’32.86”

W), located in Northwestern Costa Rica, a

semi-enclosed bay covering approximately

24 km², and reaching depths up to 42 m (Cor-

tés, 2012). This bay is an important biodiversity

hotspot and one of the major coastal tourist

destinations in Costa Rica (Sánchez-Noguera,

2012). Its reefs are dominated by branching cor-

als (Pocillopora spp.) and characterized by the

presence of massive species such as Pavona cla-

vus and Porites lobata (Jiménez, 2001; Sánchez-

Noguera et al., 2018). The seasonal upwelling

phenomenon occurs between December and

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

April, coinciding with the peak tourism season

in the country (Alfaro et al., 2015).

The bay lies within the PTGP developing

area; therefore it has promoted the establish-

ment of hotel complexes and luxury residential

infrastructure and a marina along its coastline

(Sánchez-Noguera, 2012). Its reefs support a

variety of marine-based recreational activities,

including snorkeling, SCUBA diving, boating,

wildlife watching, and sport fishing.

The three selected coral reefs (Fig. 1) are in

front of different beaches inside the bay. Both

Blanca and Virador reefs lie adjacent to a luxury

hotel, however these two sites can be visited by

the general public, as they are freely accessible

by land. Jícaro reef is a relatively isolated site

that can be only accessed by boat. During this

study, the three sites were part of the coral reef

restoration project.

Data collection: Direct observations were

conducted twice per month from April 2021 to

April 2022 on three coral reefs within the bay.

Each visit consisted of at least 4 h of observation

of the aquatic activities on the reef ’s surface

area between 08:00 and 16:00. Observations

were made using binoculars from fixed shore-

based points. The coral reefs were delimited by

buoys indicating the presence of underwater

restoration structures, ensuring that counts

were conducted within the desired area. Either

the environmental conditions and distance did

not limit visibility or affect the accuracy of

observations as none of the sites were located

more than 400 m away from the shoreline. In

order to standardize the collecting data process,

two previous visits were made to train aspects

such as identification, classification of activi-

ties and watercraft types. Moreover, a single

observer conducted all observations.

For each interaction observed, the follow-

ing attributes were recorded: type of activity

(tourism fishing, transport, patrolling, scien-

tific), aquatic vehicle involved (e.g., catamaran,

boat, panga, jet ski, kayak), number of users,

time of interaction, and duration of the interac-

tion. For this study, the duration of interaction

was the total time spent by an interacting ele-

ment (e.g. a swimmer, a diver, a vessel, etc.)

inside of the delimited coral reef area, regardless

Fig. 1. Location of the study sites (red dots) in Bahía Culebra, northwestern Costa Rica.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

the activity involved was simple transit or a lon-

ger use. For motorized vessels, the duration was

recorded regardless of whether the engine was

on or off.

Data analysis: Comparative analyses were

conducted on the following attributes: num-

ber of interactions, types of activities, number

of users, watercraft, and interaction duration.

Data normality was tested using the Shapiro-

Wilk test. As the data did not meet normal-

ity assumptions, comparisons among sites and

among months were performed using the Krus-

kal–Wallis test. When comparing the attributes

by tourism seasons, Mann-Whitney U tests

were applied. When statistically significant dif-

ferences were found, Dunn’s post hoc test with

Bonferroni correction was applied to iden-

tify pairwise differences while controlling for

Type I error.

In order to evaluate whether the frequency

distribution of activity and watercraft types

differed among sites, chi-square tests (χ²) were

performed. Additionally, Correspondence

Analysis (CA) was conducted to visualize the

relationship between activities and types of

watercraft observed at each site.

Scheirer–Ray–Hare test was used to assess

differences in interaction frequency due to

activity type and season for each reef separately.

To ensure representativeness, activities

recorded only once were excluded from inter-

action-duration analyses. All statistical analyses

were performed in R (R Core Team, 2024).

