Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026184, enero-diciembre 2026 (Publicado Feb. 04, 2026)

Short-term variability of fish larvae assemblages in Malpelo island,

an isolated oceanic island in the Eastern Tropical Pacific

Alan Giraldo1*; https://orcid.org/0000-0001-9182-888X

Felipe Muriel-Hoyos1, 2; https://orcid.org/0000-0002-5392-3605

Diego F. Córdoba-Rojas1; https://orcid.org/0000-0002-7817-9632

Juan J. Gallego-Zerrato1; https://orcid.org/0000-0002-4633-1265

1. Grupo de Investigación en Ciencias Oceanográficas, Departamento de Biología, Facultad de Ciencias Naturales y

Exactas, Universidad del Valle, Cali, Colombia; alan.giraldo@correounivalle.edu.co (*Correspondence), felipe.muriel@

correounivalle.edu.co, diego.cordoba.rojas@correounivalle.edu.co, juan.j.gallego@correounivalle.edu.co

2. Parques Nacionales Naturales de Colombia, Dirección Territorial Pacífico, Santuario de Fauna y Flora Malpelo,

Buenaventura, Colombia.

Received 23-VII-2025. Corrected 08-IX-2025. Accepted 20-I-2026.

ABSTRACT

Introduction: Oceanic islands are key biogeographic features that influence the distribution and diversity of

marine organisms, particularly in tropical ecosystems. Malpelo Island, located 380 km off the Colombian Pacific

coast, is the only oceanic island in the country and a strategic site for assessing ichthyoplankton dynamics in the

Eastern Tropical Pacific (ETP).

Objective: To characterize the taxonomic composition and spatial ecological structure of fish larvae assemblages

associated with Malpelo Island in the Colombian Eastern Tropical Pacific under prevailing hydrographic condi-

tions. Specifically, to assess differences in species richness, abundance, and diel vertical distribution between

insular and adjacent offshore sites. These patterns are interpreted in the context of localized larval retention and

vertical migratory behaviors, potentially shaped by insular oceanographic features.

Methods: Short-term variability in larval fish assemblages was evaluated using vertical zooplankton trawls

conducted in September 2012 and July 2016 across three depth strata. Fish larvae were identified to the lowest

possible taxonomic level, and hydrographic profiles of temperature and salinity were obtained using CTD casts.

Results: A total of 26 taxonomic units representing eight orders and twelve families were recorded. Assemblages

near the island included epipelagic and reef-associated taxa such as Sardinops sagax and Anchoa spp., while

offshore stations were dominated by mesopelagic species including Diogenichthys laternatus and Vinciguerria

lucetia. Although larval abundance did not differ significantly between locations (MW; p = 0.38), assemblage

similarity was low (45 %) and species turnover high (79 %). Larvae exhibited diel vertical migration, occupying

deeper strata (50-100 m) during daylight and ascending to shallower layers (0-50 m) at night, influenced by local

thermocline and halocline depth.

Conclusions: These findings highlight the ecological distinctiveness of Malpelo Island and its role in support-

ing larval fish diversity in tropical oceanic environments. The results provide baseline data for future ecological

monitoring and conservation planning in insular marine habitats of the ETP.

Key words: ichthyoplankton; oceanic islands; assemblages; diversity; vertical migration.

https://doi.org/10.15517/5h3qkd21

AQUATIC ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026184, enero-diciembre 2026 (Publicado Feb. 04, 2026)

INTRODUCTION

Oceanic islands play a vital role in shaping

the spatial distribution of marine fauna. These

open-ocean enclaves often exhibit enhanced

primary productivity and zooplankton abun-

dance compared to surrounding waters, sup-

porting high fish species richness and biomass

(Boehlert & Mundy, 1993; Lima et al., 2016;

Macedo-Soares et al., 2012; Morato et al., 2010).

They also offer favorable environmental condi-

tions for the survival and development of early

life stages of both pelagic and resident fish

species (Dower & Perry, 2001). The composi-

tion of larval fish assemblages associated with

oceanic islands appears to be driven by a com-

plex interplay of adult reproductive strategies,

local physical and biological conditions, and

species-specific ecological and behavioral traits

(Boehlert & Mundy, 1993; Harris et al., 2020;

Moyano & Hernández-León, 2011; Suthers et

al., 2006). In these habitats, the study of vertical

and horizontal distribution patterns of ichthyo-

plankton is central to larval fish ecology, closely

linked to local hydrographic processes and diel

vertical migration behaviors (Höffle et al., 2013;

Leis, 2006; Munk, 2016; Olivar et al., 2018).

