Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

Fish spawning aggregations in the Gulf of Chiriqui,

Panamanian Pacific: six years of monitoring

Yolani A. Robles P.1; https://orcid.org/0000-0003-4140-9235

Ángel J. Vega1*; https://orcid.org/0000-0002-9535-3348

Luis A. Montes S.1; https://orcid.org/0000-0002-8823-1991

Erasmo E. Sánchez-Carles1; https://orcid.org/0009-0006-6106-7745

Roberto C. Lombardo G.1,2; https://orcid.org/0000-0002-0279-8621

1. Centro de Capacitación, Investigación y Monitoreo de la Biodiversidad en el Parque Nacional Coiba (CCIMBIO-

CRUV-UP), Universidad de Panamá, Santiago, Veraguas, Panamá; yolany.robles@up.ac.pa, angel.vega@up.ac.pa*, luis.

montes@up.ac.pa, esanchezc157@gmail.com, roberto.lombardo@up.ac.pa (*Correspondencia)

2. Sistema Nacional de Investigación (SNI), Secretaría Nacional de Ciencia, Tecnología e Innovación, Panamá

(SENACYT), Ciudad del Saber, Clayton Panamá, República de Panamá.

Received 03-VII-2025. Corrected 19-XI-2025. Accepted 30-I-2026.

ABSTRACT

Introduction: Fish spawning aggregations (FSAs) are temporary concentrations of individuals of the same spe-

cies that form for the sole purpose of reproducing.

Objective: To document the species, times, and localities where FSAs occur in the Gulf of Chiriqui, Panamanian

Pacific.

Methods: From 2020 to 2025, SCUBA surveys and photographic documentation were conducted to iden-

tify FSAs within Coiba National Park (CNP) and the Islas Secas Archipelago (ISA) in the Gulf of Chiriquí.

Environmental data, including temperature, salinity, and pH were collected using a YSI EXO2 multiparameter

probe, and temperature was continuously recorded with a HOBO Water Temperature Pro v2.

Results: The FSAs were recorded for three snapper species (Lutjanus peru, Lutjanus colorado and Lutjanus

aratus); one jack (Caranx sexfasciatus); one grouper (Cephalopholis colonus); and one wrasse (Thalassoma

lucasanum). Aggregations were observed at Bajo 20, Sacramento, Sueño del Pescador, and Montaña Rusa within

CNP, and at Bajo Rizo in ISA. In the latter location, aggregations were recorded for T. lucasanum, C. colonus,

and L. colorado. During aggregations, water column stratification was observed, associated with the intrusion

of cold-water masses into the gulf, thermocline shoaling, and a decrease in dissolved oxygen concentrations, all

correlated with temperature dynamics. Spawning events were most frequently observed in the morning hours.

Conclusion: The number of reported species forming FSAs in CNP increased from three to seven, and FSAs were

documented for the first time in ISA for three species.

Key words: spawning; Coiba; Islas Secas; reproduction; snappers.

RESUMEN

Agregaciones de desove de peces en el Golfo de Chiriquí, Pacífico panameño: seis años de monitoreo

Introducción: Las agregaciones reproductivas de peces (FSAs, por sus siglas en inglés) son concentraciones

temporales de individuos de la misma especie que se forman con el único propósito de reproducirse.

https://doi.org/10.15517/0wk2ee74

AQUATIC ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

INTRODUCTION

Fish spawning aggregations (FSAs) are

temporary concentrations of individuals of the

same species that form for the sole purpose of

reproducing. These events, which are key to

the survival of the species, are determined by

adaptation to habitat characteristics and ocean

dynamics (Erisman, Cota-Nieto et al., 2017;

Erisman et al., 2018). These aggregations lead to

large-scale, recurrent spawning events, and are

considered a reproductive strategy that increas-

es offspring survival probability (Domeier &

Colin, 1997; Domeier, 2012). Spawning aggre-

gations have been documented across several

commercially important fish families, including

snappers (Lutjanidae), seabass and groupers

(Serranidae) (Choat, 2012).

Various fish species that form spawning

aggregations are known to traverse great dis-

tances to specific sites, where they spawn sea-

sonally over short time windows (Boomhower

et al., 2010; Kobara et al., 2013; Kadison et al.,

2006), while other species are resident and trav-

el shorter distance (Domeir, 2012). For exam-

ple, the kelp bass, Paralabrax clathratus (Girard,

1854), maintain small home ranges (few hun-

dred m) where individuals repeatedly use reef

habitats for spawning aggregations (Lowe et al.,

2003). In contrast, the gulf weakfish, Cynoscion

othonopterus (Jordan & Gilbert, 1882), migrates

approximately up to ~ 150 km from widespread

coastal habitats to aggregate and spawn in the

Colorado River delta. (Erisman et al., 2012).

At the extreme end, the bluefin tuna, Thunnus

thynnus (Linnaeus, 1758), travels transoceanic

distances exceeding 4 000 km, from its foraging

areas in the North Atlantic to spawning aggre-

gation sites in the Gulf of Mexico and Mediter-

ranean Sea (Block et al., 2005).

