Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025200, enero-diciembre 2025 (Publicado Nov. 29, 2025)

Habitat use of the Pacific eagle ray Aetobatus laticeps

(Myliobatiformes: Aetobatidae) in a marine protected area

of touristic importance, Banderas Bay, Mexico

Alejandra P. Castelo-Corona¹,2*; https://orcid.org/0000-0003-3378-4297

Adrián Maldonado-Gasca3; https://orcid.org/0000-0003-2067-7062

Juan C. Pérez-Jiménez4; https://orcid.org/0000-0002-7258-1700

Javier Tovar-Ávila5; https://orcid.org/0000-0001-9095-5801

1. Aetos ID, Pacific Eagle Ray Investigation and Conservation Project, Jalisco, México; alejandracasteloc@gmail.com

(Correspondence*)

2. Red de Observadores Ciudadanos, CP 23060, La Paz, Baja California Sur, México.

3. Tecnológico Nacional de México, Bahía de Banderas, Nayarit, México; adrian.mg@bahia.tecnm.mx

4. Laboratorio de Ecología Pesquera, El Colegio de la Frontera Sur. Campeche, México; jcperez@ecosur.mx

5. Instituto Mexicano de Investigación en Pesca y Acuacultura Sustentables, Centro Regional de Investigación Acuícola y

Pesquera, Bahía de Banderas, Nayarit, México; javier.tovar@imipas.gob.mx

Received 01-III-2025. Corrected 19-IX-2025. Accepted 17-XI-2025.

ABSTRACT

Introduction: Aetobatids have low reproductive potential, with coastal and benthopelagic habits, which makes

them vulnerable to anthropogenic activities. Despite this, the Pacific eagle ray Aetobatus laticeps is a poorly

studied species, which makes it difficult to assess its populations to implement or improve conservation and

management measures.

Objective: To determine the habitat use of the Pacific eagle ray in Los Arcos (LA), Banderas Bay, Central Mexican

Pacific, a protected natural area of tourist importance.

Methods: From December 2015 to July 2017, 64 marine surveys were conducted to record the activity and size

of eagle rays in three transects delimited by tourist use in LA. The variability in the seasonal presence of adults

and juveniles, the trend in the space-time encounter rate, and its relationship with sea surface temperature (SST)

were determined.

Results: The encounter rate was higher in April and May (2016 and 2017), decreasing to zero during July, August,

and September of 2016 (with the warmest SST) and February 2017 (with the coldest SST). Displacement activity

predominated, followed by feeding activity. Recorded sizes ranged from 50-130 cm disc width (DW) (mean = 89

± 17 cm). The site with the highest encounter rate coincides with the area of greatest tourist activity.

Conclusions: This study provides evidence of the gregarious behaviour of juveniles and adults, their seasonal

movements, and their relationship with SST at a limited spatial scale. Finally, it is essential to raise awareness

among local communities in LA, as tourist activity may indirectly impact A. laticeps.

Key words: batoid; natural protected area; marine megafauna; conservation management; elasmobranch.

https://doi.org/10.15517/29wq2j10

AQUATIC ECOLOGY

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INTRODUCTION

The pacific eagle ray Aetobatus laticeps

(Gill, 1867) is a species classified as “Vulner-

able” by the International Union for Conserva-

tion of Nature (IUCN); its populations are at

risk of declining due to its low fecundity and

the coincidence of its distribution with vari-

ous fisheries (Pollom et al., 2021). Therefore,

generating information is necessary as a tool

for its conservation and management. This

species was recently separated from Aetobatus

narinari (Euphrasen, 1790) based on genetic

evidence (Richards et al., 2009; Sales et al.,

2019). It is considered to be poorly studied, and

information from nearby species is relied upon

to understand some biological and ecologi-

cal aspects, such as habitat use. For instance,

A. narinari is known for seasonal migrations,

with sea temperature being the primary factor

influencing its movements (Bassos-Hull et al.,

2014; Cuevas-Zimbrón et al., 2011; Cuevas-

Zimbrón et al., 2013). Likewise, patterns of site

fidelity and return movements of A. narinari to

specific areas have been identified, such as per-

manent or seasonal residence (DeGroot et al.,

2021; Flowers et al., 2017). The repeated use of

specific sites suggests that these areas may play

a critical role in the life cycle of these species,

providing essential biotic and abiotic resources.

As a result, human activities can exert hetero-

geneous impacts across their distribution range

(Flowers et al., 2017). Hence, spatio-temporal

research on habitat function is fundamental

for assessing the interactions between elasmo-

branchs and anthropogenic activities (Becerril-

García et al., 2022).