RESULTS

Activity patterns: A total of 2 437 interac-

tions were recorded distributed across twelve

on-water activities in the study area. Two of

which were not directly related to tourism or

recreation: artisanal fishing and scientific div-

ing. Regarding the volume of interactions, sig-

nificant differences were detected among sites

(H = 26.48, p < 0.05), with Virador reef showing

significantly fewer interactions per day, while

Jícaro and Blanca reefs presented similar values

(Table 1, Fig. 2A).

Table 1

Tourism attributes of three coastal coral reefs at Bahía Culebra. * Only recorded once.

Jícaro Blanca Virador

Mean interactions-day-¹52.68 ± 17.40 36.18 ± 17.95 21.91 ± 11.39

Mean users-day-¹184.13 ± 76.91 70.68 ± 31.98 51.00 ± 24.22

Mean watercraft

interactions-day-¹

46.77 ± 16.01 35.32 ± 17.58 21.68 ± 11.39

Mean interaction

duration (min)

20.86 ± 34.88 3.12 ± 20.62 4.08 ± 25.38

Tourism use

On-reef activities

Snorkeling

Motorized tours

Non-motorized tours

Free diving*

Recreational diving*

Surface supplied air diving*

Motorized tour

Non-motorized tour

Snorkeling

Motorized tour

Snorkeling

Reef- adjacent activities Leisure boating

Transit

Boat cleaning*

Private patrolling

Leisure boating

Transit

Leisure boating

Transit

Private patrolling*

Non tourism use Artisanal fishing*

Scientific diving

Scientific diving Fishing

Scientific diving

Watercrafts Jet ski, Boat, Paddleboard, Kayak

Catamaran, Panga boat

Banana boat, Towable tube boat

Jet ski, Boat, Paddleboard

Kayak, Panga boat

Banana boat

Jet ski, Boat

Panga boat, Catamaran

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

All tourism-related activities were

observed at Jícaro reef, however, five of them

were recorded only once during the monitoring

period (Table 1). At Blanca and Virador reefs,

only five tourism-related activities were record-

ed. The activities that occurred on all reefs

were motorized tours, leisure boating, transit

and snorkeling. Of which, the motorized tours

exhibited the highest value of interactions-

day-¹ (Table 2), while snorkeling interactions

showed the lowest value of interactions-day-¹.

The rest of activities differentially occurred on

the studied reefs.

The Scheirer–Ray–Hare test (SRH test)

revealed significant differences in the rela-

tive frequency of activities at the Jícaro (H =

9.899, p = 0.042) and Virador (H = 35.379, p <

0.001) reefs, but not at Blanca (H = 7.325, p =

0.120). This indicates that activities at Blanca

exhibited more homogeneous frequencies. Post

hoc Dunn tests showed that at Jícaro, motor-

ized and non-motorized tours had significantly

different frequencies (p = 0.048). At Virador

reef, the motorized tours exhibited significantly

higher interaction values compared with lei-

sure boating (p = 0.045) and transit (p < 0.001)

Fig. 2. A. Boxplot of daily interaction rates (interactions·day-¹) across study sites. The x-axis indicates the study sites, and the

y-axis represents the number of interactions per day. Central horizontal lines within the boxes represent the median. Black

points represent outliers. B. Correspondence Analysis (CA) between on-water activities and the sites. Blue points represent

activities. Red items correspond to sites.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

(Table 2). Which indicates that interaction vol-

ume varied according to activity type.

Additionally, the distribution of activities

at the three reefs was not random and showed

a significant association between activities and

sites (χ² = 426.34, p < 0.05). The Correspon-

dence Analysis (Fig. 2B) revealed that the wide-

spread activities such as motorized tours and

transit were not specifically related to any of the

reefs, as they occurred across all sites, reflect-

ing their common practice in the inner area of

the bay. The rest of activities were differentially

associated to each site (Fig. 2B).