Spawning aggregations are critical for the

sustainability of fisheries (Chollett et al., 2020;

Kobara et al., 2013); however, their predictabil-

ity also makes fish particularly vulnerable to

fishing pressure, as fishermen exploit such pre-

dictability to locate and capture large numbers

of individuals with relatively low effort (Pitt-

man & Heyman, 2020; Erisman, Heyman et al.,

2017; Sala et al., 2001; van Overzee & Rijnsdorp,

2015). In addition to fishing, environmental

changes driven by climate change also pose

serious threats for species that form transitory

spawning aggregations. This is the case as the

timing, success, and viability of aggregations

are highly dependent on ecological conditions.

Globally, climate change affects both breeding

and non-breeding fish populations; however,

the on-going change of oceanographic fea-

tures at aggregation sites suggests that sea sur-

face temperature and thermic gradients might

Objetivo: Documentar las especies, momentos y localidades donde ocurren las FSAs en el Golfo de Chiriquí,

Pacífico panameño.

Métodos: Entre 2020 y 2025, mediante buceo autónomo y con el uso de cámaras fotográficas se documentaron las

FSAs en el Parque Nacional Coiba (PNC) y en el archipiélago de Islas Secas (AIS), Golfo de Chiriquí. Las observa-

ciones fueron complementadas con registros de temperatura, salinidad y pH con el uso de una sonda multipara-

métrica YSI EXO2 y registro continuo de temperatura con un sensor térmico HOBO Water Temperature Pro v2.

Resultados: Se documentaron las FSAs en tres especies de pargos (Lutjanus peru, Lutjanus colorado, Lutjanus ara-

tus), un carángido, Caranx sexfasciatus; un serránido, Cefalopholis colonus; y un lábrido, Thalassoma lucasanum,

en las localidades: Bajo 20, Sacramento, Sueño del Pescador y Montaña Rusa, en el PNC y bajo Rizo, en AIS, en

este último caso para T. lucasanum, C. colonus y L. colorado. Durante el periodo de agregaciones se presentó una

estratificación en la columna de agua, por la presencia de masas de agua fría que entran al golfo, acercamiento de

la termoclina a la superficie y una baja en la concentración de oxígeno disuelto, correlacionado con el comporta-

miento de la temperatura. La mayor frecuencia de desoves se observó en horas de la mañana.

Conclusiones: Se aumentó de tres a siete los reportes de especies que realizan FSAs en el PNC y se documentaron

por primera vez en el AIS, para tres especies.

Palabras clave: desoves; Coiba; Islas Secas; reproducción; pargos.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

significantly influence fish phenology, distribu-

tion and oceanic habitat suitability. In this sce-

nario, species with narrow thermal tolerances

are particularly at risk (Asch & Erisman, 2018;

Sánchez-Hernández et al., 2022). For instance,

groupers (Epinephelidae) may be more affected

than snappers (Lutjanidae), as the latter gener-

ally exhibit broader thermal spawning thresh-

olds, while groupers often rely on cooler waters

for reproduction (Gokturk et al., 2022).

By 2014, 25 % of documented spawning

aggregations were reported to be in decline,

while 4 % had been considered lost (Russell

et al., 2014). Of the documented aggregations,

52 % had not been evaluated, fewer than 35 %

were under any form of protection, and only

around 25 % had some degree of monitor-

ing (Erisman et al., 2018). These proportions

stand in stark contrast to the importance of

spawning aggregations for the ocean’s ecology

and fisheries, as well-monitored and managed

aggregations have been shown to enhance fish-

ery performance, reflected in increased yields

at nearby sites where fishing is permitted (Eris-

man, Cota-Nieto et al., 2017). Among the man-

agement strategies for spawning aggregations,

the establishment of seasonal fishing closures

have proven effective when they: a) reduce

fishing mortality of the largest individuals,

which are crucial to the reproductive potential

of the population, b) protect spawning habitats,

c) lower the risk of overexploitation in species

that form large aggregations, d) mitigate the

evolutionary effects on size at maturation and

reproductive investment, and e) reduce the

risk of overexploitation of specific spawning

components (van Overzee & Rijnsdorp, 2015).

This effectiveness translates into increased fish

biomass, higher catch rate, and improved larval

recruitment at fishing grounds (Erisman, Cota-

Nieto et al., 2017).

