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Checklist of invertebrates and conspicuous fishes in rocky reefs

and Sargassum beds in the North Pacific of Costa Rica

Alvin Suárez

1, 2

; https://orcid/org/0000-0003-2394-2685

Rafael Riosmena-Rodríguez

Jorge Cortés

; https://orcid.org/0000-0001-7004-8649

1. Universidad Autónoma de Baja California Sur, La Paz, Baja California Sur, México; SargassumGC@gmail.com

2. Centro de Estudios Biológicos, Medio Ambiente y Recursos Naturales, A.C. Felipe Carrillo Puerto, Quintana Roo,

México.

3. Centro de Investigación en Ciencias del Mar y Limnología, Universidad de Costa Rica, 11501-2060, San José, Costa

Rica; jorge.cortes@ucr.ac.cr

* Deceased

Received 31-I-2021. Corrected 22-III-2021. Accepted 04-V-2021.

ABSTRACT

Introduction: The marine biodiversity is well known in some areas and for some marine ecosystems of the

Pacific coast of Costa Rica. The marine sector of Área de Conservación Guanacaste, in the North Pacific of the

country, is a priority area for conservation of marine and coastal biodiversity.

Objective: Our aim was to increase the knowledge of marine biodiversity in a conservation priority site in Costa

Rica and in poorly or scarcely studied marine ecosystems, such as rocky reefs and Sargassum beds, respectively.

Methods: The study was conducted in Bahía Salinas and Golfo de Santa Elena in 2013. In January, four rocky

reefs were visited; while in February three rocky reefs and one Sargassum bed were visited. We identified the

macroinvertebrates and conspicuous fish species through underwater visual census methods (timed swim and

band transects). The mean density of each species was estimated through band transects method.

Results: A total of 25 invertebrates and 34 fish species were identified. We found a high number of species of

invertebrates (23) and fish species (33) in rocky reefs, but few species on Sargassum beds (eight invertebrates

and ten fish species). Species composition differed between marine ecosystems.

Conclusions: Our study further evidence the limited knowledge of invertebrates and fish communities in these

marine ecosystems in the North Pacific of Costa Rica. Considerably more surveys and field work are required

to support management decisions that are backed by sound scientific knowledge.

Key words: echinoderms, mollusks, fishes, marine ecosystems, conservation priority site, Área de Conservación

Guanacaste.

Suárez, A., Riosmena-Rodríguez, R., & Cortés, J. (2021).

Checklist of invertebrates and conspicuous fishes in

rocky reefs and Sargassum beds in the North Pacific of

Costa Rica. Revista de Biología Tropical, 69(Suppl. 2),

S180-S193. https://doi.org/10.15517/rbt.v69iS2.48316

https://doi.org/10.15517/rbt.v69iS2.48316

The marine biodiversity of the Pacific

coast of Costa Rica is well known in some

areas (i.e., Bahía Culebra, Golfo de Nicoya,

Isla del Caño and Golfo Dulce). Other marine

areas remain poorly studied, with little infor-

mation available or far from complete, like

the north Pacific of Costa Rica (Bahía Salinas

and Golfo de Santa Elena) (Cortés, 2016;

Cortés, 2017; Wehrtmann & Cortés, 2009).

The same occurs regarding coastal ecosystems

since some, such as coral reefs, have been rela-

tively well studied (Alvarado, Beita-Jiménez,

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Mena, Fernández-García & Guzmán-Mora,

2015; Alvarado, Beita-Jiménez, Mena, Fernán-

dez-García, Guzmán-Mora, & Cortés, 2016;

Alvarado Beita-Jiménez, Mena, Fernández-

García, Cortés, Sánchez-Noguera, Jiménez,

& Guzmán-Mora, 2018; Cortés, 1886/1997);

while in others the research and publications on

marine biodiversity are scarce (such as sandy

beaches and rocky coast) (Dexter, 1974; Mad-

rigal-Castro, Cabrera-Peña, Monge-Esquivel &

Pérez-Acuña, 1984; Ortega, 1987; Sibaja-Cor-

dero & Cortés, 2008; Sibaja-Cordero, Cama-

cho-García & Vargas-Castillo, 2014).

The Área de Conservación Guanacaste

(ACG) is one of eleven Conservation Areas

within Costa Rica and it was declared a UNES-

CO World Heritage Site in 1999 (SINAC,

2012; UNESCO, 2020). The marine sector of

ACG, recognized as a conservation priority

site for marine and coastal biodiversity within

Marine Ecological Unit Papagayo in the north

Pacific of Costa Rica (Alvarado, Herrera, Cor-

rales, Asch, & Paaby, 2011; SINAC-MINAET,

2008), has a high species richness and high

diversity coastal and marine ecosystems, such

as mangrove forest, beaches, bays and coves,

rocky intertidal zones, mud flats, rocky subtidal

sites, coral reefs, rhodolith beds, deep areas,

shoals, and several isolated islands (Cortés,

2017; Cortés & Joyce, 2020). During the trade

winds season (December to April-May; Rodrí-

guez, Alfaro & Cortés, 2021) it is possible to

find marine ecosystem dominated by the brown

macroalgae Sargassum liebmannii J.A. Agardh

1847, that grow significantly as a consequence

of the upwelling that brings deep cold, nutrient-

rich and CO

-rich waters to the surface (Cortés,

Samper-Villarreal & Bernecker 2014).