Concerning temporal aspects, despite

higher mean daily interactions per day were

recorded in the high season, no significant dif-

ferences were found between tourism seasons

(U = 614, p = 0.23) across all sites. Neither

at each site (Blanca: U = 78, p = 0.25; Jícaro:

U = 70, p = 0.99; Virador: U = 79, p = 0.25)

(Table 3). The RSH test also revealed that no

significant differences were found in the daily

number of interactions by activity type between

high and low season at Jícaro (H = 2.274, p =

0.132), Blanca (H = 0.046, p = 0.830), or Vira-

dor (H = 0.058, p = 0.809), despite mean daily

interaction rates were higher during the high

season (Table 4). In addition, the distribution of

activities by season was homogeneous across all

three reefs (Jícaro: H = 0.809, p = 0.937; Blanca:

H = 4.541, p = 0.338; Virador: H = 0.535, p =

0.970). Which means differences in interaction

frequency were more strongly related to the

type of activity than to the season.

On the other hand, significant monthly

variations were detected in the number of

interactions between Jícaro and Virador reefs (p

= 0.0002), where Virador reef exhibited lower

daily interaction rates (Fig. 3). The highest

number of interactions was recorded at Jícaro

reef in March 2022 (79.0 ± 15.6 interactions-

day-¹), while both Virador and Blanca reefs

showed peaks in July 2021 (46.5 ± 4.9 and 61.0

± 7.1 interactions-day-¹, respectively).

User patterns: A total of 6 728 users

were registered across the three monitored

sites. Significant differences were detected in

Table 2

Mean (+ SD) of daily tourism indicators by activity at three coastal coral reefs in 2021-2022 in Bahía Culebra, North Pacific of Costa Rica. * groups-day-¹.

Activity

Jícaro Blanca Virador

Mean

interactions-day-¹

Mean

users-day-¹

Mean time of

interaction (min)

Mean

interactions-day-¹

Mean

users-day-¹

Mean time of

interaction (min)

Mean

interactions-day-¹

Mean

users-day-¹

Mean time of

interaction (min)

Snorkeling * 5.09 ± 1.98* 34.71 ± 19.04 36.81 ± 24.81 0.63 ± 0.84* 3.40 ± 2.99 10.29 ± 7.56 0.23 ± 0.00* 1.80 ± 1.30 14.60 ± 14.25

Motorized tours 24.07 ± 11.57 44.95 ± 15.41 6.71 ± 19.45 19.86 ± 12.29 30.52 ± 16.40 0.56 ± 3.68 13.50 ± 6.30 20.19 ± 11.05 0.42 ± 1.48

Non-motorized tours 1.63 ± 3.81 5.00 ± 3.74 8.17 ± 12.89 6.95 ± 6.13 9.90 ± 9.13 4.08 ± 19.59   

Leisure boating 10.72 ± 4.27 75.50 ± 36.62 52.88 ± 46.70 2.77 ± 2.51 23.00 ±16.52 11.54 ± 50.50 2.00 ± 3.49 17.21 ± 12.93 28.63 ± 75.73

Transit 6.73 ± 9.44 31.59 ± 40.12 9.77 ± 23.29 5.72 ± 5.08 17.11 ± 12.02 3.07 ± 24.39 5.82 ± 6.14 19.04 ± 10.81 0.44 ± 1.63

Private patrolling 0.63 ± 1.01 3.71 ± 1.70 51.28 ± 46.90 

Fishing      0.31 ± 0.46 3.86 ± 0.89 64.29 ± 22.89

Scientific diving 0.59 ± 0.74 5.88 ± 3.36 56.46 ± 39.99 0.22 ± 0.57 7.66 ± 2.81 75.80 ± 53.02   

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

Table 3

Mean (± SD) of daily tourism indicators at coastal reefs during high and low seasons in Bahía Culebra, North Pacific of

Costa Rica.

