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Revista de Biología Tropical, ISSN electrónico: 2215-2075, Vol. 69(S1): 219-253, March 2021 (Published Mar. 30, 2021)

Echinoderms (Echinodermata) from the Central Mexican Pacific

Rebeca Granja-Fernández*

Brenda Maya-Alvarado

Amílcar L. Cupul-Magaña

A. Paola Rodríguez-Troncoso

Francisco A. Solís-Marín

Rosa C. Sotelo-Casas

1. Laboratorio de Ecología Marina. Centro Universitario de la Costa, Universidad de Guadalajara. Av. Universidad,

Delegación Ixtapa. Puerto Vallarta, México; beckygranja@gmail.com (*Correspondence), amilcar.cupul@gmail.com,

pao.rodriguezt@gmail.com

2. Programa de Maestría en Ciencias en Biosistemática y Manejo de Recursos Naturales y Agrícolas, Centro

Universitario de Ciencias Biológicas y Agropecuarias, Universidad de Guadalajara, Av. Ing. Ramón Padilla Sánchez,

Zapopan, Jalisco, México; brenda.maya9471@alumnos.udg.mx

3. Colección Nacional de Equinodermos “Ma. E. Caso Muñoz”, Laboratorio de Sistemática y Ecología de Equinodermos.

Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, Ciudad de México, México;

fasolis@cmarl.unam.mx

4. Unidad Multidisciplinaria de Docencia e Investigación Sisal Facultad de Ciencias, Universidad Nacional Autónoma

de México. Puerto de Abrigo S/N, Sisal, Yucatán, México; rosacarmensotelocasas@gmail.com

Received 30-XVI-2020. Corrected 18-VIII-2020. Accepted 19-X-2020.

ABSTRACT

Introduction: The echinoderms from the Central Mexican Pacific are of high scientific interest and, prior to this

present work, there was a lack of basic information that included incomplete checklists with inconsistencies in

systematics and spatial distribution. Objective: To provide a historical review, and an updated checklist with a

more complete richness of echinoderms for each state and island of the region. Methods: A checklist was elabo-

rated based on an exhaustive literature search of the Echinodermata, and was complemented with taxonomical

revisions of Ophiuroidea scientific collections. All the geographical coordinates of the records were validated.

Results: The region harbors 187 species of Echinodermata: three Crinoidea, 35 Asteroidea, 67 Ophiuroidea, 32

Echinoidea, and 50 Holothuroidea. We detected 52 records in the literature that must be considered as invalid

and five as doubtful. We provide 16 new records of Ophiuroidea from different states and islands; of them, four

are new records for the region. Jalisco presented the highest number of species (84), followed by the coast of

Nayarit (74), Michoacán (63), and Colima (55); among the islands, Revillagigedo showed the major number

of species (85) followed by Marías (81), Marietas (48), and Isabel (44). Conclusions: The numbers of species

known in the region are mostly related to both sampling effort and environmental characteristics that promote

high biodiversity. The Central Mexican Pacific is an oceanographic region with mixed conditions from the

North and South of the Mexican Pacific, and therefore, with a biogeographical importance reflected in its spe-

cies richness.

Key words: Echinodermata; biodiversity; checklist; historical review; new records.

Granja-Fernández, R., Maya-Alvarado, B., Cupul-Magaña,

A.L., Rodríguez-Troncoso, A.P., Solís-Marín,

F.A., & Sotelo-Casas, R.C. (2021). Echinoderms

(Echinodermata) from the Central Mexican Pacific.