The Eastern Tropical Pacific (ETP) is typi-

fied by warm, low-salinity surface waters and

a shallow thermocline, shaped by the South

Equatorial Current and the intrusion of cooler,

saline subtropical waters to the NorthWest. Sea-

sonal and interannual dynamics, particularly

driven by the El Niño–Southern Oscillation,

influence temperature, currents, and biological

productivity. Despite seasonal coastal upwelling

RESUMEN

Variabilidad a corto plazo de los ensamblajes de ictioplancton en isla Malpelo,

una isla oceánica aislada en el Pacífico Oriental Tropical

Introducción: Las islas oceánicas son elementos biogeográficos clave que influyen en la distribución y diversidad

de organismos marinos, especialmente en ecosistemas tropicales. La isla de Malpelo, ubicada a 380 km de la costa

pacífica colombiana, es la única isla oceánica del país y constituye un sitio estratégico para evaluar la dinámica

del ictioplancton en el Pacífico Tropical Oriental (POT).

Objetivo: Caracterizar la composición taxonómica y la estructura ecológica espacial de los ensamblajes de larvas

de peces asociados a la Isla de Malpelo, en el Pacífico Oriental Tropical colombiano, bajo las condiciones hidrográ-

ficas predominantes. Específicamente, se evalúan las diferencias en la riqueza de especies, la abundancia y la dis-

tribución vertical diaria entre sitios insulares y oceánicos adyacentes. Estos patrones se interpretan en el contexto

de mecanismos locales de retención larval y comportamientos migratorios verticales, posiblemente modulados

por las características oceanográficas insulares.

Métodos: Se examinó la variabilidad temporal a corto plazo de los ensamblajes de larvas de peces mediante arras-

tres verticales de zooplancton realizados en septiembre de 2012 y julio de 2016, en tres estratos de profundidad.

Las larvas fueron identificadas al nivel taxonómico más bajo posible, y se obtuvieron perfiles hidrográficos de

temperatura y salinidad mediante lanzamientos de CTD.

Resultados: Se registraron 26 unidades taxonómicas pertenecientes a ocho órdenes y doce familias. Los ensam-

blajes cercanos a la isla incluyeron especies epipelágicas y asociadas a arrecifes como Sardinops sagax y Anchoa

spp., mientras que las estaciones alejadas estuvieron dominadas por especies mesopelágicas como Diogenichthys

laternatus y Vinciguerria lucetia. Aunque no se detectaron diferencias significativas en la abundancia larval entre

las estaciones (MW; p = 0.38), la similitud entre ensamblajes fue baja (45 %) y el recambio de especies alto (79 %).

Las larvas mostraron migración vertical diurna, ubicándose en estratos profundos (50-100 m) durante el día y

ascendiendo a capas superficiales (0-50 m) por la noche, influenciadas por la profundidad local de la termoclina

y haloclina.

Conclusiones: Estos resultados destacan la singularidad ecológica de la isla de Malpelo y su papel en el manteni-

miento de la diversidad larval de peces en ambientes oceánicos tropicales. El estudio aporta datos de referencia

esenciales para el monitoreo ecológico y la planificación de conservación en hábitats marinos insulares del POT.

Palabras clave: ictioplancton; islas oceánicas; ensamblajes; diversidad; migración vertical.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026184, enero-diciembre 2026 (Publicado Feb. 04, 2026)

that enhances productivity, iron limitation

often constrains phytoplankton growth. Below

the thermocline, oxygen minimum zones and

low pH levels contribute to challenging bio-

geochemical conditions (Fiedler & Talley, 2006;

Kessler, 2006; Pennington et al., 2006; Wang

& Fiedler, 2006). The Colombian sector of the

ETP comprises the easternmost portion of this

region and is part of the Panama Bight marine

ecoregion (Amador et al., 2016; Spalding et al.,

2007). For most of the year (April–December),

this area is influenced by persistent southWest-

erly winds, fostering surface thermal homoge-

neity and stratification. Reported hydrographic

values include a 21 ºC isotherm at 35 m, a 17

ºC isotherm at 125 m, and a mixed layer depth

of 22 m (Amador et al., 2016; Corredor-Acosta

et al., 2020; Rodríguez-Rubio & Giraldo, 2011).