Management requires science-based mea-

sures that can be provided to administrators

and stakeholders, through scientific research

and monitoring (Erisman, Cota-Nieto et al.,

2017). In the Eastern Pacific, spawning aggre-

gations have been reported for Lutjanus argenti-

ventris, Lutjanus novemfasciatus, Mycteroperca

prionura (Rosenblatt & Zahuranec, 1967), Myc-

teroperca jordani (Jenkins & Evermann, 1889),

Mycteroperca rosacea (Streets, 1877), Paranthias

colonus (Valenciennes, 1846), Caranx sexfas-

ciatus (Quoy & Gaimard, 1825), Seriola lal-

andi (Cuvier &Valenciennes, 1833) (Sala et

al., 2003), and C. othonopterus (Erisman et al.,

2012) in the Gulf of California; Tylosurus pacifi-

cus (Steindachner, 1876) in Colombia (Correa-

Herrera et al., 2017), Dermatolepis dermatolepis

(Boulenger, 1895) in Costa Rica (Erisman et al.,

2009); and Lutjanus peru and Lutjanus guttatus

in Coiba National Park (CNP) Panama (Vega,

Maté et al., 2016). The limited number and fre-

quency of spawning aggregation reports in the

Eastern Tropical Pacific likely reflects the lack

of long-term monitoring programs, rather than

a true absence of spawning aggregations (Sala

et al., 2003). Therefore, it is in the public inter-

est to develop research and monitoring plans to

increase current knowledge on fish spawning

aggregations in the region. Following the first

report of fish spawning aggregations in Panama

(Vega, Maté et al., 2016), and in recognition of

their ecological importance and relevance to

sustainable fisheries, a research and monitoring

program was established in 2020. The objective

of the present study is to document the species,

timing, and locations where spawning aggrega-

tions occur, based on six years of monitoring

(2020 - 2025) within CNP and the Islas Secas

Archipelago (ISA), in the Panamanian Pacific.

MATERIALS AND METHODS

Study area: The CNP is a protected area

in the Gulf of Chiriqui (GCH), Eastern Tropi-

cal panamanian Pacific. It extends a surface

of 2 701.25 km2, of which 2 165.43 km2 cor-

responds to sea surface (Maté et al., 2015;

Cardiel et al., 1997). The ISA consists of five

main volcanic islands and a number of smaller

islets located in the GCH, approximately 20 km

off the Western coast of Panama (Angehr et al.,

2021). During the dry season, the GCH is influ-

enced by the shoaling of the thermocline to

depths of up to 30 m due to trade winds, lead-

ing to cooling of surface waters and increased

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nutrient availability (D’Croz & O’Dea, 2007;

D’Croz & O’Dea 2009). However, analyses of

the vertical structure of the thermocline and

halocline show greater stratification during the

dry season (January and February), with these

layers located at depths ranging from 15 m and

30-35 m (Olivera et al., 2023).

Dive sites: Vega, Maté et al. (2016) report-

ed the presence of snapper FSAs at Parque

Nacional Coiba (PNC) during full-moon peri-

ods in the dry season, between February and

April. Research on FSAs resumed in 2020, and

fieldwork to document the occurrence of aggre-

gations was carried out during each dry season

from January to April (2020-2025). Through-

out this period, research was interrupted due

to COVID-19 in March 2020, and in April

2022 the start of fieldwork was delayed due to

budget setbacks.

The sites studied were: Bajo 20 (7°40’14.76”

N & 81°40’55.32” W), a submarine rocky pin-

nacle with mats of azooxanthellate coral and

patches of accumulated white sand. The shal-

lowest part of this site is about 28 m deep

according to the tide amplitude (Vega et al.,

2019). Bajo 20 is one of the two sites in the

CNP and the only one in the Eastern Tropi-

cal Pacific where spawning aggregations have

been recorded for Pacific red snapper (L. peru)

and spotted rose snapper (L. guttatus) (Vega,

Maté et al., 2016). Sacramento (7°45’25.29” N

& 81°40’51.43” W) is seamount located West

of Bajo 20, with its shallowest point at a depth

of 32 m. The bottom is densely covered with

black corals, which host significant aggrega-

tions of reef fishes. Montaña Rusa (7°53’8.93”

N & 81°48’50.41” W) consists of a rocky pin-

nacle, oriented North to South, with a mini-

mum depth of 12 m at its summit. The surface

is covered with algal mats, crustose coralline

algae, and corals of the genus Pocillopora, along

with octocorals, sponges, and other inverte-

brates. Sueños del Pescador (7°53’20.62” N &

81°48’50.46” W) is a rocky pinnacle that rises

to about 8 m below the surface, also oriented

North to South, similar to Montaña Rusa. The

summit is covered with complex arrangements

of scleractinian corals, and the rock walls are

lined with a diverse sea fan (A. Vega, personal

observation, May 13, 2023). In 2025, Bajo Rizo

(7°55’1.43” N & 82°0’30.25” W) was included

in the fieldwork campaigns. This site is located

in the ISA, 17 km West of the Contreras

Islands, outside the boundaries of CNP. Bajo

Rizo lies at a depth of 26 m at its shallowest

point and is characterized by a black coral for-

est scattered with sandy channels and rocky

formations (L. Montes, personal observation,

March 27, 2025) (Fig. 1).

Water column conditions: To correlate

fish behavior with prevailing water column

conditions, a YSI (EXO2) multiparameter

probe was deployed upon arrival at each site.

Fig. 1. Sampling sites for fish breeding aggregations in

Coiba National Park, Panamanian Pacific. Coiba National

Park (CNP), Islas Secas Archipelago (ISA), Contreras

Islands (CON), Bajo Rizo (BR), Sueños del Pescador (SP),

Montaña Rusa (MR), Sacramento (SA), Bajo 20 (B20). The

black outline indicates CNP boundaries.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

This instrument allowed the measurement of

temperature (°C), salinity (ups), dissolved oxy-

gen (mg/l) and pH from the surface down to 30

at 5 m intervals. In addition, the probe aided in

thermocline depth estimation. Additionally, In

July 2023, a temperature data logger (HOBO

Pro v2) was installed at a depth of 28 m in

Bajo 20. Temperature and depth values cor-

responding to fish aggregation and spawning

events were obtained from both the logger and

the dive computer. Water column were ana-

lyzed using the Pearson correlation. Differences

between categorical variables, when expressed

as proportions, were tested using a goodness-

of-fit chi-squared test. Statistical significance

was set at α = 0.05 (Zar, 2010).