Revista de Biología Tropical, 69(S1), 219-253. DOI

10.15517/rbt.v69iSuppl.1.46356

DOI 10.15517/rbt.v69iSuppl.1.46356

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Revista de Biología Tropical, ISSN electrónico: 2215-2075 Vol. 69(S1): 219-253, March 2021 (Published Mar. 30, 2021)

The Central Mexican Pacific (CMP) region

comprises the states of Nayarit, Jalisco, Colima

and Michoacán as well as their coastal (Isabel

and Marietas) and oceanic islands (Marías and

Revillagigedo). The region is characterized by

numerous geographic features such as bays,

archipelagos and islands, as well as a hetero-

genic bottom that combines hard and soft com-

ponents. The CMP is located in the Tropical

Eastern Pacific Province nearby to the Warm

Temperate Northeast Pacific Province (Spald-

ing et al., 2007) and for this reason, it shelters

northern and southern fauna of the Mexican

Pacific (Arriaga-Cabrera et al., 1998). Due to

its relevant flora and fauna, most of the islands

of the CMP are catalogued as Natural Protected

Areas (CONANP, 2005; CONANP, 2007a;

CONANP, 2007b; CONANP, 2019).

The study regarding the Echinodermata

in the CMP began in 1840 when Gray (1840)

reported the asteroids Astropecten regalis and

Luidia columbia in Nayarit. Until 1891, efforts

in the collection of echinoderms mostly from

deep-waters were performed by foreign expe-

ditions such as Albatross and Challenger. As

a result, many new records and new spe-

cies of Crinoidea (Hartlaub, 1895), Asteroi-

dea (Sladen, 1889), Ophiuroidea (Lütken &

Mortensen, 1899), Echinoidea (Agassiz, 1872;

Agassiz,1898), and Holothuroidea (Ludwig,

1894) were described.

The 20

century was very productive in

terms of the number of published works (75).

During this period, numerous foreign expedi-

tions (Albatross, Velero, and Zaca) collected

samples from shallow and deep-waters along

the region resulting in new records, new spe-

cies and extensive taxonomic monographs (e.g.

Ludwig, 1905; Fisher, 1928a; Fisher, 1928b;

Deichmann, 1937; Ziesenhenne, 1937; Ziesen-

henne, 1940; Mortensen, 1948a; Mortensen,

1948b). The first national work regarding echi-

noderms from the CMP was the master´s

thesis of Caso (1943) who worked with the

Asteroidea from México. Caso published dis-

sertations and numerous manuscripts regarding

taxonomy (Caso, 1948a; Caso, 1951; Caso,

1961; Caso, 1962a; Caso, 1965; Caso, 1966;

Caso, 1974a; Caso, 1974b; Caso, 1978b; Caso,

1980; Caso,1983b), including descriptions of

new species (Caso, 1954; Caso, 1983a; Caso,

1986a), and general accounts on the biodiversity

of the CMP (e.g. Caso, 1948b; Caso, 1962b).

Their findings helped to better understand the

echinoderm fauna from shallow-waters of the

region and inspired other Mexican researchers

to work with echinoderms from Revillagigedo

Islands (Villalobos, 1992; Chacón-Torres et al.,

1993; Bautista-Romero, Reyes-Bonilla, Lluch-

Cota, & Lluch-Cota, 1994), Marietas Islands

(Nepote-González, 1998), Colima (Espino-

Barr, Cibrián-Rodríguez, & García-Boa, 1996)

and even, with species collected in several

areas from the CMP that resulted as new taxa

(Solís-Marín & Laguarda-Figueras, 1999). At

the end of the 20

century, the first ecological

studies in the CMP were published (Holguin-

Quiñones, 1994; Reyes-Bonilla, 1995; Espino-

Barr et al., 1996).

The first two decades of the 21

cen-

tury have produced a vast number of works

(55) regarding topics such as new distribution

records (Frontana-Uribe, Hernández-Alcánta-

ra, & Solís-Weiss, 2000; Sotelo-Casas, Cupul-

Magaña, Solís-Marín, & Rodríguez-Troncoso,

2015; Valdés de Anda, Hendrickx, & Solís-

Marín, 2018), description of new species

(Honey-Escandón, Solís-Marín, & Laguarda-

Figueras, 2011; Solís-Marín, Arriaga-Ochoa,

Galván-Villa, & Laguarda-Figueras, 2018),

ecology (Holguin-Quiñones & Michel-Morfin,

2006; López-Uriarte et al., 2009; Hermosillo-

Nuñez et al., 2015; Herrero-Pérezrul, Ramírez-

Ortiz, Rosales-Estrada, & Reyes-Bonilla, 2015;