Banderas Bay (BB) is situated in the central

Mexican Pacific (Fig. 1) and is recognized as an

important shark and ray area (ISRA), function-

ing as both a breeding and feeding ground for

A. laticeps (International Union for Conserva-

tion of Nature - Shark Specialist Group [IUCN

SSC], 2023). This site includes the Flora and

Fauna Refuge, Los Arcos (LA), an insular area

located 105 m offshore, where pacific eagle rays

aggregate. Furthermore, commercial and sport

fishing is prohibited (Secretaría de Industria

RESUMEN

Uso de hábitat de la raya águila del Pacífico, Aetobatus laticeps (Myliobatiformes: Aetobatidae)

en un área marina protegida de importancia turística, Bahía de Banderas, México

Introducción: Los Aetobátidos tienen un bajo potencial reproductivo, con hábitos costeros y bentopelágicos, lo

que los hace vulnerables a las actividades antropogénicas. A pesar de ello, la raya águila del pacífico Aetobatus

laticeps es una especie poco estudiada, lo que dificulta la evaluación de las poblaciones para implementar o mejo-

rar medidas de conservación y gestión.

Objetivo: Determinar el uso de hábitat de la especie en Los Arcos (LA), Bahía de Banderas, Pacífico central

mexicano, un área natural protegida de importancia turística.

Métodos: Desde diciembre 2015 hasta julio 2017, se realizaron 64 censos marinos para registrar la actividad y el

tamaño de las rayas águila en tres transectos delimitados por el uso turístico en LA. Se determinó la variabilidad

en la presencia estacional de juveniles y adultos, la tendencia en la tasa de encuentro espacio-temporal y su rela-

ción con la temperatura superficial del mar (TSM).

Resultados: La tasa de encuentro fue mayor en abril y mayo (2016 y 2017), disminuyendo a cero durante julio,

agosto y septiembre de 2016 (con la TSM más cálida) y febrero 2017 (con la TSM más fría). Predominó la acti-

vidad de desplazamiento, seguida de la actividad de alimentación. Las tallas registradas fueron de 50-130 cm de

ancho de disco (promedio = 89 ± 17 cm). El lugar con la mayor tasa de encuentro coincide con la zona de mayor

actividad turística.

Conclusiones: Este estudio aporta pruebas del comportamiento gregario de juveniles y adultos, sus movimientos

estacionales y su relación con la TSM a una escala espacial limitada. Por último, es importante sensibilizar a las

comunidades locales en LA, ya que la actividad turística puede afectar indirectamente a A. laticeps.

Palabras clave: batoideo; área natural protegida; megafauna marina; manejo para la conservación; elasmobranquio.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025200, enero-diciembre 2025 (Publicado Nov. 29, 2025)

y Comercio, 1975) in LA, with tourism being

the main activity and an important source of

income for local communities. However, there

are currently no regulations in place to man-

age the number of tourists visiting the area in

LA, this has resulted in modifications to the

environment (Balzaretti-Merino et al., 2021;

Medina-Rosas, 1997). Considering the gregari-

ous and philopatric characteristics of aetobatids

in certain areas, it can be inferred that these

species may be vulnerable to changes in local

environmental factors or the impact of human

activities, such as fishing or unsustainable tour-

ism practices. In this context, understanding

the habitat use of A. laticeps in aggregation

areas like LA is valuable for establishing man-

agement measures.

Direct observations can be used to assess

habitat use in a non-invasive manner, provid-

ing accurate data on the activities of organ-

isms in situ, at small spatial scales and over

long periods of time (Carraro & Gladstone,

2006; Economakis & Lobel, 1998). This method

yields information on the location and number

of organisms, as well as their behaviour and

biological characteristics, such as size and asso-

ciations with physical, biological factors or

human disturbances (Simpfendorfer & Heupel,

2004). The aim of this study was to evaluate

the habitat usage of A. laticeps in LA during

peak tourist activity hours, considering their

behaviour, individual size, and the number of

sightings in relation to sea surface temperature,

through direct observations.

MATERIALS AND METHODS

Study area: Los Arcos (LA) is comprised

of five islets located between 105 and 497 m

offshore, Southwest of Puerto Vallarta, Jalisco,

Mexico (Fig. 1). The area is dominated by rocky

reef and some patches of sand (Medina-Rosas

& Cupul-Magaña, 2002). These islets are eas-

ily accessible, since they are close to the coast

where there are several communities dedicated

to tourism, and is one of the most frequently

visited sites in BB for scuba diving, free diving,

kayaking, paddle boarding and boat tours. The

Fig. 1. Map of Los Arcos in Banderas Bay. On gray, tourist activity and boat transit zone. Lines indicate different transects.