In ACG, including Bahía Salinas (BSa)

and Golfo de Santa Elena (GSE), there are

some publications on invertebrates and fish

communities in coral reef, rocky reef and sandy

areas (Alvarado et al., 2018; Cordero-Umaña

& Santidrián-Tomillo, 2020). Thus, the aim of

the present study is to increase the knowledge

of marine biodiversity in this conservation

area, specifically on BSa and GSE, and on

poorly studied ecosystems like rocky reefs and

Sargassum beds. The study provides a checklist

of invertebrates and conspicuous fishes species

that occur on shallow (6-14 m depth) rocky

reefs and Sargassum beds in ACG. In addition,

patterns of geographic distribution, number

of species, species density and abundance

are discussed.

MATERIAL AND METHODS

Study site: The study was conducted dur-

ing January and February 2013 in BSa and

GSE, within ACG in the North Pacific of Costa

Rica (Fig. 1, Table 1).

Field methodology: In BSa we visited

two sites, while in the GSE six. The marine

ecosystems monitored were shallow (6 to 13.5

m depth) rocky reefs (boulders < 30 cm, small

rocks < 30 cm – 1 m, big rocks 1 m – 3m, and

rocks > 3m) and Sargassum beds (rocky sub-

strate covered with 70% to 100% Sargassum)

Fig. 1. Sampling sites in the North Pacific of Costa Rica.

Rocky reefs: S1, S2, S3, S4, S5, S7 and S8; Sargassum

beds: S6.

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(Fig. 1, Table 1). Underwater Visual Census

(UVC) methods using SCUBA were conducted

at the eight sites during two sampling times

(January and February 2013). In January, four

sites were visited (S2, S5, S7 and S8). At each

site the timed swim method, 50 to 60-minute,

were done to register the invertebrates and

conspicuous fish species. While in February,

the other four sites (S1, S3, S4 and S6) were

visited. Census followed standard band tran-

sect methodology described in previous studies

(Alvarado et al., 2015; Alvarado et al., 2016).

In each marine ecosystem, 10 m long tran-

sects, parallels to shore line and separated 10

m between them, were conducted by a single

diver swimming to register the composition

and abundance of invertebrates and conspicu-

ous fishes. Three to six transects were per-

formed at each site. Two runs were made along

each transect. The first run was done while the

line was being unrolled, recording all conspicu-

ous fishes (> 5 cm total length) in the water col-

umn. We focused on conspicuous species rather

than cryptobenthic species, or small fishes (<

~5 cm in total length) that are behaviorally

cryptic and are difficult to quantify by visual

surveys due to their close association with the

substratum (Allen, Bouvier & Jensen, 1992).

In the second run, the diver recorded all inver-

tebrates (>2.5 cm length) over the benthos and

between crevices. For both UVC techniques,

individuals under rocks were not registered

to avoid altering the substrate. The categories

were mollusks (gastropods and bivalves) and

echinoderms (sea stars, sea urchins, sea cucum-

bers and brittle stars). Cryptic species (brittle

stars) registered in transect bands method were

not quantify to avoid an underestimation. For

fishes, each band transect covered an area of

50 m

(10 x 5 m) and for invertebrates an area

of 20 m

(10 x 2 m). Mean density (# ind m

-2

relative abundance (# ind of one species/total

# individuals), and frequency of occurrence

(# transects observed/total # transects) were

calculated for each species.

Fish species nomenclature and taxonomic

designations followed Fricke, Eschemer & Van

der Laan (2020) and Van der Laan, Fricke &

Eschemeyer (2020), and for invertebrates the

World Register of Marine Species (WoRMS

Editorial Board, 2020). Fish species distribu-

tion were based on Froese & Pauly (2019) and

Fricke et al. (2020), and categorized as fol-

lows: CT, Circumtropical; EP, Eastern Pacific;

TABLE 1

Localities and survey sites in the North Pacific of Costa Rica. UVC: Underwater Visual Census; n: size of the sample.

Locality Survey site ID Geographic coordinates Depth (m) UVC method n

Bahía Salinas Isla Bolaños 1 S1 11°03’02.65’’N

85°42’39.62’’W

9 Band transect 3

Isla Bolaños 2 S2 11°02’45.19’’N

85°42’32.78’’W

9 Timed swim 1

Golfo de Santa Elena La Mesa S3 11°01’37.13’’N

85°45’46.44’W

9 Band transect 6

Isla Loro 1 S4 11°00’10.14’’N

85°44’51.43’W

9 Band transect 6

Isla Loro 2 S5 11°00’03.14’’N

85°44’50.86’’W

13.5 Timed swim 1

Isla Muñecos 1 S6 10°58’53.37’’N

85°43’01.78’’W

9 Band transect 6

Isla Muñecos 2 S7 10°58’51.67’’N

85°43’00.82’’W

11 Timed swim 1

Isla David S8 10°57’23.76’’N

85°43’21.98’’W

6 Timed swim 1

ID: Code of the survey sites in the map.