Recent studies in other insular systems

have revealed that ichthyoplankton assemblag-

es are often structured by mesoscale oceano-

graphic features such as eddies, fronts, and

upwelling zones, which influence larval reten-

tion, transport, and survival (Aceves-Medina

et al., 2018; Zhang et al., 2021). These features

can create localized hotspots of larval abun-

dance and diversity, particularly around sub-

marine ridges and island slopes where vertical

mixing and nutrient enrichment occur. For

example, research conducted above the Ninety

East Ridge in the Eastern Indian Ocean dem-

onstrated that ichthyoplankton abundance was

positively correlated with sea surface salinity,

temperature, and chlorophyll-a concentration,

suggesting that insular productivity gradients

play a key role in shaping larval distributions

(Zhang et al., 2021). Such findings underscore

the importance of integrating hydrographic and

biological data to understand larval fish ecol-

ogy in oceanic island contexts.

Malpelo Island is the sole oceanic island in

this Colombian portion of the ETP. It represents

the emergent peak of the Malpelo Ridge, a sub-

marine mountain system rising from depths of

nearly 4 000 m, featuring a central rocky prom-

ontory reaching 300 m above sea level and asso-

ciated outcrops (Rodríguez-Rubio & Giraldo,

2011). Unlike other oceanic areas in the ETP,

biological and ecological data on ichthyoplank-

ton around Malpelo remain limited (Comisión

Colombiana del Océano, 2018). This is largely

due to research efforts traditionally focusing

on neritic and coastal zones (Beltrán-León &

Rios-Herrera, 2000; Calle-Bonilla et al., 2017;

Escarria et al., 2007; Martínez-Aguilar et al.,

2010; Medina-Contreras et al., 2014; Ramírez-

Mártinez et al., 2022; Valencia et al., 2024).

Nonetheless, recent surveys have reported lar-

vae of Synchiropus atrilabiatus (Beltrán-León

et al., 2016), and Diplophos proximus (Beltrán-

León & Ríos-Herrera, 2018), in nearshore Mal-

pelo waters.

In the Colombian Pacific, comparative

studies across neritic and oceanic areas—

including Malpelo island—have shown that

ichthyoplankton composition varies markedly

with distance from shore and habitat type (Bel-

trán-León & Morales-Osorio, 2021). Coastal

areas tend to be dominated by Engraulidae

and other neritic taxa, while oceanic zones

such as Malpelo exhibit higher proportions of

mesopelagic families like Phosichthyidae and

Bregmacerotidae. These patterns reflect both

the biogeographic isolation of oceanic islands

and the influence of regional oceanography

on larval dispersal and assemblage structure.

Understanding these dynamics is essential for

assessing the ecological connectivity of insular

systems and informing conservation strategies

that account for larval transport and recruit-

ment processes.

This study examines the taxonomic com-

position and spatial ecological patterns of fish

larvae assemblages associated with an iso-

lated oceanic island in the Colombian East-

ern Tropical Pacific (ETP), under prevailing

hydrographic conditions. We hypothesize that

ichthyoplankton assemblages near Malpelo

Island exhibit significantly greater species rich-

ness and abundance compared to adjacent off-

shore sites. Furthermore, we propose that both

areas display comparable diel vertical stratifi-

cation, with larvae occupying distinct depth

strata during daytime and nighttime periods.

These patterns are interpreted as evidence of

localized retention mechanisms and vertical

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026184, enero-diciembre 2026 (Publicado Feb. 04, 2026)

migratory behaviors shaped by insular oceano-

graphic features.

MATERIALS AND METHODS

Study area: Malpelo Island (4°00’08’’ N &

81°36’31’’ W) lies approximately 380 km off the

Colombian mainland (Fig. 1). Designated as a

Sanctuary of Fauna and Flora (SFF) in 1995, it

forms a critical node within the Eastern Tropi-

cal Pacific Marine Corridor (CMAR), a trans-

boundary conservation initiative (Enright et al.,

2021; Murillo, 2015). The pelagic environment

surrounding Malpelo is shaped by the interac-

tion of three distinct water masses, Superfi-

cial Tropical Water (ATS): Characterized by

temperatures > 25 °C and salinity < 34, this

water mass dominates the upper 100 m and is

typical of regions NorthNorth of the equator;

Equatorial Surface Water (ASE): With tempera-

tures < 25 °C and salinity > 34, ASE originates

from equatorial upwelling and influences sur-

face conditions along the equatorial belt, and

Subtropical Subsurface Water (ASSST): Found

between 100 and 115 m depth, this water mass

exhibits intermediate properties (19-25 °C,

34.6-35.4 salinity) and is likely derived from

Central Pacific Water intrusions (Rodríguez-

Rubio & Giraldo, 2011). A general description

of the surface circulation patterns in the study

area during March and September can be found

in Valencia et al. (2013).