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025200, enero-diciembre 2025 (Publicado Nov. 29, 2025)

sites where the pacific eagle ray surveys were

conducted are shallow (3-12 m).

Data collection: From December 2015 to

July 2017, 64 surveys were conducted (one per

week), by free diving during the hours of great-

est tourist influx (08:00 to 15:00 h), in three

transects that were previously delimited based

on the characteristics of the area, such as depth,

type of seabed, areas of boat traffic and tourist

activity (Fig. 1). During the surveys, the num-

ber of eagle ray observations was recorded, and

the disc width (DW) of each eagle ray was esti-

mated at a distance of ~ 30 to 50 cm from the

individuals, using a “T” ruler measuring 50 cm

(horizontal) and 100 cm (vertical), with gradu-

ations every 10 cm (English et al., 1997). When

rays were observed in groups (> 2 individuals),

DW was visually estimated to avoid disturb-

ing other organisms. From April 2016 to July

2017, the activity of eagle rays was documented

following the classification by González-Pérez

and Cubero-Pardo (2010), categorizing them as

cleaning (symbiotic fishes associated with eagle

rays, which remove parasites from their bodies,

clean wounds, or feed on dead skin), swimming

(when individuals move in a specific direction),

foraging (individuals searching or feeding in

the sand or among rocks in search of food),

or reproduction (individuals observed during

the mating or courtship process, according by

Swider et al., 2017).

Data analysis: to standardize the number

of eagle ray observations by sampling time, the

encounter rate (ER) was calculated:

ER=n/t

Where: n is equal to the number of eagle ray

observations per sample, and t is the total num-

ber of hours for each sampling.

In order to describe the activities of A.

laticeps in each transect, the activity index (Kar-

czmarski et al., 2000) was used:

AI=B/T

Where: B is the number of observations of eagle

rays engaged in a particular activity, and T is

the total number of observations. The activity

index indicates dominance of an activity when

it is close to one, and scarcity of the activity

when it is close to zero.

The habitat preference index (Ajemian et

al., 2012) was used to define temporal prefer-

ence between transects:

Wi=Oi/πi

Where: Oi is the proportion of the number of

eagle rays observations per month in transect

‘x’ relative to the total number of rays from all

transects per month, πi is the proportion of the

area of transect ‘x’ relative to the total area of all

transects. This index indicates positive selec-

tion for an area when it is greater than one, and

avoidance when it is less than one.

The Kruskal-Wallis test was used to deter-

mine whether there is a difference in length

between months and in monthly ER. If sig-

nificant differences were found, the Nemenyi

post-hoc test was applied (Pohlert, 2014). A

polynomial regression test was conducted to

assess the relationship between ER and sea

surface temperature (SST). The SST data were

acquired from MODIS-Aqua (NASA OBPG,

2019) for the 2015-2017 period through the

Environmental Research Division Data Access

Program (ERDDAP) hosted by the NOAA

Southwest Fisheries Science Center (Nation-

al Oceanic and Atmospheric Administration

[NOAA], 2023).

RESULTS

During the 64 marine surveys conducted, a

total of 181 sightings of A. laticeps were record-

ed, with an overall average per survey of 2.82 ±

3.67 observations events. The average ER was

1.56 ± 2.18, with the highest ER observed dur-

ing April and May, whereas no eagle ray sight-

ings were recorded in August and September

(Fig. 2), with significant differences between

May with August/September in 2016 (p < 0.05).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025200, enero-diciembre 2025 (Publicado Nov. 29, 2025)

A total of 147 activity records of A. laticeps

were obtained, where “swimming” was the

activity most observed (87.07 %), followed by

“foraging” (10.88 %), “cleaning” (1.36 %), and

“breeding” (0.68 %) (Table 1). The habitat pref-

erence index was > 1 for transect three com-

pared to transects one and two in most months

(January, March-June, and October-December

2016, and January, March-July 2017), which

coincided with the highest ER values (Fig. 2).

The recorded lengths of A. laticeps dur-

ing the surveys ranged 50 to 130 cm DW

(mean 88.83 ± 17 cm DW) approximately.

For this study, individuals ≥ 90 cm DW were

considered adults based on previous length

at maturity observations (Vázquez 2003 in

Salomón-Aguilar, 2015), which also coincide

with our records of some pregnant females

with ≥ 90 cm DW in BB (Fig. 3). From the

181 sightings, 114 were classified as adults and

67 as juveniles (Fig. 4), with a ratio of 1:0.58.