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IP, Indo-Pacific; ETP, Eastern Tropical Pacific;

WA, Western Atlantic. The IUCN Red List

(IUCN 2020) was used to categorize the con-

servation status of each species recorded in this

study as follows (in order of threat level): NE,

not evaluated; DD, data deficient; LC, least

concern. The categories mentioned correspond

only to the species observed in this study.

RESULTS

A total of 25 invertebrate species in 25

genera and 18 families were observed in BSa

and GSE (Table 2) with the combined use of

timed swim (n = 3) and band transects meth-

ods (n = 21). The most speciose invertebrate

families were the Muricidae (3 spp.), Ophidia-

steridae (2 spp.), Diadematidae (2 spp.), Toxo-

poneustidae (2 spp.), Cucumaridae (2 spp.) and

Ophiocomidae (2 spp.). Hexaplex (2 spp.) was

the most species rich genera, the rest of genera

were represented by one specie. The highest

number of species was observed in the rocky

reef (23 spp.) while in Sargassum beds eight

spp. (Table 2).

Based on classification of the IUCN Red

List, the conservation status of invertebrates

species observed in BSa and GSE is unknow

TABLE 2

Taxonomic list of invertebrates recorded in rocky reefs (RRs) and Sargassum beds (SBs) in the North Pacific

of Costa Rica during January and February 2013, with the IUCN Red List classification. NE: Not Evaluated

Taxon

Marine Ecosystem

IUCN

Red List

RRs SBs

Phylum Mollusca

Class Gastropoda

Family Conidae

Conus spp. Linnaeus, 1758

S1, S5 NE

Family Fasciolariidae

Opeatostoma pseudodon (Burrow, 1815)

S3, S4, S8 NE

Family Muricidae

Babelomurex hindsi (Carpenter, 1857)

S2 NE

Hexaplex erythrostomus (Swainson, 1831)

S3, S4 NE

Hexaplex princeps (Broderip, 1833)

S1, S2, S4, S5 S6 NE

Family Turbinellidae

Vasum caestus (Broderip, 1833)

S6 NE

Family Plakobranchidae

Elysia diomedea (Bergh, 1894)

S4, S7 S6 NE

Class Bivalvia

Family Margaritidae

Pinctada mazatlanica (Hanley, 1856)

S1, S2, S5 NE

Family Spondylidae

Spondylus limbatus G. B. Sowerbey II, 1847

S1, S5 NE

Phylum Echinodermata

Class Asteroidea

Family Mithrodiidae

Mithrodia bradleyi Verrill, 1867

S5, S7 NE

Family Ophidiasteridae

Pharia pyramidata (Gray, 1840)

S5 NE

Phataria unifascialis (Gray, 1840)

S1, S3, S4, S5, S7 NE

Family Oreasteridae

Nidorellia armata (Gray, 1840)

S3 NE

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because those species have not been assessed

by the IUCN (Table 2).

In the case of fishes, a total of 34 species

in 30 genera and 15 families were observed

(Table 3) with the combined use of timed swim

(n = 3) and band transects methods (n = 21).

The most speciose families were Serranidae (5

spp.), Pomacentridae (5 spp.), Labridae (5 spp.)

and Tetraodontidae (4 spp.). Haemulon (3 spp.)

and Halichoeres (3 spp.) were the most species

rich genera. Two genera were represented by

two species, and the rest by one species. The

highest number of fish species was observed

in the rocky reef (33 spp.), while in Sargassum

beds only 10 species (Table 3).

Three percent of species are restricted to

the Eastern Tropical Pacific (Gulf of California

to Peru), whereas 94% occur throughout the

Eastern Pacific (California, USA to Chile).

Only four species are found outside the Eastern

Pacific, one of which is circumtropical (Diodon

holocanthus), one is amphioceanic (Haemulon

steindachneri) and the remaining two occur in

the Indo-Pacific (Arothron hispidus, A. melea-

gris) (Table 3).

Classification by IUCN Red List status

revealed that the fish observed in BSa and GSE

are not species in threatened categories, only of

least concern (Table 3).