Site

Mean interactions-day-¹

(nhigh = 10)

(nlow= 12)

Mean users-day-¹

(nhigh = 10)

(nlow= 12)

Mean aquatic vehicles

interactions-day-¹

(nhigh = 10) (nlow= 12)

Mean interaction

duration (min)

Jícaro high = 52.6 ± 19.7

low =52.8 ± 16.1

high = 178.1 ± 67.2

low = 189.2 ±86.8

high = 48.1 ± 18.6

low = 46.7 ± 14.8

high = 21.9 ± 35.8 (n = 526)

low = 20.0 ± 34.1 (n = 633)

Blanca high = 40.3 ± 16.0

low = 32.8 ± 19.5

high = 85.8 ± 34.2

low = 58.1 ± 24.7

high = 38.6 ± 15.9

low = 36.7 ± 19.5

high = 4.8 ± 28.2 (n = 403)

low = 1.4 ± 6.4 (n = 393)

Virador high = 23.8 ± 9.6

low = 20.3 ± 12.9

high = 59.2 ± 27.2

low = 44.5 ± 20.0

high = 23.5 ± 9.4

low = 20.2 ± 13.0

high = 4.3 ± 32.4 (n = 238)

low = 3.9 ± 15.9 (n = 244)

Table 4

Seasonal mean (± SD) of interactions-day-¹ of shared activities in three reefs from Bahía Culebra, Costa Rica.

Activity Jícaro Blanca Virador

Leisure boating high = 12.0 ± 4.9

low = 9.7 ± 3.5

high = 5.8 ± 1.8

low = 1.9 ± 1.4

high = 4.6 ± 2.6

low = 1.7 ± 0.5

Motorized tour high = 32.7 ± 12.8

low = 22.3 ± 8.3

high = 23.7 ± 12.3

low = 18.2 ± 12.3

high = 16.1 ± 6.3

low = 7.6 ± 7.6

Non-motorized tour high = 2.5 ± 1.0

low = 5.2 ± 4.9

high = 6.6 ± 3.9

low = 8.5 ± 7.6

Transit high = 2.3 ± 1.5

low = 12.2 ± 10.2

high = 6.3 ± 4.3

low = 8.2 ± 5.7

high = 4.1 ± 3.3

low = 6.2 ± 7.4

Snorkeling high = 4.8 ± 1.5

low = 5.8 ± 2.3

Fig. 3. Monthly variation of interactions during the monitoring period. Asterisks (*) indicate significant differences (p <

0.05), including the p-value from Dun’s test. Error bars represent the standard deviation.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

the number of users (H = 38.93, p < 0.05)

among reefs (Fig. 4A). Jícaro reef exhibited

significantly higher number of users, (a total of

4 055 users, representing 60.3 % of the total).

Whereas Blanca and Virador reefs had a similar

number of users (p = 0.055). See Table 1 for the

daily average of users of each site.

Additionally, significant differences were

detected in the number of users associated

with each activity (H = 131.31, p < 0.05) across

sites. Only transit exhibited similar values of

users-day-¹ across all sites (Table 2). At Jícaro

reef, most of activities showed more associated

users than at the other reefs, excepting for non-

motorized tours had significantly more users at

Blanca reef (H = 3.76, p < 0.05).

Regarding the temporal aspects of user pat-

terns, the overall number of users did not differ

significantly (U = 646.5, p = 0.10) between the

high (111.0 ± 67.3 users-day-¹) and low seasons

Fig. 4. A. Box plot of number of users-day-¹ of each study site. The x-axis indicates the study sites, and the y-axis represents

the number of users per day. Central horizontal lines within the boxes represent the median. Black points represent outliers.

B.Monthly variation of interactions during the monitoring period. Asterisks (*) indicate significant differences (p < 0.05),

including the p-value from Dun’s test. Error bars represent the standard deviation.

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

(97.1 ± 84.1 users-day-¹). Similarly, when the

analysis was conducted separately for each reef,

no significant seasonal differences in the num-

ber of users·day-¹ were detected at Blanca (U

= 88, p = 0.06), Jícaro (U = 60.5, p = 0.99), or

Virador (U = 84.5, p = 0.11) (Table 3).