Hermosillo-Nuñez, Rodríguez-Zaragoza, Ortiz,

Calderon-Aguilera, & Cupul-Magaña, 2016;

Sotelo-Casas, Cupul-Magaña, Rodríguez-Zara-

goza, Solís-Marín, & Rodríguez-Troncoso,

2018), and reproduction (Sotelo-Casas, Cupul-

Magaña, Solís-Marín, & Rodríguez-Troncoso,

2016). It is important to emphasize that most

of these studies are focused in the compilation

of species of echinoderms inhabiting in dif-

ferent areas of the CMP (Chávez-Dagostino,

Nepote-González, Solís-Marín, & Medina-

Rosas, 2000; Fuentes-Farías, Villarroel-Melo,

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& Solís-Marín, 2005; Honey-Escandón, Solís-

Marín, & Laguarda-Figueras, 2008; Ríos-Jara,

Galván-Villa, & Solís-Marín, 2008a; Ríos-Jara

et al., 2008b; Santos-Beltrán & Salazar-Silva,

2011; Ríos-Jara et al., 2013; Rodríguez-Tron-

coso et al., 2013; Solís-Marín et al., 2013a;

Granja-Fernández, Herrero-Pérezrul, López-

Pérez, Hernández-Morales, & Rangel-Solís,

2015a; Solís-Marín, Laguarda-Figueras, &

Durán-González, 2016b; Granja-Fernández et

al., 2017; Padilla-Pérez, Rodríguez-Troncoso,

Sotelo-Casas, & Cupul-Magaña, 2017; Ríos-

Jara, Juárez-Carrillo, & Galván-Villa, 2017;

Nava-Bravo et al., 2019).

Despite the above, the records have shown

many inconsistencies such as: 1) the names

of some species have been mentioned (writ-

ten or registered) incorrectly across time (e.g.

misspelling), 2) many of the recorded species

for the CMP are invalid, and 3) the generated

checklists are not complete because they are

based only in museum records or material col-

lected mostly in shallow-waters. Thus, the aim

of the present manuscript is to provide an accu-

rate and updated checklist of the valid names

and synonyms of the Echinodermata from the

CMP according to literature, including new dis-

tribution records of Ophiuroidea. Moreover, we

provide a historical review of the study of each

class of Echinodermata, as well as an updated

number of species for each state and island in

the region.

MATERIALS AND METHODS

An exhaustive search of published lit-

erature for the CMP regarding the classes Cri-

noidea, Asteroidea, Ophiuroidea, Echinoidea,

and Holothuroidea was performed (records

dating from 1840 to 2019). Moreover, we

visited reference collections in order to taxo-

nomically identify specimens of Ophiuroidea:

ICML-UNAM: Colección de Equinodermos

“Dra. Ma. Elena Caso”, Instituto de Ciencias

del Mar y Limnología, Universidad Nacional

Autónoma de México, Ciudad de México,

México; LACM: Natural History Museum, Los

Angeles County Museum, Los Angeles, United

States of America (USA); and, MCZ: Museum

of Comparative Zoology, Harvard University,

Cambridge, USA.

When available, the geographical coordi-

nates of the provided localities in the literature

were validated using ArcMap 10.4.1 software,

to corroborate distribution within the CMP

maritime zone. Records were assigned to any

of the following areas: coastlines of the states

of Colima (COL), Jalisco (JAL), Michoacán

(MICH), and Nayarit (NAY), and the islands

Isabel (ISA), Marías (MAR), Marietas (MAT),

and Revillagigedo (REV).

With this information, a checklist of the

valid names and synonyms of Echinoderma-

ta from the CMP was constructed. The list

includes those synonyms used exclusively in

this area, yet some species may have larger

synonym lists for other geographic areas. Sys-

tematics arrangements and valid names agree

with Kroh and Mooi (2020), Mah (2020),

Messing (2020), Stöhr, O´Hara, and Thuy

(2020), and WoRMS (2020).