We counted a total of 1 964 invertebrate

individuals with the use of band transects (n

= 21). The highest number of individuals was

recorded in the rocky reef, 1 862 ind., while

in Sargassum beds only 102 ind. In the rocky

reef, Diadema mexicanum accounted for 81.3%

of all individuals observed. D. mexicanum

(100%) and Eucidaris thouarsii (93%) ranked

highest in frequency of occurrence, and three

species were observed between 40 to 50% of

the transects (Toxopneustes roseus, Opeatos-

toma pseudodon and Phataria unifascialis). D.

mexicanum has the highest mean density (5.04

Taxon

Marine Ecosystem

IUCN

Red List

RRs SBs

Class Echinoidea

Family Arbaciidae

Arbacia stellata (Blainville, 1825; ?Gmelin, 1791)

S1, S2, S3, S8 NE

Family Diadematidae

Astropyga pulvinata (Lamarck, 1816)

S7 S6 NE

Diadema mexicanum A. Agassiz, 1863

S1, S2, S3, S4, S5, S7, S8 S6 NE

Family Cidaridae

Eucidaris thouarsii (L. Agassiz & Desor, 1846)

S1, S2, S3, S4, S5, S7, S8 NE

Family Toxopneustidae

Toxopneustes roseus (A. Agassiz, 1863)

S1, S2, S3, S7, S8 S6 NE

Tripneustes depressus A. Agassiz, 1863

S2, S7 S6 NE

Class Holothuroidea

Family Cucumaridae

Cucumaria flamma Solis-Marin & Laguarda-Figueras, 1999

S2 NE

Pseudocnus spp. Panning, 1949

S2, S7 NE

Family Holothuriidae

Holothuria (Stauropora) fuscocinerea Jaeger, 1833

S2 NE

Class Ophiurodea

Family Ophiocomidae

Ophiocoma aethiops Lütken, 1859

S4, S5 NE

Ophiocomella alexandri (Lyman, 1860)

S6 NE

Family Ophionereididae

Ophionereis annulata (Le Conte, 1851)

S2 NE

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TABLE 3

Taxonomic list of conspicuous fishes recorded in rocky reefs (RRs) and Sargassum beds (SBs) in the North Pacific of

Costa Rica during January and February 2013, showing geographic distribution and IUCN Red List category.

CT: Circumtropical; EP: Eastern Pacific; EP + IP: Eastern Pacific and Indo-Pacific; EP + WA: Eastern Pacific

and Western Atlantic; ETP: Eastern Tropical Pacific. LC: Least Concern

Taxon

Marine Ecosystem

Distribution

IUCN

Red List

RRs SBs

Order Myliobatiformes

Family Urotrygonidae

Urobatis halleri (Cooper, 1863)

S2, S5 S6 EP LC

Order Anguilliformes

Family Muraenidae

Muraena lentiginosa Jenyms, 1842

S4, S5, S7, S8 EP LC

Order Acanthuriformes

Family Pomacanthidae

Holacanthus passer Valenciennes, 1846

S2, S3, S4, S5, S8 EP LC

Pomacanthus zonipectus (Gill, 1862)

S4, S8 EP LC

Family Chaetodontidae

Chaetodon humeralis Günther, 1860

S1, S2, S3, S4, S5, S7,

S6 EP LC

Johnrandallia nigrirostris (Gill, 1862)

S3, S4, S5, S7, S8 EP LC

Order Tetraodontiformes

Family Diodontidae

Diodon holocanthus Linnaeus, 1758

S2, S3, S4, S5, S7, S8 S6 CT LC

Family Tetraodontidae

Arothron hispidus (Linnaeus, 1758)

S5, S8 EP + IP LC

Arothron meleagris (Anonymus, 1798)

S2, S5 EP + IP LC

Canthigaster punctatissima (Günther, 1870)

S2, S5, S8 S6 ETP LC

Sphoeroides lobatus (Steindachner, 1870)

S2 S6 EP LC

Family Balistidae

Balistes polylepis Steindachner, 1876

S5, S7, S8 S6 EP LC

Order Carangiformes

Family Carangidae

Caranx caballus Günther, 1868

S2, S3, S8 EP LC

Order Perciformes

Family Serranidae

Alphestes immaculatus Breder, 1936

S1, S2, S7 EP LC

Cephalopholis panamensis (Steindachner, 1876)

S2, S3, S4, S5, S7 EP LC

Epinephelus labriformis (Jenyns, 1840)

S1, S2, S3, S5, S7, S8 EP LC

Paranthias colonus (Valenciennes, 1846)

S8 EP LC

Serranus psittacinus Valenciennes, 1846

S6 EP LC

Family Apogonidae

Apogon pacificus (Herre, 1935)

S2, S5, S7, S8 EP LC

Family Lutjanidae

Lutjanus argentiventris (Peters, 1869)

S1, S2, S3, S4, S5, S7,

EP LC

Family Haemulidae

Haemulon maculicauda (Gill, 1862)

S2 EP LC

Haemulon scudderii Gill, 1862

S8 EP LC

Haemulon steindachneri (Jordan & Gilbert, 1882)

S3, S4 S6 EP + WA LC

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± 2.02 ind m

-2

) (Table 4). While in Sargas-

sum beds, T. roseus accounted for 87.3% of

all individuals observed and ranked highest in

frequency of occurrence (83%) and in mean

density (0.74 ± 0.75 ind m

-2

) (Table 4).