Regarding monthly variation, significant

differences in the number of monthly users

were found among sites (H = 21.5, p < 0.05),

with Jícaro reef showing the highest monthly

mean users (p = 0.004) (Fig. 4B), whereas no

significant differences were observed between

Blanca and Virador reefs (p = 0.56). However,

we did not find significant differences in the

number of users among the monitored months

(HBlanca = 12.83, HJícaro = 14.97, HVirador

= 17.53, p > 0.05). Despite the absence of sta-

tistically significant monthly differences within

sites, descriptive patterns indicated marked

temporal fluctuations, with a decrease in user

numbers across all sites in June 2021, followed

by a sharp increase in July.

Watercrafts: We recorded a total of 2 284

interactions involving watercrafts, represent-

ing 93 % of all recorded interactions. Eight

watercraft types were identified in the area

(Table 1), of which jet skis (64.1%) and boats

(23.4%) were the most abundant, followed by

paddleboards (5.2%), kayaks (3.0%), catama-

rans (1.5%), panga, which is an artisanal short

scale boat (1.5%), banana boats (1.4%), and

towable tubes (< 1%). Mean interactions day-¹

of jet skis and boats in each reef are shown

in Table 5. When analyzing the number of

watercraft interactions across sites, significant

differences were detected (H = 22.59, p < 0.05).

Jícaro and Blanca reefs showed similar values

(p = 0.07) while Virador reef recorded signifi-

cantly fewer watercraft interactions (Refer to

Table 1 for mean watercraft interactions day-¹

of each reef).

Fig. 5. Relationship between watercrafts and sites. Red points represent sites. Blue points represent watercrafts. The proximity

between items indicates a stronger association between them.

Table 5

Mean (± SD) interactions day-¹of jet skis and boats on three

reefs from Culebra Bay.

Jícaro Blanca Virador

Jet ski 28.31 ± 12.29 22.86 ± 12.79 12.07 ± 10.92

Boat 13.18 ± 5.28 5.90 ± 2.51 5.25 ± 3.32

A Correspondence Analysis (CA) revealed

associations between specific watercrafts and

the monitored sites (χ² = 312.37, p < 0.05)

(Fig. 5). Non-motorized watercraft such as

paddleboards and kayaks were primarily

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

associated with Blanca reef, whereas catama-

rans and banana boats were more frequent at

Jícaro reef. Interactions involving pangas were

mainly related with Virador reef. In contrast,

jet skis and boats were the most frequent across

all sites, showing no strong site-specific pattern.

Regarding the temporal attributes of water-

crafts’ occurring, no significant differences

were found between high and low tourism

seasons (38.0 ± 17.3 vs. 33.2 ± 19.0 interactions-

day-¹; U = 612, p = 0.24), nor across individual

months (H = 0.99, p = 0.61).

Interaction duration: In general, duration

ranged from 0 to 480 min. Most interactions

lasted less than one minute (65.6 %, n = 1 599),

while 27.5 % (n = 670) lasted between 1 and

60 min. Longer interactions were less frequent:

5.7 % (n = 138) lasted between 61-120 min,

and only 1.2 % (n = 30) exceeded 120 min.

Significant differences were found among sites

(H = 294.3, p < 0.005). As shown in Table 1,

the mean interaction duration was similarly

short at both Blanca and Virador reefs, with no

significant difference between them (p = 0.71).

In contrast, interactions at Jícaro reef were

significantly longer. In addition, significant

differences were detected in interaction dura-

tion among activities. Transit, motorized tours,

leisure boating, and snorkeling lasted longer

at Jícaro compared to Blanca and Virador

(Table 2),while non-motorized tours and fish-

ing showed similar durations in the sites where

they occurred.