RESULTS AND DISCUSSION

Historical review

Crinoidea: Six references documented the

Crinoidea from the CMP; one during the 19

one in the 20

, and four in the 21

century.

Among the Echinodermata, Crinoidea repre-

sents the less studied class of the CMP. The

study of the Crinoidea from the CMP began

in 1895 when Hartlaub (1895) recorded the

species Florometra tanneri (as Antedon rhom-

boidea) for MAR at 1 236 m depth. Several

years later, using a submersible, Roux (2004)

collected one specimen of Hyocrinus foelli in

COL at 3 030 m depth. Finally, during deep-

water dredging by the TALUD cruises aboard

the R/V “El Puma” of the Universidad Nacio-

nal Autónoma de México, Florometra serratis-

sima was collected in two stations located in

COL between 1 040 and 1 106 m depth (Valdés

de Anda et al., 2018).

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Asteroidea: 57 references addressed the

Asteroidea distributed along the CMP. During

the 19

century only two works were pub-

lished, meanwhile the 20

and 21

centuries

presented similar numbers of works, 28 and

27, respectively. The 19

century represents

the era with the lowest addition of Asteroidea.

The first reported Asteroidea for the CMP were

Luidia columbia and Astropecten regalis, both

collected in San Blas, NAY (Gray, 1840).

The 20

century was the period with the

highest addition of Asteroidea to the CMP (25

species). Ludwig (1905) described Ampheraster

marianus, Henricia gracilis, and Sarkaster

validus (= Lophaster furcilliger) for MAR and

other areas of the Eastern Pacific. According

to samples from the Albatross, Fisher (1911)

recorded Hippasteria lepidonotus and Henricia

clarki for deep-waters from MAR and REV,

respectively. Clark (1916) provided the original

descriptions of Saraster insignis (= Nearchast-

er (Nearchaster) aciculosus) and Cyllaster

seminuda (= Henricia seminudus) from Clarion

Island, REV. In 1936, the Templeton Crocker

Expedition collected echinoderms during the

expedition of the Zaca along the north of

México and as a result, Ziesenhenne (1937)

mentioned Astropecten armatus, Acanthaster

ellisii (= Acanthaster planci), Linckia columbi-

ae, Pauliella aenigma (= Paulia horrida), and

Sclerasterias heteropaes for shallow-waters

(up to 100 m depth) from Clarion Island. Dur-

ing 1937-1938, the Zaca continued performing

collections of shallow-water echinoderms, and

Clark (1940) reported Luidia bellonae (Man-

zanillo, COL and Chamela Bay, JAL), Luidia

foliolata, Pharia pyramidata, and Phataria

unifascialis (Tenacatita Bay, JAL). Later, Caso

(1943) reported the species Heliaster micro-

brachius and Luidia superba for Manzanillo,

COL, as well as Nidorellia armata for the same

locality and MAR. In later years, the addition

of species for the CMP was sporadic, with

mentions of just one species per work: Patiria

miniata (Socorro Island, REV; Adem et al.,

1960), Astropecten californicus (Manzanillo,

COL; Caso, 1961), Mithrodia bradleyi (REV;

Caso, 1962b), Pentaceraster cumingi (Socorro

and Clarion Islands, REV; Hertlein, 1963), and

Asteropsis carinifera (REV; Maluf, 1991).

During the 21

century, only eight new

records for the CMP have been documented.

Fuentes-Farías et al. (2005) provided a list

of echinoderms for MICH including the new

records of Luidia tessellata and Heliaster heli-

anthus. Two years later, the Mexican govern-

ment decreed the MAR as a Biosphere Reserve,

publishing a management and conservation

program (CONANP, 2007a), which provided

lists of the marine and terrestrial flora and

fauna, and included the record of Echinas-

ter (Othilia) tenuispina for the area. Honey-

Escandón et al. (2008) and Solís-Marín et al.