Aggregation monitoring: Each field trip

began two days prior to the new and full moons

and concluded two days after those lunar phas-

es. The presence of fish schools was confirmed

prior to diving using a Garmin Echomap Ultra

106sv echosounder. During each dive, a team of

three to four divers descended, equipped with

two Sony Cyber-Shot DSC-RX100M6 cameras,

housed inside Nauticam NA-RX100VI frames,

one of which was fitted with a wide-angle lens

(130°-67 mm). Additionally, two GoPro Hero

10 Black cameras were used. Horizontal visibil-

ity was estimated by the divers as the maximum

distance at which a diving partner remained

clearly visible. Dives took place in the morning

(6:30-11:00 h) and afternoon (14:00-18:00 h),

each lasting 30-40 min, with two dives com-

pleted in the morning and two in the afternoon.

Monitoring activities at each site aimed to cap-

ture as much as possible of the present species,

their behavior, and depth at which spawning

occurred. In the laboratory, collected photo-

graphic and video materials were reviewed

with Adobe Premier Pro (Adobe, 2019) to

study the spawning behavior of each species

in the observed aggregations (Heyman et al.,

2004; Sala et al., 2003). To determine whether

aggregations qualified as FSAs, we record-

ed the species forming the aggregation, the

occurrence of spawning, and several indicators,

including color changes in individuals, swollen

abdomens, courtship behavior, the presence of

spawn remnants, predator presence, and ther-

mal stratification in the water column. In addi-

tion to these indicators, a spawning aggregation

event in this study was defined as one occurring

during either a low-tide or high-tide cycle.

RESULTS

Water column conditions: Significant cor-

relation was found between temperature, salin-

ity, dissolved oxygen and pH (Pearson, p <

0.05). Dissolved oxygen and temperature had

the strongest correlation (R = 0.84), while cor-

relation between the other variables was lower

yet significant (Table 1).

The greatest variations in temperature and

dissolved oxygen between the surface and 30

m depth occurred between January and April.

During this period, the water column was

stratified into three main layers. In the first

layer, the water was clearer and warmer in the

upper 10-12 m, with roughly 25 m of visibil-

ity. This was followed by a denser and more

turbid second layer extending to about 20 m

depth, where visibility dropped to roughly 7 m.

The third layer, reached the bottom, and was

clearer again (~ 15 m visibility) but significantly

colder, at 16 °C.

These variations were driven by the intru-

sion of a cold-water mass into the GCH during

dry season, causing the thermocline to raise

closer to the surface, especially between Feb-

ruary and mid-April. This resulted in thermal

stratification, with surface waters exceeding

Tabl e 1

Pearson correlations (r) between the different para m

recorded in Bajo 20, Panamanian Pacific, during 2024.

DO (mg/l) S (ups) pH

S (ups) R -0.38*

n1 914

pH R 0.39*-0.10*

n1 755 1 755

T (°C) R 0.84*-0.28*0.32*

n1 914 1 914 1 755

Salinity (S), temperature (T), and dissolved oxygen (DO).

Asterix represents significance.

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

30 °C and relatively cooler waters below 20 m

depth (Fig. 2). This change in the water column

structure coincided with fluctuations in dis-

solved oxygen concentrations, which dropped

below 3 mg/l and were associated with the

thermocline between 20-30 m depth; this was

particularly evident in March, when thermal

minima were also recorded at that depth (Fig. 3).

Salinity showed minor variation at all

depths in March across all monitoring years,

coinciding with the presence of the cool water

mass that flows through the PNC, with its influ-

ence being most pronounced between 20-30 m

depth (Fig. 4). The pH held between 7.5 and

8.5, with the highest values observed at the sur-

face in March, compared to those at the 20-30

m depth interval, particularly in 2021 and 2024

(Fig. 5).

Continuous temperature records at Bajo

20, from January and April 2024, show daily

oscillations ranging from 16 to 30 °C at approx-

imately 28 m depth (Fig. 6). During this period,

the lowest temperatures were associated with

full and new moon phases, coinciding with a

turbid water mass rich in organic matter and

low dissolved oxygen.

Aggregation monitoring: A total of 187

dives were completed, representing approxi-

mately 122 diver-hours. Between 2020 to 2025,

spawning aggregations were observed during

full and/or new moon phases. The six species

confirmed to form spawning aggregations in

CNP were L. peru, Lutjanus colorado, Lutja-

nus aratus (Günther, 1864), C. sexfasciatus,

Cephalopholis colonus, and Thalassoma lucasa-

num (Gill, 1862) (Table 2). Aggregations were

documented at Bajo 20, Sacramento, Sueño del

Pescador and Montaña Rusa in CNP, and Bajo

Rizo in the ISA. In the latter, aggregations were

Fig. 2. Temperature records in Bajo 20, between surface and 30 m depth, Coiba National Park, Panamanian Pacific.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

recorded for T. lucasanum, C. colonus, and L.

colorado (Jordan & Gilbert, 1882).