Fig. 1. A. Geographic location of Malpelo Island within the Tropical Eastern Pacific, emphasizing its position in the

Colombian Pacific region. B. Bathymetric representation of the study area within the Malpelo Fauna and Flora Sanctuary

(SFF Malpelo), indicating the positions of biological sampling stations surveyed during the 2012 (black dot) and 2016 (red

star) expeditions.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026184, enero-diciembre 2026 (Publicado Feb. 04, 2026)

The primary climatological forcing in the

study area is driven by seasonal variability

in the equatorial trade winds system, which

modulates mesoscale circulation patterns and

influences regional productivity (Corredor-

Acosta et al., 2020; Devis-Morales et al., 2008;

Rodríguez-Rubio & Stuardo, 2002; Rodríguez-

Rubio et al., 2003). Average surface circula-

tion exhibits meridional characteristics, with

southeastward flow observed in March (mean

current velocity: 0.59 m/s) and Northward flow

in September (mean current velocity: 0.54 m/s)

(Rodríguez-Rubio & Giraldo, 2011).

Field methodology: This study was con-

ducted during two oceanographic campaigns

involving the collection of stratified vertical

zooplankton samples. The first campaign took

place aboard the ARC Providencia in Septem-

ber 2012, as part of the ERFEN XV-CPC LI

oceanographic cruise. Sampling was performed

across a predefined grid comprising ten bio-

logical stations. The second campaign occurred

aboard the ARC Buenaventura in July 2016, tar-

geting a fixed station located in shallow anchor-

ing zones near Malpelo Island. Sampling was

conducted at two-hour intervals between 20:00

h on June 30 and 04:00 h on July 1 (Fig. 1).

Temperature and salinity profiles were

recorded from the surface to a depth of 150 m,

or to the seafloor in shallow anchoring zones,

using a Seabird 19 CTD in 2012 and a YSI-

CastAway CTD in 2016. Vertical variability

in hydrographic conditions was assessed via

continuous profile analyses. Stratified vertical

zooplankton sampling was conducted at each

station using a 1.9 m long conical net (0.3

m mouth diameter, 250 µm mesh), equipped

with a General Oceanic opening and clos-

ing system. In the 2012 campaign, tows were

performed in three discrete depth intervals:

0-50 m, 50-100 m, and 100-150 m. During the

2016 campaign, modified depth strata were

sampled at 0-20 m, 20-50 m, and 50-90 m, tai-

lored to the shallower bathymetry near Malpelo

Island. Zooplankton samples were preserved in

buffered formaldehyde prepared in seawater at

a final concentration of 4 % and transported

to the laboratory for taxonomic analysis, and

were subsequently sorted, counted, and identi-

fied to the lowest feasible taxonomic level using

standard ichthyoplankton identification keys

(Beltrán-León & Rios-Herrera, 2000; Moser,

1996; Richards, 2005). The identified speci-

mens were deposited in the Zoological Practice

Collection of the Department of Biology at

Universidad del Valle.

Sampling analysis: Standardized larval fish

abundance was calculated according to estab-

lished protocols (Smith & Richardson, 1977;

Moser, 1996), with methodological updates

reflecting recent advances in ichthyoplankton

survey design (Ma et al., 2025). Abundance was

expressed as individuals per square meter (ind/

m²) by dividing the number of larvae collected

by the volume of filtered water per tow and

subsequently multiplying by the vertical extent

of the sampled stratum. Total larval abun-

dance per station was derived by summing the

abundance values obtained across all stratified

depth intervals.

Species richness of marine fish larvae in

the study area was estimated to be using the

non-parametric Jackknife one estimator. This

approach, which emphasizes the frequency of

singleton taxa across sampling units, is par-

ticularly suited to ichthyoplankton commu-

nities marked by high proportions of rare

species and constrained sampling effort (Cas-

tilho et al., 2016). By reducing bias associated

with underrepresentation of low-abundance

taxa, Jackknife 1 provides a robust approxima-

tion of expected richness. Its application is

especially recommended in tropical aquatic

environments, where spatial and temporal het-

erogeneity frequently limits sampling represen-

tativeness (Jaonalison et al., 2020).