(2013b) provided the most-complete checklist

of echinoderms for the Mexican Pacific and

the Eastern Pacific, respectively, including

the new records of Heliaster polybrachius

(JAL), Mediaster transfuga and Meridiastra

modesta (REV), and Narcissia gracilis (NAY).

Recently, Nava-Bravo et al. (2019) included

the record of Astropecten verrilli for MICH.

Ophiuroidea: 59 references have been

published regarding this class. The 19

cen-

tury was the least productive since only one

work was published. During the 20

and 21

centuries, similar numbers of manuscripts were

published, 30 and 28, respectively. In 1891,

the Albatross conducted an expedition along

the Eastern Pacific, and as a result, Lütken and

Mortensen (1899) described the first species

and subspecies of ophiuroids from deep-waters

of the CMP (MAR): Amphiura serpentina,

Astrodia excavata, Dougaloplus notacanthus,

Ophiacantha costata, Ophiacantha hirta,

Ophiacantha moniliformis, Ophiernus adsper-

sus annectens, Ophiomitra granifera, Ophi-

omitra partita, Ophiosphalma variabile, and

Ophiura (Ophiura) scutellata.

During the 20

century a total of 30

species were added to the list of ophiuroids

of the CMP, representing the highest num-

ber, in comparison to the other two centuries

accounted for in this review. Döderlein (1911)

conducted a revision of the world’s Euryalida,

reporting Asteroschema sublaeve for MAR.

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With specimens collected in shallow-waters of

Clarion Island, Ziesenhenne (1937) described

Ophiacantha pyriformis, and reported for the

first time in the CMP, the presence of Ophi-

actis savignyi, Ophiocoma aethiops, Ophio-

derma variegatum, Ophionereis annulata, and

Ophiothrix galapagensis. Later, Clark (1940)

found the conspicuous Diopederma daniana,

Ophiolepis variegata, and Ophiothrix (Ophio-

thrix) spiculata in JAL and COL. In the same

year, Ziesenhenne (1940) published the results

obtained by the cruise Velero III performed in

1938, and described Amphichondrius laevis,

Amphiodia sculptilis, Ophiolepis plateia, and

Ophiophragmus lonchophorus, all of them col-

lected in Tenacatita Bay, JAL. Later, the addi-

tions of new records of ophiuroids per work

were reduced: Ophiocomella alexandri (REV;

Steinbeck & Ricketts, 1941), Ophioderma teres

and Ophiolepis pacifica (Puerto Vallarta, JAL;

Caso, 1951), Ophioderma panamense (REV;

Ziesenhenne, 1955), Ophiomyxa panamensis

(REV; Hertlein, 1963), Ophiostigma tenue (ISA;

Clark, 1970), Ophiomusa lymani, Ophiothela

mirabilis, and Ophiuroglypha irrorata irrorata

(JAL and REV; Luke, 1982), Amphichondri-

us unamexici (= Amphichondrius granulatus;

NAY; Caso, 1983a), Amphiura seminuda and

Ophiophragmus marginatus (JAL and NAY;

Caso, 1986b), Amphiura (Amphiura) assimilis

and Gymnophiura mollis (MAR; Maluf, 1988),

Ophiactis simplex (REV; Maluf, 1991), and

Ophiuroconis bispinosa (REV; Hendler, 1996).

The addition of new records of ophiuroids

(22) during the 21

century has been important.

Frontana-Uribe et al. (2000) collected speci-

mens of Ophiocomella schmitti associated with

coral rubble in localities from Socorro Island

(REV). Fuentes-Farías et al. (2005) included

the record of Ophiothrix (Ophiothrix) rudis

for MICH; in the same year, Maluf and Brusca

(2005) published a checklist of the echino-

derms from the Gulf of California, including

the general distribution of the species, and the

new records of Amphipholis elevata, Astroca-

neum spinosum, Microphiopholis puntarenae,

Ophiocomella sexradia, Ophiocnida hispida,

Ophiolepis crassa, Ophiopholis bakeri, and

Ophiophthalmus diplasia (MAR and REV).