Between 2020 and 2025, 61 aggregation

events were recorded, where 32 events cul-

minated in spawning, while 29 showed signs

of reproductive activity but did not end in

spawning. During the morning at low tide,

21 spawning events were recorded and 11 in

the afternoon at high tide. On a full moon,

17 events were recorded and 15 during a new

moon (Table 2). There was no difference in the

proportions among these categories (spawn vs.

no spawn: χ2 = 0.15, p = 0.70; spawning at low

vs. high tide: χ2 = 3.13, p = 0.08; full vs. new

moon: χ2 = 0.13, p = 0.72).

The six species spawned during both full

and new moons, and during both high and low

tides, except for T. lucasanum, which spawned

only during low tide. Within the snapper group,

L. peru spawned two days apart in 2025, on

the evening of 12 March at Sacramento and

the morning of 14 March at Bajo 20, two sites

approximately 10 km apart within PNC. L.

aratus spawned on consecutive days, 7 and 8

April 2025, with afternoon and morning events

recorded at Bajo 20. A similar pattern occurred

Fig. 3. Dissolved oxygen (mg/l) profiles, between the surface and 30 m depth in Bajo 20, Coiba National Park, Panamanian

Pacific.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

in C. sexfasciatus (Carangidae), which spawned

on 24 and 25 March 2025, with morning and

afternoon events documented at Bajo 20 and

Sacramento (Table 2).

Lutjanus peru: With the arrival of the

cold-water mass, L. peru concentrates on or

around underwater mounds at depths between

25-40 m, and as the cold-water mass moves to

shallower depths, individuals follow it to begin

the reproductive process. A total of two events

were recorded during the new moon (Janu-

ary 25, 2020; January 22, 2023) and six during

the full moon (March 09, 2023; February 23

and March 23, 2024; February 12, March 12

and 14, 2025) (Table 2). Spawning aggrega-

tions generally occurred in the early morning

hours, although two afternoon events were also

observed. As the cold-water mass ascended, the

school separated off the rock pinnacle where

smaller groups of 5-200 individuals formed.

These groups began courtship behavior char-

acterized by chasing, where some individuals

interacted with the lead fish. During the chase,

fish clustered and circled, while swimming

turned slow rising in the water column while

nipping and rubbing against the leader until

it spawned. When the leader spawned, nearby

individuals were stimulated to spawn as well,

leaving behind a cloud of eggs and sperm in the

water column (Fig. 7).

Depending on the size of the aggregation,

spawning groups ranged from as few as ten to

as many as 200 individuals; however, in large

spawning groups, not all individuals were able

to release their gametes. Those located at the

periphery of the spawning core regrouped with

other fish to carry out a subsequent episode

Fig. 4. Salinity (ups) records between surface and 30 m depth in Bajo 20, Coiba National Park, Panamanian Pacific.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

of gamete release. The end of spawning was

signaled when individuals swam downslope

to rejoin the main aggregation. These pro-

cesses occurred at the interface between cold

and warm water masses. Observations indicate

spawning individuals were over 50 cm in total

length, although smaller individuals were pres-

ent, they did not participate in the spawning

process. L. peru spawning aggregations were

documented at Bajo 20 and Sacramento in CNP.

Lutjanus colorado: This species was

observed forming schools consisting of hun-

dreds of individuals around or near underwater

pinnacles. In some cases, fish are aggregated

at the interface between warm and cold water,

between 10-20 m depth. Within the school,

spontaneous spawning, either by individual

fish or as a mass event, occurred without

segregation into spawning subgroups. Spawn-

ing was observed in both the morning and

afternoon, with two events during the new

moon (April 9, 2024; March 28, 2025) and one

during the full moon (March 31, 2024) (Table

2). Aggregations were recorded at Sueño del

Pescador (SP) and Montaña Rusa (MR) in

Coiba National Park, and at Bajo Rizo (BR) in

Islas Secas (ISA), outside the park boundaries

(Fig. 8).

Lutjanus aratus: Similar to what was

observed in L. colorado, L. aratus formed

schools of hundreds of individuals around

submarine mounds or in open water column

at depths between 10-20 m. Spawning pulses

were observed within schools, one during full

moon (March 26, 2024) and three in new moon

(March 12, and April 6 and 7, 2024) in both

Fig. 5. pH records from the surface to 30 m depth at Bajo 20, Coiba National Park, Panamanian Pacific.

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

Fig. 6. Daily thermal profile in Bajo 20 between January and April 2024 at Coiba National Park, Panamanian Pacific. Data

corresponds to records from a fixed sensor at 28 m depth. Black line is the daily average temperature.

Fig. 7. Fish spawning aggregations in Pacific Red Snapper Lutjanus peru in Bajo 20, Coiba National Park. A. Spawning group

in pursuit. B. Spawning group.

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Tabl e 2

Summary of species observed in spawning aggregations from 2020 to 2025 in Coiba National Park, Pacific Panama.