Vertical variations in larval fish abundance

were evaluated using data from the 2012 sam-

pling campaign. To quantify inter-strata abun-

dance shifts, the Vertical Distribution Index

(VDI) was calculated as: VDI = ln (n1/n2),

where n1 and n2 represent larval abundances

(ind/m²) in two contiguous depth layers. This

index accounts for relative abundance gradients

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in the vertical plane and is sensitive to diel

fluctuations, given that sampling tows were

conducted at various times throughout the

day. Additionally, a complementary graphical

assessment of larval abundance patterns was

conducted at a fixed station during the 2016

campaign. Samples were collected every two

hours across three predefined depth strata,

enabling temporal resolution of vertical migra-

tory behavior under island-influenced hydro-

graphic conditions.

To characterize the structure and heteroge-

neity of larval fish assemblages, we applied two

complementary ecological diversity indices:

Shannon (H’) and Simpson (D). The Shannon

index, is sensitive to species richness and even-

ness, making it well-suited for detecting spatial

shifts in community diversity (Sponaugle et

al., 2002). The Simpson index, by contrast,

emphasizes dominance by weighting the most

abundant taxa more heavily, providing insight

into community resilience and species even-

ness (Magurran, 2004). These indices have

been widely validated in marine ecological

studies and are recommended for biodiversity

assessments in tropical environments (Cowen

& Sponaugle, 2009; Guyah et al., 2021; Paris &

Cowen, 2004). To assess statistical differences

in diversity and dominance between near-

shore and offshore assemblages, we employed a

permutation-based comparison using 999 itera-

tions. Permutation tests are non-parametric

and distribution-free, making them especially

appropriate for ecological datasets that exhibit

non-normality and zero inflation (Anderson,

2001). This approach enhances inference qual-

ity while minimizing assumptions about under-

lying data structure.

To compare total larval fish abundance,

community composition, and species turn-

over between nearshore (0.5 km) and off-

shore (12 km) zones around Malpelo Island,

three statistical approaches were employed:

the non-parametric Mann-Whitney U test, the

Bray-Curtis similarity index, and Whittaker’s

beta diversity metric. The Bray–Curtis index

quantified compositional dissimilarity based

on relative abundance data, offering robustness

against the disproportionate influence of domi-

nant taxa and suitability for ecological datasets

(Hardersen & La Porta, 2023). Whittaker’s beta

diversity provided an integrative measure of

species turnover across spatial units, enabling

assessment of heterogeneity and connectivity

in pelagic larval assemblages (Maslakova et al.,

2022). Both metrics have demonstrated efficacy

in recent ichthyoplankton research, allowing for

the detection of spatial aggregation, dispersal

patterns, and biophysical coupling modulated

by environmental gradients (Zhao et al., 2025).

RESULTS

Surface temperature ranged from 26.6 to

27.3 °C across both sampling periods, while

surface salinity varied between 30.5 and 32.5

UPS in 2012, and from 33.1 to 33.9 UPS in

2016 (Fig. 2). The 21 °C isotherm and the 34

UPS isohaline were found at shallower depths

near Malpelo Island (30 m, 2016 campaign)

compared to the surrounding oceanic zone (50

m, 2012 campaign), indicating localized strati-

fication patterns influenced by topographic and

hydrographic conditions (Fig. 2).

A total of 93 fish larvae were collected

during two oceanographic sampling campaigns

around Malpelo island in the Eastern Tropi-

cal Pacific: 71 specimens in 2012 and 22 in

2016. These larvae were identified and grouped

into 26 taxonomic units (TUs), representing

eight orders and twelve families. In 2012, the

assemblage was dominated by the family Myc-

tophidae, which accounted for 66.3 % of all

specimens (Table 1), with Diogenichthys lat-

ernatus and Vinciguerria lucetia as the most

abundant taxa. In contrast, the 2016 assemblage

was characterized by a higher relative abun-

dance of the families Engraulidae (27.5 %) and

Clupeidae (22.7 %), with Sardinops sagax and

Anchoa sp. as the dominant taxa (Table 1).

The sampling effort yielded a complete-

ness of 78 %, based on a first-order Jackknife

richness estimator that projected 33.37 spe-

cies, compared to the 26 species recorded.