Concerning the interaction duration of

watercrafts, significant differences were detect-

ed among sites (H = 689.4, p < 0.05). With

the exception of paddleboards (p = 0.093) and

panga boats (p = 0.263) which exhibited simi-

lar interaction duration across sites (Fig. 6). In

contrast, jet skis (H = 75.1, p < 0.05), boats (H

= 165, p < 0.05), and kayaks (H = 16.0, p < 0.05)

showed significantly longer interaction times at

Jícaro reef.

Concerning temporal aspects, interaction

duration did not differ significantly between

seasons at any of the reefs (Blanca: U = 83825,

p = 0.49; Jícaro: U = 170960, p = 0.29; Virador:

U = 29888, p = 0.41) (Table 3). In contrast,

monthly differences in interaction duration

Fig. 6. Box plot representing the variability of interaction duration (min) across sites by the recorded watercrafts. The

x-axis indicates the study sites, and the y-axis represents interaction duration (min). Central horizontal lines within

the boxes represent the mean interaction duration. Black points represent outliers. Asterisks (*) indicate significant

differences (p < 0.05).

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

were found across all sites (HBlanca = 26.94;

HJícaro = 19.61; HVirador = 26.69; p < 0.05), with

the longest mean durations occurring in April

2022 at Jícaro reef (32.39 ± 46.22 min), in Feb-

ruary 2022 at Blanca reef (8.31 ± 33.27 min),

and in January 2022 at Virador reef (19.59 ±

92.13 min).

DISCUSSION

Spatial characterization: The differences

observed in the spatial distribution of tourism

intensity among sites could be influenced by a

combination of natural (Anfuso et al., 2017),

social (Dolnicar et al., 2017; Giorgio et al.,

2018), and economic factors (Spalding et al.,

2017; Xiao et al., 2020). These include biodiver-

sity richness, and ecosystem quality (McKenna

et al., 2011), tourist preferences (Blanco-Pimen-

tel et al., 2022), and the availability of tourism

facilities and accessibility (Carey et al., 1997;

Giorgio et al., 2018).

Jícaro reef received the highest propor-

tion of tourism interactions (47% of the total),

despite its lack direct infrastructure and

restricted access by sea only. This phenomenon

suggests that the site’s pristine and exclusive

environment may exert a stronger attraction

than infrastructure-based convenience. Fur-

thermore, Jícaro’s natural characteristics, such

as its limited exposure to waves, the shallow

reef close to the shore, and its marine biodi-

versity, likely enhance its suitability for leisure

boating and snorkeling (Wolf et al., 2019)

compared to the other sites. This pattern is

consistent with global tourism trends showing a

growing tourist preference for pristine environ-

ments (Dolnicar, 2020; Dolnicar et al., 2017).

Accordingly, Jícaro reef recorded the highest

mean number of users significantly exceeding

that of the other sites.

In contrast, the adjacent hotel infrastruc-

ture and better accessibility of Blanca and

Virador reefs attract a different tourist segment

(Dong, 2025), generating distinct patterns of

site use (Mihai et al., 2023). At Jícaro reef, tour-

ism intensity was characterized by group-based

recreational activities, influenced by multiple

external departure points to Jícaro reef (mainly

Playa del Coco and Playa Hermosa), rather

than by hotels within the immediate area (Cor-

dero-Ulate, 2010; Sánchez-Noguera, 2012).

Meanwhile, Blanca and Virador (Güiri-Güiri)

reefs are more associated with logistical and

operational uses linked to the nearby luxury

hotel operations and services. This difference

is reflected in the interaction duration: recre-

ational activities at Jícaro reef lasted significant-

ly longer than those at Blanca and Virador reefs,

where interactions were shorter and primarily

transit-related (Table 2). These findings suggest

that Jícaro reef functions as a receiver of exter-

nal tourism flows, contributing to its higher use

intensity compared to the other two reefs.