Species Date Lunar phase Site Time of day Tide

Lutjanus peru Jan 25, 2020 N B20 a.m. L

Jan 22, 2023 N B20 a.m. L

Mar 9, 2023 F SA a.m. L

Feb 23, 2024 F SA p.m. H

Mar 23, 2024 F B20 a.m. L

Feb 12, 2025 F B20 p.m. L

Mar 12, 2025 F SA p.m. H

Mar 14, 2025 F B20 a.m. L

Lutjanus colorado Mar 31, 2024 FSP a.m. L

Apr 9, 2024 NSP p.m. H

Mar 28, 2025 NBR p.m. H

Lutjanus aratus Mar 12, 2024 N B20 a.m. L

Mar 26, 2024 F B20 a.m. L

Apr 6, 7, 2024 N B20 p.m., a.m. H, L

Caranx sexfasciatus Jan 27, 2021 F B20 a.m. L

Mar 14, 2021 N B20 a.m. L

Jan 23, 2023 N B20 a.m. L

Feb 3, 6, 2023 FB20, SA a.m., p.m. L, H

Mar 24, 25, 2024 FB20, SA a.m., p.m. L, H

Feb 25, 2025 NBR a.m. L

Mar 13, 2021 NSP p.m. H

Cephalopholis colonus Mar 29, 2021 FSP p.m. H

Jan 21, 2023 N B20 a.m. L

Jan 15, 2025 F B20 a.m. L

Feb 25, 2025 NBR a.m. L

Thalassoma lucasanum Apr 26, 2021 F MR a.m. L

Jan 16, 2025 F B20 a.m. L

Apr 13, 2025 F MR pm H

Jan 31, 2025 N B20 a.m. L

Fig. 8. Red snapper (Lutjanus colorado) spawning aggregation in Bajo 20 and Sueño del Pescador, Coiba National Park. A.

Note difference between the warm-water layer (WWL, ~27 °C) and the cold (CWL, ~18 °C) water in Bajo 20. B. White arrows

point to released gametes from spawning fish at Sueño del Pescador.

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

morning and afternoon, all in Bajo 20 (Table

2, Fig. 9).

Caranx sexfasciatus: This species formed

schools of hundreds to thousands of individu-

als, especially in the warm, clearer water layer.

While swimming over or around submarine

mounds in the colder water layer, the fish, in

response to an unknown stimulus, ascended

vertically in mass toward the warmer layer.

Within the school, dark-colored individuals

were observed separating from the group and

chasing lighter-colored individuals in a court-

ship ritual, positioning themselves beneath

their partner. Spawning occurred individually,

leaving a trail of gametes after the pair. Five

spawning events were observed during full

moon (January 27, 2021; January 3 and 6, 2023;

March 24 and 25, 2024) and three in new moon

(March 14, 2021; January 23, 2023 and Febru-

ary 25, 2025) (Table 2, Fig. 10).

Cephalopholis colonus: This species was

observed forming schools over coral reefs and

Fig. 9. A. Aggregation of mullet snapper (Lutjanus aratus) at Bajo 20. B. Spawning activity at Sueño del Pescador, indicated

by white arrows.

Fig. 10. Bigeye crevalle-jack (Caranx sexfasciatus) aggregation in Bajo 20, Coiba National Park. A. Specimens moving

vertically from the bottom to the water column, moving from cold to warm water. B. Aggregation. C. Pair with dimorphism

in courtship. D. Spawning pair in Bajo 20. Note the linear trail (white arrows) of gametes.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

rocky mound pinnacles. The spawning aggre-

gation segregated into multiple smaller groups

of about 12 individuals that swam vertically

to spawn. It was also observed forming large

school in the open water column, from which

smaller groups detached to carry out simulta-

neous spawning pulses. Two spawning events

occurred during the full moon (March 29, 2021

and January 15, 2025) and tree during the new

moon at Sueño del Pescador (March 13, 2021),

Bajo 20 (January 21, 2023), and Bajo Rizo, the

latter located outside the CNP boundaries (Feb-

ruary 25, 2025) (Table 2, Fig. 11).

Thalassoma lucasanum: Spawning aggre-

gations of this small fish formed groups of

10-60 individuals, over coral or rocky reef habi-

tats. Individuals ascended into the water col-

umn where females released eggs, followed by

males releasing sperm. Thee spawning events

were observed during full moon in Montaña

Rusa (April 26, 2021), Bajo 20 (January 16,

2025) and Montaña Rusa (April 13, 2025) and

one during new moon at Bajo 20 (31 January,

2025) (Table 2, Fig. 12).

Spawning aggregation indicators: During

spawning aggregations distinct water column

stratification was observed between surface lay-

ers down to 30 m depth. The top layer was warm

and clear followed by a cooler, turbid layer with

copious sediment and organic matter. The bot-

tom layer was cold and visibility improved. Tur-

bidity was associated to suspended particles of

organic origin, as well as plankton, cnidarians

and sediment (Fig. 8A). Another noteworthy

aspect was the presence of predators in the

water column such as Seriola rivoliana (Cuvier

& Valenciennes, 1833), Seriola peruana (Stein-

dachner, 1881), Caranx melampygus (Cuvier &

Valenciennes, 1833), Decapterus sp. (Bleeker,

1851), Elagatis bipinnulata (Quoy & Gaimard,

1825), Triaenodon obesus (Rüppell, 1837), Car-

charhinus falciformis (Müller & Henle, 1839),

Fig. 11. Pacific creole fish (Cephalopholis colonus) aggregation at Bajo Rizo, Islas Secas Archipelago, Panamanian Pacific. A.