This discrepancy suggests that additional sam-

pling could reveal further diversity, particularly

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among rare or low-abundance taxa. Composi-

tional similarity between zones was relatively

low (Bray-Curtis index = 0.41), underscoring

pronounced ecological differentiation across

the marine landscape. Furthermore, no sta-

tistically significant differences in larval fish

abundance were detected between nearshore

and offshore zones surrounding Malpelo Island

(Mann–Whitney test, p = 0.41). However,

significant differences were observed in both

diversity and dominance metrics. Shannon

diversity was notably higher offshore (Shan-

non index: Inshore = 1.48, Offshore = 2.03, p

= 0.001), while dominance was lower offshore

as indicated by the Simpson index (Inshore =

0.70, Offshore = 0.79, p = 0.045), suggesting

greater evenness and heterogeneity in offshore

assemblages. Additionally, β-diversity analysis

revealed a high species turnover rate (Whit-

taker index = 0.71) between the zones, indi-

cating substantial species replacement along

an approximately 12 km spatial gradient. This

turnover would be strongly influenced by infre-

quent taxonomic groups (Table 1), implying the

presence of environmental discontinuities or

oceanographic fronts that may modulate larval

distribution patterns in the region.

Fish larvae exhibited distinct diel verti-

cal distribution patterns throughout the sam-

pling periods. During daylight hours, larvae

predominantly aggregated within the 50-100

m depth stratum, whereas at night they were

Fig. 2. Vertical variation of temperature and salinity around Malpelo Island during the study period. The average record

(black line) and the maximum and minimum records (gray lines) are presented. n2012 = 10, n2016 = 5. Dotted line shows depth

of 21°C isotherm and 34 UPS isohaline.

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primarily concentrated in the surface layer

(0-50 m) of the water column (Fig. 3A). This

behavior suggests short-scale vertical migra-

tory movements, likely associated with predator

avoidance and foraging efficiency. A similar

trend was observed during the 2016 sampling

period, when larval abundance peaked in the

upper stratum between 20:00 and 02:00, fol-

lowed by a gradual decline thereafter (Fig. 3B).

DISCUSSION

This study provides the first comprehen-

sive characterization of larval fish assemblag-

es around Malpelo Island, revealing distinct

spatial and vertical distribution patterns. The

assemblages exhibited high taxonomic diver-

sity, including mesopelagic, epipelagic, and

reef-associated taxa, with diel vertical migra-

tion modulated by thermocline depth. In oce-

anic island systems, larval fish assemblages are

typically structured by complex interactions

among hydrographic conditions, habitat het-

erogeneity, and dispersal dynamics (Cowen

& Sponaugle, 2009; Sponaugle et al., 2002).

Around Malpelo Island, elevated species rich-

ness and high turnover rates highlight the eco-

logical complexity of the nearshore–offshore

gradient. These patterns align with previous

findings that island wakes, mesoscale eddies,

Table 1

Identities of the fish larvae taxonomic units (TU) collected during the 2012 campaign and 2016 sampling at Malpelo island.

Orden Familia Especie 2012 AB (%) 2016 AB (%)

Anguilliformes Muraenidae TU Sp. 1 1.6

Clupeiformes Clupeidae Sardinops sagax 22.7

Engraulidae Anchoa sp. 18.1

Cetengraulis mysticetus 9.4

Stomiiformes Phosichthyidae Vinciguerria lucetia 26.2 9.1

Aulopiformes Paralepipidae Lestidiops neles cf. 1.6

Myctophiformes Myctophidae Diaphus sp. 3.3

Diogenichthys laternatus 42.6 9.1

Hygophum sp. 2.1

Lampanyctus sp. 1 1.6

Lampanyctus sp. 2 1.6

Myctophum sp. 2.1

Nannobrachium sp. 1 1.6

Nannobrachium sp. 2 1.6

Symbolophorus sp.1 1.5

TU Sp. 2 1.6

TU Sp. 3 1.6

TU Sp. 4 2,1

TU Sp. 5 1.6

TU Sp. 6 3.5

Gadiformes Bregmacerotidae Bregmaceros bathymaster 1.6

Beloniformes Hemiramphidae TU Sp. 7 2.1

Carangiformes Coryphaenidae Coryphaena hippurus 9.4

Perciformes Haemulidae Anisotremus sp. 9.1

Blenniformes Gobiesocidae Gobiesox sp. 4.5

Labriformes Labridae Xyrichthys sp. 4.5

poor condition 0.6 4.1

AB: relative abundance (% Larvae/m2). Phylogenetic organization according to Betancur et al. (2017).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026184, enero-diciembre 2026 (Publicado Feb. 04, 2026)

and bathymetric discontinuities can generate

retention zones or dispersal barriers, thereby

influencing larval community composition

over relatively short spatial scales (Lobel &

Robinson, 1986; Macedo-Soares et al., 2012).