Additionally, there were differences in the total

number of sightings between juveniles and

adults (x2 = 12.2, p ≤ 0.01), but not among the

sizes per month (Kruskall-Wallis, p = 0.8195,

x² =10.835, df = 9).

During the sampling period the SST ranged

23-30.5 °C. The correlation between the ER and

Table 1

Activity index (AI) of Aetobatus laticeps per transect at Los Arcos from December2015 to July 2017.

Activity AI transect 1 AI transect 2 AI transect 3 Total AI

Foraging 0.31 (5) 0.71 (5) 0.04 (6) 0.10

Breeding 0.00 0.00 0.008 (1) 0.006

Swimming 0.68 (11) 0.28 (2) 0.92 (115) 0.87

Cleaning 0.00 0.00 0.01 (2) 0.013

Number of observations events are in parenthesis.

Fig. 2. Monthly encounter rate (shaded area) and habitat preference index for three transects surveyed from December 2015

to July 2017 in Los Arcos.

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SST was low, but significant (r = 0.2128, p <

0.001), it is possible that there are other factors

that influence ER, such as salinity, tidal flow,

and dissolved oxygen. It is important men-

tioning that when the SST fluctuated between

25-27 °C, the highest ER occurred, whereas

when the highest (30 °C) and lowest (23 °C)

SST occurred, eagle rays sightings decreased to

zero (Fig. 5).

DISCUSSION

This study presents novel information on

the habitat use of A. laticeps obtained through

Fig. 3. Female pacific eagle rays visibly in gestation period, with approximately 90 cm DW.

Fig. 4. Number of juveniles and adults of pacific eagle rays observed during the sampling period.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025200, enero-diciembre 2025 (Publicado Nov. 29, 2025)

direct observations in a tourist area in the Mex-

ican Pacific. Los Arcos harbors a high diversity

and abundance of mollusks in Banderas Bay

(Ríos-Jara et al., 2001), which are an important

part of the diet of pelagic eagle rays (Ajemian

et al., 2012; Schluessel et al., 2010; Serrano-

Flores et al., 2018). This area is one of the

well-known spots for divers in the BB where A.

laticeps forms aggregations (Balzaretti-Merino

et al., 2021). Considering the observations of

individuals feeding, it is likely that eagle rays

frequent LA due to the high availability of prey,

as is the case of A. narinari in Bermuda and the

Southern Gulf of Mexico (Ajemian et al., 2012;

Serrano-Flores et al., 2018).

The area where eagle rays showed habitat

preference was specifically located between

two islets where strong currents form and vary

in direction and intensity throughout the day.

This could benefit the rheotaxis of Aetobatus

laticeps by facilitating water flow through its

gills. Aquatic activities are common in this area,

with high boat traffic. Occasionally, eagle rays

have been observed quickly fleeing when boats

pass nearby, similar to the behaviour described

by Berthe and Lecchini (2016) for Aetobatus

ocellatus. However, in more recent years, an

eagle ray with fresh propeller-inflicted wounds

was recorded in LA, although the frequency of

such incidents is still uncertain (Castelo-Coro-

na & Arnés-Urgellés, 2025). Furthermore, the

long-term effects of interrupting their activities,

such as cleaning or feeding in an aggregation

area like LA, remain unknown.

During most of the sampling period, both

adult and juvenile individuals were observed

in LA, with adults being more prevalent (62

% of the sightings). Their primary recorded

activities were swimming and feeding. Previous

studies have shown that habitat preference can

vary during ontogeny in species of the family

Aetobatidae (Bassos-Hull et al., 2014; Cuevas-

Zimbrón et al., 2011; Degroot et al., 2021), as

well as in several elasmobranch species (Hoyos-

Padilla et al., 2014; Yamaguchi & Taniuchi,

2000). This has mainly been attributed to varia-

tions in food requirements, differences in the

distribution and abundance of prey, and the

ability to evade predators (Palacios et al., 2021).

In Southern BB, the continental shelf is narrow

Fig. 5. Encounter rate variability of Aetobatus laticeps in Los Arcos in relation to sea surface temperature (SST).

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025200, enero-diciembre 2025 (Publicado Nov. 29, 2025)

and drops steeply to over 1 400 m (Plata &

Filonov, 2007). Given the shallow-water prefer-

ence of aetobatids waters (Ajemian et al., 2012;

Ajemian & Powers, 2014), both juveniles and

adults likely remain nearshore. This represents

a lower energy expenditure as they make short-

er vertical foraging trips.