Vertical movement of congregated fish. B. Spawning group.

Fig. 12. Aggregation of Cortez rainbow wrasse (Thalassoma lucasanum) at Bajo 20, Coiba National Park, Panamanian Pacific.

A. Vertical movement of spawning group. B. Spawning event with patches of released gametes (white arrows).

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

Hypanus longus (Garman, 1880), Aetobatus

laticeps (Gill, 1865), and Carcharhinus limbatus

(Valenciennes, 1839). Whale sharks, Rhincodon

typus (Smith, 1828), common CNP visitors,

were occasionally observed during the dry sea-

son, feeding at the spawning sites.

DISCUSSION

Since 2012, when spawning aggregations

of L. peru and L. guttatus were first reported

(Vega, Maté et al., 2016), no further monitor-

ing had been conducted. In 2020, monitoring

activities resumed and continued through 2025.

This effort has made it possible to document

the spawning aggregations of six additional

species in the region, expanding the known

group. Additionally, four new aggregation sites

within CNP and one in the ISA were identified,

all located within GCH.

Spawning aggregations have been linked to

oceanographic conditions, and the importance

of thermal stratification in the water column

has been highlighted by many studies, in fact

deepening of the thermocline, reduced verti-

cal mixing, and rising temperatures, disrupt

the spawning habitat suitability for Pacific cod,

Gadus macrocephalus (Tilesius, 1810), with

severe consequences for the fishery (Laurel

& Rogers, 2020). In contrast, during the dry

season, a cold-water mass enters the GCH sys-

tem, causing the thermocline to rise (D’Croz &

O’Dea, 2009). This process appears to promote

fish aggregation for spawning on or near sub-

marine mound pinnacles during the new and/

or full moon.

According to Russell et al. (2014), snapper

species exhibit the highest spawning frequency

during full moon and at sunset. While this

all aligns with our findings regarding lunar

phase, it differs in the time of day, as most

of the recorded spawning events occurred in

the morning. Nevertheless, our results suggest

thermal stratification of the water column is

critical given that fish species that form spawn-

ing aggregations exhibit behavioral adaptations

to spawn in habitats with temperatures that

optimize egg-larval development, highlighting

the importance of thermal regimes in reproduc-

tive success (Laurel & Rogers, 2020; Pankhurst

& Munday, 2011).

L. peru is the primary species in catches

from the GCH, accounting for 60 % of total

landings and approximately 90 % of the snap-

per catch. It is fished at depths between 20-100

m, with the highest frequency around 50 m.

Gonad analysis showed the species reproduc-

es year-round with peaks in May, July and

November (Vega, Robles-P et al., 2016). Spawn-

ing aggregations for L. peru were originally

described between February and April (Vega,

Maté et al., 2016), a period confirmed by our

monitoring from 2020 to 2025, which also

includes January as part of the spawning aggre-

gation season. Reproduction in L. peru through

spawning aggregations may be associated with

the thermal profile of the GCH, implying that

oceanographic conditions, particularly tem-

perature, influence the reproductive aggrega-

tion process. According to Kobara et al. (2013),

changes in water temperature, current velocity,

tidal cycles, and ambient light operate at inter-

mediate spatial scales (regional to local). These

same criteria can be applied to explain the

reproductive process of L. peru congeners: L.

colorado and L. aratus, species that form spawn-

ing aggregations at the same sites and periods

as L. peru, although their aggregation behavior

and density differ.

Furthermore, C. sexfasciatus was fre-

quently observed at high density, hundreds to

thousands of individuals, in the first m of the

water column where the water was warmer.

This species displayed sexual dimorphism and

distinct courtship behavior, which has been

reported from other localities, for example in

the Gulf of California (Sala et al., 2003) and

Thailand, associated to an oil and gas platform

(Madgett et al., 2022). Both report male color

change and fast spawning, a scenario in line

with the spawning aggregations observed at

CNP. Interestingly, this same behavior pat-

tern was also observed in congeners from

the Caribbean and other species within the

Carangidae (Graham & Castellanos, 2005). For

C. colonus, reproduction was described by Sala

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

et al. (2003) in the Gulf of California, where

spawning-related behaviors such as group for-

mation, color changes, and spawning events

were reported. In Cabo Pulmo, Rowell et al.

(2019) reported groups of 12 to 34 individu-

als spawning in multiple bouts. These same

behaviors were observed in CNP and the ISA,

where groups of approximately 12 individuals

gathered to spawn at depths between 12-25 m,

ascending through the stratified water column

after separating from the main school. The pat-

tern of ascension from the cold to the warm

water layer was also observed in T. lucasanum,

which is a species associated with coral com-

munities, and rocky mounds (Robertson et al.,

2024). Its spawning aggregations were reported

at Bajo 20 in CNP (Vega et al., 2019), consistent

with observations of Thalassoma bifasciatum

(Bloch, 1791) in the U.S. Virgin Islands. In both

cases, females were observed rapidly swimming

upward for 1-2 min to release eggs, followed

by accompanying males that released sperm

(Warner, 1995).