We counted a total of 622 fish individuals

with the use of band transects (n = 21). The

highest number of individuals was recorded

in the rocky reef (539 ind.), while only 83 ind.

in Sargassum beds. In the rocky reef, three

Taxon

Marine Ecosystem

Distribution

IUCN

Red List

RRs SBs

Family Pomacentridae

Abudefduf troschelii (Gill, 1862)

S2, S4, S5, S7, S8 EP LC

Chromis atrilobata Gill, 1862

S3, S4, S5, S7, S8 EP LC

Microspathodon dorsalis (Gill, 1862)

S3, S4, S5, S8 EP LC

Stegastes acapulcoensis (Fowler, 1944)

S2, S3, S4 EP LC

Stegastes flavilatus (Gill, 1862)

S2, S4, S5, S7, S8 S6 EP LC

Family Labridae

Bodianus diplotaenia (Gill, 1862)

S2, S3, S4, S5, S7, S8 EP LC

Halichoeres chierchiae Di Caporiacco, 1948

S3, S5 EP LC

Halichoeres dispilus (Günther, 1864)

S3, S4, S7, S8 S6 EP LC

Halichoeres nicholsi (Jordan & Gilbert, 1882)

S5, S7 EP LC

Thalassoma lucasanum (Gill, 1862)

S1, S3, S5, S7, S8 EP LC

Family Scorpaenidae

Scorpaena mystes Jordan & Starks, 1895

S7 EP LC

TABLE 4

List of invertebrates observed in rocky reefs and Sargassum beds in the North Pacific of Costa Rica organized by

total abundance, relative abundance, frequency of occurrence, and mean density (± SD)

Species

Abundance

(# inds)

Relative abundance

(%)

Frequency

(%)

Mean Density

(#ind m

-2

)

ROCKY REEFS

Diadema mexicanum

1513 81.3 1.00 5.04 2.02

Eucidaris thouarsii

192 10.3 0.93 0.64 0.57

Toxopneustes roseus

73 3.9 0.47 0.24 0.47

Opeatostoma pseudodon

43 2.3 0.47 0.14 0.24

Arbacia stellata

14 0.8 0.13 0.05 0.14

Phataria unifascialis

9 0.5 0.40 0.03 0.05

Hexaplex princeps

7 0.4 0.27 0.02 0.05

Hexaplex erythrostomus

3 0.2 0.13 0.01 0.03

Spondylus limbatus

3 0.2 0.07 0.01 0.04

Pinctada mazatlanica

2 0.1 0.07 <0.01 0.03

Conus sp.

1 0.1 0.07 <0.01 0.01

Elysia diomedea

1 0.1 0.07 <0.01 0.01

Nidorellia armata

1 0.1 0.07 <0.01 0.01

SARGASSUM BEDS

Toxopneustes roseus

89 87.3 0.83 0.74 0.75

Diadema mexicanum

4 3.9 0.50 0.03 0.04

Hexaplex princeps

3 2.9 0.33 0.03 0.04

Astropyga pulvinata

3 2.9 0.33 0.03 0.04

Vasum caestus

1 1.0 0.17 <0.01 0.02

Elysia diomedea

1 1.0 0.17 <0.01 0.02

Tripneustes depressus

1 1.0 0.17 <0.01 0.02

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species (Chromis atrilobata, Halichoeres dispi-

lus and Stegastes acapulcoensis) accounting

for 69.8% of all individuals observed. S. aca-

pulcoensis (60%), Bodianus diplotaenia (60%)

and Chaetodon humeralis (53%) ranked high-

est in frequency of occurrence, and five species

were observed on 33% of transects. According

to their mean density (Table 5), C. atrilobata

(0.34 ± 0.52 ind m

-2

), H. dispilus (0.09 ± 0.21

ind m

-2

), S. acapulcoensis (0.07 ± 0.10 ind

-2

) and H. steindachneri (0.05 ± 0.18 ind m

-2

)

ranked highest. While in Sargassum beds, H.

dispilus accounted for 77.1% of all individuals

observed and ranked highest in frequency of

occurrence (83%) and in mean density (0.21 ±

0.18 ind m

-2

) (Table 5).