The pronounced turnover observed between

sampling zones may reflect shifts in habitat

structure or hydrographic discontinuities that

compartmentalize larval habitats, particularly

for species with brief planktonic durations or

specialized behavioral traits.

The thermal and saline conditions record-

ed during this study were comparable to those

reported by Rodríguez-Rubio and Giraldo

(2011) for the second semester of the year.

Notably, the thermocline and halocline were

shallower near Malpelo Island than in adja-

cent offshore areas, likely due to the island

mass effect on local circulation (de Falco et al.,

2022). This phenomenon—where the physi-

cal presence of an island obstructs flow and

induces vertical compression of water masses—

is known to enhance micro-upwelling and

nutrient availability, thereby increasing surface

chlorophyll-a concentrations (Andrade et al.,

2014; Bourdin et al., 2025; Caldeira et al.,

2005; Hasegawa et al., 2009; Messié et al.,

2020; Rodríguez et al., 2001). Such conditions,

combined with the availability of habitat and

refugia, contribute to the designation of oce-

anic island systems as biodiversity hotspots and

critical sources of eggs and larvae (Boehlert &

Mundy, 1993; Bowen et al., 2013; Cortés, 2012;

Cowie & Holland, 2006; de Santana et al., 2020;

Hernández-León, 2009; Morato et al., 2010;

Requena et al., 2020).

Larval assemblages near Malpelo Island

were taxonomically diverse, comprising meso-

pelagic species such as Diogenichthys laternatus

and Vinciguerria lucetia (Myctophidae), epi-

pelagic taxa including Sardinops sagax (Clu-

peidae) and Anchoa sp. (Engraulidae), and

reef-associated species like Anisotremus sp.

(Haemulidae) and Coryphaena hippurus (Cory-

phaenidae). This mixed composition is charac-

teristic of oceanic island environments, where

Fig. 3. A. Vertical Distribution Index (IDV) of fish larvae in relation to sampling time at Malpelo Island during 2012. The

shaded region indicates the nocturnal period. B. Temporal variation in fish larval abundance across three depth strata from

June 30 to July 1, 2016, near Malpelo Island, highlighting diel vertical migration patterns.

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 74: e2026184, enero-diciembre 2026 (Publicado Feb. 04, 2026)

larval fish from reef, epipelagic, and mesopelag-

ic habitats co-occur due to localized retention

and habitat complexity (Boehlert & Mundy,

1993; Hunte et al., 1995; Kitchens & Rooker,

2014; Logerwell & Smith, 2001; Macedo-Soares

et al., 2012; Oxenford, 1999; Sassa et al., 2004;

Moyano et al., 2009).

Vertical distribution patterns revealed diel

migratory behavior, with larvae aggregating

at 50-100 m during daylight and ascending

to 0-50 m at night. This behavior has been

widely documented in both coastal and oceanic

systems (Auth et al., 2007; Boehlert & Mundy,

1993; Boehlert et al., 1992; Compaire et al.,

2021; Govindarajan et al., 2023; Hawes et al.,

2020; Sabatés, 2004; Suntsov, 2002; Suthers et

al., 2006). The depth of the thermocline appears

to modulate the extent of these movements,

serving as a key determinant of larval vertical

positioning (Ahlstrom, 1959; Alvarez et al.,

2021; Boehlert et al., 1985; Leis, 2006; Moser

& Pommeranz, 1999; Smith & Suthers, 1999;

Sutton, 2013). Numerous hypotheses have been

proposed to explain diel vertical migration,

including predator avoidance (Röpke, 1993),

prey acquisition (Munk et al., 1989), transport

optimization (Ospina-Alvarez et al., 2018; Paris

& Cowen, 2004), energetic efficiency (Gray,

1996; Lin et al., 2012), and UV radiation avoid-

ance (Heath et al., 1988).

The dominance of Myctophidae in 2012

may reflect mesopelagic spawning synchro-

nized with stratified upper layers and stable

retention conditions (Gartner-Jr, 1991; Rodrí-

guez & Castro, 2000). In contrast, the preva-

lence of Sardinops sagax and Anchoa sp. in

2016 near the fixed station appears linked to

episodic upwelling and the intrusion of Equa-

torial Surface Water, as observed in ichthyo-

plankton studies correlating spawning intensity

with nutrient pulses (Beltrán-León & Morales

Osorio, 2021; González-Quirós et al., 2003;