The ascent of fish into warmer layers may

facilitate gamete release through thermal con-

trast, a mechanism known to regulate reproduc-

tion. In the tropics, environmental regulation of

aggregating species appears to operate across

a hierarchy of variables with temperature and

photoperiod as primary drivers (Pankhurst &

Porter, 2003). The observed temperature strati-

fication may explain the upward swimming

behavior of aggregating fish as they enter the

warm layer to spawn. Importantly, the forma-

tion of spawning aggregations appears to be

facilitated by the seasonal intrusion of cold-

water masses, which raise the thermocline and

bring the fish closer to the warm surface layer.

This interface likely provides optimal condi-

tions for gamete fusion, egg development, and

overall offspring survival (Domeier & Colin,

1997; Domeier, 2012). During new and full

moon periods, a cold-water mass rises from the

bottom toward the coast, bringing the thermo-

cline closer to the surface. This causes spawning

fish groups to move into shallower areas, where

they enter the warm surface layer to spawn,

likely induced by the associated thermal and

pressure changes. The spawned products are

rapidly dispersed by surface currents, which

increase in intensity as the tidal stage changes.

According to Sánchez-Hernández et al. (2022),

shifts in oceanographic conditions at spawning

aggregation sites can be used by various spe-

cies to release gametes, promote fertilization,

and facilitate subsequent egg dispersal, thereby

reducing predation.

Fish spawning aggregations present a chal-

lenge for both fishers and managers, as fishing

during these reproductive events might be eco-

nomically profitable in the short term, but often

leads to overfishing and population collapse

(Tobin et al., 2013). For example, the Nassau

grouper, Epinephelus striatus (Bloch, 1792), has

undergone severe reductions in number and

density of its spawning aggregation across its

distribution range, and it is now considered a

threatened species (De Mitcheson et al., 2008).

In this context, among the species in this study,

snappers (especially L. peru), represent the

primary target of the fisheries in CNP (Vega,

Robles-P et al., 2016). These species are pro-

tected under the park’s fishing regulations,

which include a seasonal closure during the

spawning aggregation period and an absolute

ban on fishing within one nautical mile of any

island, islet, or emergent area inside the park

(Maté et al., 2015). This implies the spawning

aggregations are protected by mechanisms of

control and surveillance; nonetheless, landings

continued between January and April 2024, a

year in which both registered and unregistered

fishing vessels were observed operating at night

within the protected area (A. Vega, personal

observation, March 12, 2024).

In 2025, the panamanian ministry of envi-

ronment increased its personnel and started

radar surveillance within the protected area.

These management measures might signifi-

cantly improve control over illegal fishing in

the park. This is particularly important given

that fish spawning aggregations can result from

long-distance migratory patterns, concentrat-

ing spawning events at specific sites which are

repeated in time and space (transient aggre-

gations), or they can be resident, involving

16 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026203, enero-diciembre 2026 (Publicado Feb. 12, 2026)

short-range migratory patterns (Domeier &

Colin, 1997; Domeier, 2012). For the spawning

aggregations described in the present study,

further research is needed, particularly for

snappers due to their commercial value, to

determine whether these spawning aggrega-

tions are transient or resident, and whether

reproduction occurs exclusively within CNP or

also in other areas of the GCH or the Panama-

nian Pacific. Such research could help identify

future candidate sites requiring special protec-

tion and management.

Heidmann et al. (2024) recommended

increasing the protection of Lutjanus analis

(Cuvier, 1828) spawning aggregation sites in St.

Croix, U.S. Virgin Islands, by modifying exist-

ing regulations, improving enforcement, and

involving fishers in co-management actions.

In this context, the current regulations in CNP

for snapper species include both spatial and

temporal protection during the spawning sea-

son; however, much work remains regarding

co-management. Therefore, it is critical to

improve the efficacy of these regulations, par-

ticularly during nighttime hours, when illegal

fishing targeting spawning aggregations occurs.

Moreover, engaging communities within the

park’s area of influence is essential to raise

awareness about the importance of conserv-

ing these reproductive events to ensure the

reproductive potential of the species and the

long-term sustainability of snapper fisheries

and other associated species.

The next steps involve acoustic studies

to track individuals from species that form

FSAs, in order to determine their origins and

movements between aggregation sites and the

localities where they are caught by artisanal

fisheries in the Gulf of Chiriquí, as documented

by onboard observer programs (Vega, Robles-P

et al., 2016). Acoustic tracking would refine our

understanding of aggregation dynamics and

occurrence patterns, particularly whether the

same individuals spawn at nearby sites within

a one- to two-day window or whether distinct

groups contribute to the observed spatio-tem-

poral patterns.

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

We thank Secretaría Nacional de Ciencia,

Tecnología e Innovación de Panamá (SENA-

CYT), for funding to the projects EIE17-014 y

PFID21-165, which contributed to the develop-

ment of our monitoring program in 2020, 2022

and 2023. Our gratitude also goes to Interna-

tional Conservancy, Islas Secas Foundation

for their financial support of the aggregation

monitoring efforts in 2021, 2024, and 2025 and

to the University of Panama and Geoversity

Foundation for their administrative support.

We are grateful to our vessel captain, Franco

Camarena and to Dive Base Coiba for logisti-

cal support, and to the marine biology thesis

students at CCIMBIO for their support dur-

ing fieldwork. We also thank the anonymous

reviewers, who provided valuable comments

that helped improve the manuscript.

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