DISCUSSION

The North Pacific of Costa Rica is a

region of great research interest due to its

TABLE 5

List of conspicuous fishes observed in rocky reefs and Sargassum beds in the North Pacific of Costa Rica organized by

total abundance, relative abundance, frequency of occurrence, and mean density (± SD)

Species

Abundance

(# inds)

Relative abundance

(%)

Frequency

(%)

Mean density

(#ind m

-2

)

ROCKY REEFS

Chromis atrilobata

255 47.3 0.33 0.34 0.52

Halichoeres dispilus

65 12.1 0.33 0.09 0.21

Stegastes acapulcoensis

56 10.4 0.60 0.07 0.10

Haemulon steindachneri

40 7.4 0.27 0.05 0.18

Thalassoma lucasanum

23 4.3 0.13 0.03 0.10

Bodianus diplotaenia

19 3.5 0.60 0.03 0.03

Chaetodon humeralis

14 2.6 0.53 0.02 0.02

Diodon holocanthus

13 2.4 0.33 0.02 0.04

Johnrandallia nigrirostris

10 1.9 0.27 0.01 0.03

Epinephelus labriformis

9 1.7 0.40 0.01 0.02

Abudefduf troschelii

8 1.5 0.33 0.01 0.02

Microspathodon dorsalis

8 1.5 0.33 0.01 0.02

Holacanthus passer

4 0.7 0.20 <0.01 0.01

Stegastes flavilatus

4 0.7 0.13 <0.01 0.02

Lutjanus argentiventris

3 0.6 0.20 <0.01 0.01

Cephalopholis panamensis

2 0.4 0.13 <0.01 0.01

Halichoeres chierchiae

2 0.4 0.13 <0.01 0.01

Muraena lentiginosa

1 0.2 0.07 <0.01 0.01

Pomacanthus zonipectus

1 0.2 0.07 <0.01 0.01

Caranx caballus

1 02 0.07 <0.01 0.01

Alphestes immaculatus

1 0.2 0.07 <0.01 0.01

SARGASSUM BEDS

Halichoeres dispilus

64 77.1 0.83 0.21 0.18

Chaetodon humeralis

4 4.8 0.33 0.01 0.02

Haemulon steindachneri

4 4.8 0.17 0.01 0.03

Serranus psittacinus

3 3.6 0.33 0.01 0.02

Sphoeroides lobatus

2 2.4 0.33 <0.01 0.01

Stegastes flavilatus

2 2.4 0.17 <0.01 0.02

Urobatis halleri

1 1.2 0.17 <0.01 0.01

Diodon holocanthus

1 1.2 0.17 <0.01 0.01

Canthigaster punctatissima

1 1.2 0.17 <0.01 0.01

Balistes polylepis

1 1.2 0.17 <0.01 0.01

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high biological diversity and ocean-atmosphere

phenomena, and for its great national impor-

tance due to economic activities that take

place there (Cortés, 2014; Cortés, 2016). The

scientific information about species in marine

ecosystems in an area is critical to understand,

conserve and management the biodiversity in

an integral way (Margules & Pressey, 2000).

Although our records of invertebrates and

fish species in BSa and GSE are typical of the

biogeographic province of the Eastern Tropi-

cal Pacific (ETP) of Costa Rica (Alvarado,

Chacón-Monge, Solís-Marín, Pineda-Enríquez,

Caballero-Ochoa, Solano-Rivera & Romero-

Chaves, 2017; Alvarado & Chiriboga, 2008;

Alvarado & Fernández, 2005; Alvarado, Solís-

Marín & Ahearn, 2010; Alvarado et al., 2015;

Alvarado et al., 2018; Cordero-Umaña & San-

tidrián-Tomillo, 2020; Cortés, 2017; Dominici-

Arosemena, Brugnoli-Olivera, Cortés-Núñez,

Molina-Ureña & Quesada-Alpizar, 2005;

Murase, Angulo, Miyazaki, Bussing & López,

2014;), the distribution geographic record of

invertebrates and fishes observed in this study

contribute to the knowledge of marine biodi-

versity in conservation priority areas (North

Pacific of Costa Rica) and/or sites (marine sec-

tor of conservation areas) identified in the coun-

try (Alvarado et al., 2011; Cortés, 2012; Cortés,

2014; Cortés, 2017; SINAC-MINAET, 2008).

Moreover, it contributes to the research efforts

on the biodiversity of marine ecosystems of

the Pacific shores of Costa Rica (Cortés, 2012;

Cortés, 2016) that have received little (e.g.,

rocky reefs) (Cordero-Umaña & Santidrián-

Tomillo, 2020; Dominici-Arosemena et al.,

2005; Espinoza & Salas 2005) or scarce atten-

tion (e.g., Sargassum beds) compared with

others, e.g., coral reefs (Alvarado & Chiriboga,

2008; Alvarado et al., 2015; Alvarado et al.,

2018; Cortés, 1996/1997).

Our estimates of number of macroinver-

tebrates and conspicuous fish species would

be larger with more spatial and temporal rep-

lication. Because estimates are sample-design

dependent, comparisons to number of spe-

cies at other sites where different sampling

designs were used should be considered only

suggestive. At the rocky reefs, for mollusks

(gastropods and bivalves) and echinoderms

(sea stars, sea urchins, sea cucumber and brittle

stars) we recorded a lower number of species

(23) than reported in rocky reefs in the Gulf

of Papagayo (46 species) (Cordero-Umaña, &

Santidrián-Tomillo, 2020) and in coral reefs

(35 to 37 species) in the ETP of Costa Rica

(Alvarado et al., 2015; Alvarado et al., 2018).