Twatwa et al., 2005). These taxonomic transi-

tions reinforce Malpelo Island’s role as both a

larval retention hotspot and a potential node

within the Eastern Tropical Pacific Marine

Corridor (CMAR). Biophysical modeling stud-

ies have shown that larval dispersal across

the ETP is modulated by mesoscale eddies

and intermittent current reversals, which can

create transient corridors for export or reten-

tion (León-Chávez et al., 2010; Romero-Torres

et al., 2018). Malpelo’s steep topography and

surrounding stratification may promote local-

ized larval residency, particularly for endemic

and low-dispersal species such as Axoclinus

rubinoffi and Lepidonectes bimaculatus, whose

life histories suggest high site fidelity and lim-

ited recruitment beyond the island’s influence

(Rojas-Vélez et al., 2021). Collectively, these

findings underscore the ecological significance

of Malpelo’s oceanographic setting and its stra-

tegic value for regional conservation planning

within the CMAR framework.

Comparative studies from other oceanic

islands, such as the Hawaiian archipelago (e.g.,

Boehlert & Mundy, 1993; Cowie & Holland,

2006), provide valuable context for interpreting

our findings. These systems similarly exhibit

mixed larval assemblages composed of reef-

associated, epipelagic, and mesopelagic taxa,

shaped by localized retention, diel vertical

migration, and stratification-driven larval posi-

tioning. Malpelo Island shares several of these

hydrographic and ecological features, includ-

ing a compressed thermocline and halocline,

illustrated in figure 2 through mean water col-

umn profiles of temperature and salinity. These

patterns suggest vertical compression of water

masses due to topographic obstruction, consis-

tent with the island mass effect. However, Mal-

pelo’s extreme isolation, steep bathymetry, and

location within the CMAR corridor introduce

distinct dispersal constraints and endemism

dynamics not typically observed in multi-island

systems. While the island mass effect is not the

central focus of this study, the observed hydro-

graphic structure and high β-diversity between

nearshore and offshore assemblages suggest its

localized presence. We propose this mechanism

as a plausible factor influencing larval retention

and spatial turnover, and recommend future

studies incorporate detailed hydrographic pro-

filing to further elucidate these dynamics.

Species identification was based on estab-

lished morphological keys (Moser, 1996;

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 74: e2026184, enero-diciembre 2026 (Publicado Feb. 04, 2026)

Richards, 2005), but we recognize that larval

stages of several taxa exhibit overlapping traits.

Genetic confirmation was not conducted in

this study, representing a limitation that future

research should address through DNA barcod-

ing approaches (Lira et al., 2023). However,

our study reveals that larval fish assemblages

around Malpelo Island exhibit distinct spatial

and vertical distribution patterns, with mesope-

lagic species dominating offshore waters and a

heterogeneous mix of mesopelagic, epipelagic,

and reef-associated taxa concentrated near the

island. This compositional gradient mirrors

patterns documented around other oceanic

islands, underscoring the ecological singular-

ity of insular marine systems. Moreover, the

observed diel vertical migration—characterized

by larval aggregation in deeper strata during

daylight hours and ascent to shallower layers at

night—is modulated by the local thermocline

depth and reflects behavioral strategies con-

sistent with those reported in both coastal and

pelagic environments. These findings reinforce

the ecological significance of oceanic islands as

biodiversity hotspots and critical habitats for

early fish life stages.

To our knowledge, this is the first study to

analyze larval fish assemblages around Malpelo

Island, addressing a notable gap in ichthyo-

plankton research for the region, and provide

essential baseline data for biodiversity assess-

ments and inform conservation planning with-

in this UNESCO World Heritage Site. Future

research should adopt integrative approaches

that combine larval ecology, biophysical mod-

eling, and long-term monitoring to elucidate

the mechanisms driving assemblage structure

and support evidence-based management of

isolated island ecosystems in the Eastern Tropi-

cal Pacific.

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

section. A signed document has been filed in

the journal archives.

ACKNOWLEDGMENTS

We thank Vanessa Izquierdo, Marisol Rive-

ra, and the crews of the B.O. ARC Providen-

cia and ARC Buenaventura for their valuable

help during the field stage of this research.

This work was supported by the Universidad

del Valle (Oceanographic Sciences Research

Group), General Maritime Directorate of the

Republic of Colombia (DIMAR), Pacific Pollu-

tion Control Center (CCCP), National Natural

Parks of Colombia (NNP) - Malpelo Fauna

and Flora Sanctuary (SFFM), and the Young

Researchers and Innovators program Virginia

Gutiérrez de Pineda of Colciencias (grant 617-

2013 to FMH).

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