Underwater activities were concentrated at

Jícaro reef (Table 1), though their low frequency

suggests they were isolated events. Other reefs

in and near the bay (e.g., Monkey Head, Mur-

ciélago Island, Catalinas Islands) offer better

conditions for diving, such a as greater vis-

ibility, depth and biodiversity (Giglio et al.,

2015; Giorgio et al., 2018; Sánchez-Noguera,

2012). At Blanca reef, the prevalence of non-

motorized tours likely reflects the availability

of kayak and paddleboard rentals at the nearby

hotel, rather than a genuine preference for these

activities. This reinforces the idea that tourism

infrastructure and natural reef features joint-

ly shape activity distribution (Spalding et al.,

2017). The case of Virador reef is interesting,

as no direct reef-use touristic activities were

identified there, but it was found that transit

is the only tourism-related activity that occurs

systematically there.

Fishing activity showed spatial shifts. While

Jiménez (1997) and Jiménez (2001) reported 24

commercial dives off Playa Blanca (1995-1996),

and none at Virador reef (Jiménez, 2001), our

study found that fishing now occurs mainly at

Virador reef, predominantly through snorkel-

ing. We recorded 0.31 ± 0.46 interactions-day-¹,

with an average interaction time of 65.60 ±

21.48 min. This shift may be related to recent

improvements in ecosystem conditions fol-

lowing coral restoration efforts at Virador reef

(Güiri-Güiri) (Fabregat-Malé et al., 2023).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

Temporal characterization: During the

monitoring period, Costa Rica was transition-

ing from COVID-19 restrictions toward gradual

reopening of the tourism sector (Gómez-Melé-

ndez, 2022; Instituto Costarricense de Turismo

[ICT], 2023). In this context, we found that

the mean number of interactions-day-¹, users-

day-¹, interactions with aquatic vehicles-day-¹,

and interaction time (min) were all higher dur-

ing the high tourism season (December–April)

compared with the low season (May–Novem-

ber). However, no statistically significant differ-

ences were detected between seasons or among

individual sites. This absence of strong sea-

sonality suggests a gradual recovery of coastal

tourism one year after the “anthropause” (Rutz

et al., 2020) caused by the COVID-19 shutdown.

To contextualize these dynamics, interna-

tional tourist arrivals from the nearest airport

were used as a proxy for regional tourism flow

(Fig. 7). We observed a progressive increase in

international arrivals between April 2021 and

April 2022 (ICT, 2023), with marked peaks in

July 2021 and March 2022. Although this pat-

tern was not identical to the number of users

in Bahía Culebra, similar peaks were evident

in July 2021 and the first four months of 2022

(Fig. 7B). These temporal patterns may reflect

local or national events, (e.g., national holi-

days, school breaks, spring break season), that

Fig. 7. A. International arrivals to International Airport Daniel Oduber Quirós, North Costa Rica, from national tourism

authority (Instituto Costarricense de Turismo, ICT) data 2021-2022. B. Monthly variation of the number of users on the

three sites.

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74 (S1): e20267730, abril 2026 (Publicado May. 20, 2026)

temporally amplify visitor pressure. Compa-

rable effects have been documented in other

coral reef areas during special events (Widmer

& Underwood, 2004), which often increase

visitor numbers and alter behavior (e.g., alcohol

consumption, lack of awareness of local regula-

tions and the location of coral reefs and resto-

ration sites). Such scenarios require adaptive

management strategies to mitigate occasional

surges in human impact and promote sustain-

able visitor behavior.

Ethical statement: The authors declare

that they all agree with this publication and

made significant contributions; that there is no

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

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

ACKNOWLEDGMENTS

The present study was performed within

the framework of the coral reef restoration

project “Culebra Reef Gardens”, which would

not have been possible without the support of

Centro de Investigación en Ciencias del Mar

y Limnología (CIMAR), and to the founding

of Vicerrectoría de Investigación from Univer-

sidad de Costa Rica. We also thank Península

Papagayo and the German Agency for Inter-

national Cooperation (GIZ) for logistical and

financial support.

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