On the other hand, the number of fish species

(33) is less than reported previously (46 to

81 species) by Cordero-Umaña & Santidrián-

Tomillo (2020), Dominici-Arosemena et al.

(2005) and Espinoza & Salas (2005) in rocky

reefs in the North Pacific of Costa Rica. Our

lower number of species maybe due to our

lower sampling compared to previous studies,

our results suggest that the number of macroin-

vertebrates and fish species in rocky reef could

be higher at BSa and GSE.

In the case of Sargassum beds, our study

is the first biodiversity report for this marine

ecosystem in the ETP of Costa Rica. With eight

macroinvertebrates (mollusks and echinoderms

groups) and ten fish species, the number of

species recorded are lower than reported in Sar-

gassum beds for these faunistic groups (44 to

73 species) from others latitudes, like the Gulf

of California in México (Foster, McConnico,

Lundsten, Wadsworth, Kimball, Brooks et al.,

2007; Suárez-Castillo, 2008; Suárez-Castillo,

2014; Suárez-Castillo, Riosmena-Rodríguez,

Hernández-Carmona, Méndez-Trejo, López-

Vivas, Sánchez-Ortiz et al., 2013). Our results

suggest that the number of invertebrates and

fish species could be higher in GSE, this too

possibly due lower sampling effort than used in

other studies, and because we visit the Sargas-

sum beds in February, before the algae reached

their maximum length (102.0 cm in April)

(Cortés et al., 2014). The mean length of S.

liebmanni thallus observed was 7.67 cm (Alvin

Suárez, personal observation). Sargassum spe-

cies are considered foundation species because

they harbor a vast diversity of species because

of the habitat produced by the algae itself (Fos-

ter et al., 2007; Suárez-Castillo, 2008; Suárez-

Castillo, 2014).

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The variation of nearshore flora and fauna

is often attributed to habitat structure and

seasonal changes in environmental conditions

(Aburto-Oropeza & Balart, 2001; Beukhof,

Dencker, Pecuchet & Lindegren, 2019; Domi-

nici-Arosemena & Wolff, 2006; Foster et al.,

2007; McCourt, 1984; Palacios & Zapata,

2014; Scrosati, 2001). However, without phys-

ical descriptors of marine ecosystems and

temporal replication and more quantitative,

seasonal records of species composition in

these marine ecosystems from this study, the

consistency of these temporal patterns and

their causes are largely unknow. The commu-

nities of invertebrate and fish of both marine

ecosystems have different species composition.

At the rocky reefs, the black sea urchin, D.

mexicanum, and slate pencil urchin, E. thouars-

sii, were the most abundant invertebrates (1

513 and 192 ind., respectively) and frequently

observed (100% and 93%, respectively). This

result suggest that our estimate is reasonable

for this marine ecosystem, where it has been

recognized that these two species are the most

dominant grazers in the ETP (Guzmán & Cor-

tés, 1993). It has been reported that these two

echinoids exert a strong influence on the com-

munity structure (Andrew, 1989; Glynn, Wel-

lington & Birkeland, 1979; Lawrence, 1975;

Underwood, 1992), so the values of mean

density of D. mexicanum (5.04 ind m

-2

) and

E. thouarsii (0.57 ind m

-2

) observed in this

study are important to consider the periodical

monitoring of community structure in rocky

reefs in BSa and GSE. The mean density value

recorded for D. mexicanum in this study are

higher than was has been previously recorded

in the northern Pacific region of Costa Rica

(0.20 to 2.19 ind m

-2

) (Alvarado, Cortés &

Reyes-Bonilla, 2012). The fish species compo-

sition observed in this study, high abundance

of C. atrilobata (255 inds.) and serranids,

pomacentrids, labrids and tetraodontids fishes

with highest number of fish species, is similar

to recorded for rocky reef in the North Pacific

of Costa Rica (Cordero-Umaña & Santidrián-

Tomillo, 2020; Dominici-Arosemena et al.,

2005; Espinoza & Salas, 2005), indicating that

the estimate for our survey sites is reasonable

for this type of community.

Comparisons of communities of inverte-

brate and fish species in Sargassum beds from

this study with the observed in other sites in

the northern Pacific region of Costa Rica can’t

be done, because our work represent the first

report of this type for this marine ecosystem

in the region. The composition of inverte-

brates and fish species recorded are similar

to observed in Sargassum beds in the Gulf of

California (Foster et al., 2007; Suárez-Castillo,

2008; Suárez-Castillo, 2014; Suárez-Castillo et

al., 2013) suggesting that our observations are

typical for this marine ecosystem. However,

it was possible to observe invertebrate spe-

cies (Vasum caestus, Astropyga pulvinata) that

have not been previously recorded in Sargas-

sum beds, and differences in the invertebrate

and fish species frequently observed and with

highest mean densities compared with Sargas-

sum beds in other sites (Suárez-Castillo, 2008;

Suárez-Castillo et al., 2013). Therefore, taxo-

nomic and community structure studies of the

fauna in Sargassum beds in the North Pacific of

Costa Rica are necessary.