The classification of individuals ≥ 90 cm

DW as adults in this study is supported by the

only available reference on the species’ size at

sexual maturity (Vázquez, 2003 in Salomón-

Aguilar, 2015), although this corresponds to

unpublished gray literature and is not easily

accessible. Nevertheless, this criterion is further

supported by our direct observations (n = 16)

of pregnant females ≥ 90 cm DW recorded dur-

ing and after the study period. This agreement

reinforces the reliability of using the 90 cm DW

threshold to distinguish adults from juveniles,

despite the limited published information on

the reproductive biology of A. laticeps.

In other areas, temperature has been dem-

onstrated as one of the main factors influencing

the movement of pelagic eagle rays (Bassos-Hull

et al., 2014; Bernal et al., 2012; Cuevas-Zimbrón

et al., 2013). This aligns with the absence of

Aetobatus laticeps sightings in LA during peri-

ods of either the warmest or coldest tempera-

tures. This variability in temperature preference

may reflect A. laticeps affinity for intermediate

temperatures to optimize metabolic processes,

potentially due to physiological constraints

(Gleiss et al., 2022). For example, in the case

of A. narinari, higher catches were recorded in

Campeche when sea surface temperature (SST)

was 23.9 °C (Cuevas-Zimbrón et al., 2013),

and observations in Florida occurred primar-

ily within the 23-31 °C range (Bassos-Hull et

al., 2014). Similarly, A. flagellum was captured

in Ariake Sound when SST exceeded 15 °C.

Temperature has been consistently identified as

a key factor driving elasmobranch movements

(Bernal et al., 2012). However, other seasonally

variable factors, such as dissolved oxygen, salin-

ity, and pH, can also influence elasmobranch

behaviour (Schlaff et al., 2014), but were not

considered in this study, leaving the effects of

these factors on the movement of A. laticeps in

the area unknown. Future studies should incor-

porate these environmental variables to better

understand their impact on the habitat use of

this species.

According to the results of this study, LA

represents an aggregation area for Aetobatus

laticeps, where juvenile and adult individuals

primarily engage in feeding activities. Addi-

tionally, recent information indicates that the

area is an important area for the reproduction

of the species (IUCN SSC, 2023). At the same

time, this species holds significant importance

for recreational tourism activities in the area

(Balzaretti-Merino et al., 2021). Despite LA

being a 3 km² protected natural area since 1975,

there is no constant surveillance at the site, and

illegal fishing continues to occur (Stokes et

al., 2019). Furthermore, the area lacks regula-

tions to manage tourism activities. Nearly three

decades ago, (1997) noted that boating, snor-

keling, scuba diving, and fishing in LA already

caused coral damage, invertebrate extraction,

pollution, and sedimentation from anchor-

ing. He suggested implementing conservation

measures to regulate these tourism activities.

Additionally, considering the benthos as the

physical environment that provides the food

source for Aetobatus laticeps, it is likely that

these activities also impact the area as a feeding

zone for this species.

Considering the surface habits of Aeto-

batus laticeps, it is likely to be affected by

collisions with boats, as recently recorded in

Banderas Bay with this species (Castelo-Corona

& Arnés-Urgellés, 2025). Similar incidents have

been documented with other megafauna spe-

cies, such as whales and giant manta rays in

Banderas Bay (Dominguez-Sanchez et al., 2024;

Ransome et al., 2021). Therefore, regulating

boat traffic and speed in this and surrounding

areas is necessary to protect megafauna with

epipelagic habits, ensuring that the movements

of these animals are not disrupted.

Medina-Rosas (1997) estimated a daily vis-

itation of 2 000 to 3 000 people in LA. Current-

ly, considering that tourism demand in Puerto

Vallarta has increased significantly over the last

decade (Secretaría de Turismo de Gobierno del

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025200, enero-diciembre 2025 (Publicado Nov. 29, 2025)

Estado de Jalisco, 2024), it is likely that visita-

tion in LA has also risen due to its easy access.

Therefore, in addition to establishing a tourism

management program, it is imperative to raise

awareness among local residents with con-

tinuous influence in the area. This would aim

to encourage local tourism service providers

and fishers to adopt environmentally respectful

behaviours, fostering a better balance between

social equity, economic growth, and environ-

mental protection in LA and surrounding com-

munities over the long term.

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 all the volunteers from Aetos ID

who participated in the fieldwork at Los Arcos,

and the members of the fisheries laboratory at

ECOSUR-Campeche for their support in struc-

turing the project.

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