The original approach of this research by

one of the authors (A. Suárez) was to monitor

the community of invertebrate and fish spe-

cies in Sargassum beds in the northern Pacific

region of Costa Rica, where has been reported

the absence of S. liebmannni since 2013, fol-

lowing a population outbreak of the black sea

urchin D. mexicanum (Cortés et al., 2014). The

substantial decrease or total disappearance of

these seaweed beds by multiple factors (sea-

sonal patterns related to temperature changes,

introduction of alien species, biotic interac-

tions, among others) has been reported in many

coastal areas around the world (Aburto-Oro-

peza, Sala, Paredes, Mendoza & Ballesteros,

2007; Britton-Simmons, 2004; Haraguchi &

Sekida, 2008; Rivera & Scrosati, 2006; Viejo,

1997; Xuan-Vy & Huu-Dai, 2011). The losses

of these foundation species and the shift from

algal dominated substrate to bare substrate,

as observed on rocky reefs in our survey

sites in BSa and GSE (Alvin Suárez, personal

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observation), may have implications in the

decline of marine biodiversity associated with

them. Therefore, more studies of marine biodi-

versity in Sargassum beds in the North Pacific

of Costa Rica are necessary.

Our study further evidences the poorly and

scarce knowledge of communities of inver-

tebrates and fish species in rocky reefs and

Sargassum beds in the North Pacific of Costa

Rica, respectively. Which require considerably

more survey and field work to locate such

marine ecosystems, better characterize their

marine biodiversity that occur in them and

better understand of the community structure

of fauna and the factors that determine them.

Information that is required to support man-

agement decisions that are backed by sound

scientific knowledge. These marine ecosystems

deserve special consideration in efforts to con-

serve biodiversity.

Ethical statement: authors declare that

they all agree with this publication and made

significant contributions; that there is no con-

flict of interest of any kind; and that we

followed all pertinent ethical and legal proce-

dures and requirements. All financial sources

are fully and clearly stated in the acknowled-

gements section. A signed document has been

filed in the journal archives.

ACKNOWLEDGMENTS

We dedicate this paper to the memory

of Rafael Riosmena-Rodríguez (1966-2016),

mentor, colleague and friend. The study was

possible by CONACYT (México) who granted

a Research Fellowship (Beca Mixta No. 34677)

to Alvin Suárez during his Doctorate (Scholar-

ship number 202159) for a research visit to the

Centro de Investigación en Ciencias del Mar

y Limnología, Universidad de Costa Rica. We

thank the editor and two anonymous reviewers

for their valuable comments and suggestions,

which greatly improved the paper. A special

thank to Eleazar Ruiz and Davis Morera for

their assistance in the field, and Imelda Amador

for support in the elaboration of the map.

RESUMEN

Listado de invertebrados y peces conspicuos

en arrecifes rocosos y mantos de Sargassum en

el Pacífico Norte de Costa Rica

Introducción: La biodiversidad marina es bien conocida

en algunas áreas y para algunos ecosistemas marinos de la

costa Pacífica de Costa Rica. El sector marino del Área de

Conservación Guanacaste, en el Pacífico norte del país, es

un área prioritaria para la conservación de la biodiversidad

marina y costera.

Objetivo: Nuestro objetivo fue el de incrementar el cono-

cimiento de la biodiversidad marina en un sitio prioritario

de conservación de Costa Rica y en ecosistemas marinos

poco o nulamente estudiados, como son los arrecifes roco-

sos y mantos de Sargassum, respectivamente.

Métodos: El estudio se realizó en Bahía Salinas y Golfo de

Santa Elena en el 2013. En enero, cuatro arrecifes rocosos

fueron visitados; mientras que en febrero tres arrecifes

rocosos y un manto de Sargassum fueron visitados. Identi-

ficamos las especies de invertebrados y peces conspicuos

a través métodos de censos visuales submarinos (buceos

cronometrados y transectos de banda). La densidad pro-

medio de cada especie se estimó mediante el método de

transectos de banda.

Resultados: Se registró un total de 25 invertebrados y

34 especies de peces. Encontramos un mayor número de

especies de invertebrados (23) y peces (33) en los arrecifes

rocosos, pero pocas especies en el manto de Sargassum

(ocho especies de invertebrados y diez de peces). La com-

posición de especies difirió entre ecosistemas marinos.

Conclusiones: Nuestro estudio evidencia el limitado cono-

cimiento de las comunidades de invertebrados y peces en

estos ecosistemas marinos en el Pacífico Norte de Costa

Rica. Considerablemente más encuestas y trabajo de campo

son requeridos para apoyar decisiones de manejo que sean

respaldados por conocimientos científicos solidos.

Palabras clave: equinodermos, moluscos, peces, sitio prio-

ritario de conservación, Área de Conservación Guanacaste.

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