The genus Ophiophthalmus is an invalid junior

synonym and a new assignment is needed

(Hendler, 1996), but in the present work we

kept this temporary genus in order to avoid sys-

tematic confusions. Later, Honey-Escandón et

al. (2008) reported 33 species of ophiuroids for

the CMP, of which, the amphiurids Amphiodia

occidentalis, Amphiodia platyspina, Amphiodia

violacea, Amphiodia (Amphispina) digitata,

Amphiodia (Amphispina) urtica, Amphipho-

lis pugetana, and Microphiopholis platydisca,

represented new records in several states and

islands from the CMP. In 2011, Astrodictyum

panamense (Ayala-Bocos, Reyes-Bonilla, Her-

rero-Pérezrul, Walther-Mendoza, & Castañeda-

Fernández de Lara, 2011) and Hemipholis

gracilis (Hendler, 2011) were reported in REV

and JAL, respectively. García-Hernández et

al. (2014) compared the ecological diversity

and species composition of benthic macroin-

vertebrates in rocky reefs of JAL and MAT,

finding new records for Amphipholis squamata

and Ophionereis perplexa. Finally, Granja-

Fernández et al. (2017) provided a checklist

of ophiuroids from the CMP, adding the new

record of Ophiophragmus papillatus.

Echinoidea: 73 references documented

the presence of the Echinoidea from the CMP,

representing the major number of works among

all the classes. The 19

century is represented

only by two publications, the 20

century for

39, and the 21

century for 32 works. In the

century all the work regarding Echinoidea

was published by Alexander Agassiz whom

documented for the first time in the CMP

the species Eucidaris thouarsii and Echino-

metra vanbrunti, both collected in Manza-

nillo, COL (Agassiz, 1872), and who described

Brisaster townsendi and Plexechinus cinctus

for stations located in deep-waters of MAR

(Agassiz, 1898).

The 20

century was by far the most pro-

ductive in terms of new addition of echinoids

(27) in the CMP. Clark described Hesperocidar-

is perplexa and Echinometra insularis (Clark,

1907; Clark, 1912), and reported Echinometra

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oblonga and Tripneustes gratilla from REV

(Clark, 1912). With specimens collected by the

Zaca, Ziesenhenne (1937) reported Clypeaster

europacificus, Diadema mexicanum, and Tri-

pneustes depressus in shallow-waters of Socor-

ro and Clarion, REV. In 1938, was published

an important work regarding fossil and living

Echinoidea from the west American Cenozoic,

in which were mentioned the new records of

Astropyga pulvinata (JAL) and Toxopneustes

roseus (ISA) (Grant & Hertlein, 1938). Poste-

riorly, Clark (1940) added Clypeaster ochrus

and Clypeaster speciosus from ISA and NAY,

respectively, and Caso (1948a) included Arba-

cia stellata to NAY. Clark (1948) reported the

highest number of new records of echinoids

for the CMP (Brissopsis pacifica, Clypeaster

rotundus, Encope micropora, Encope micro-

pora insularis, Lovenia cordiformis, Meoma

ventricosa grandis, and Rhyncholampas pacifi-

cus), based on the collections of the Velero III

along the Eastern Pacific during the years 1931-

1941; E. micropora insularis was described in

this work and its holotype corresponds to

Socorro Island, REV. The records in posterior

works were sporadic: Hesperocidaris asteris-

cus (JAL and REV; Caso, 1961), Heterocentro-

tus mamillatus (REV; Chan, 1974), Dendraster

excentricus and Lanthonia longifissa (ISA and

COL, respectively; Caso, 1980), Agassizia

scrobiculata (NAY; Caso, 1983b), and Mellita

notabilis (MICH; Harold & Telford, 1990).

Despite the number of published works

during the 21

century is similar to the previ-

ous century, only three new records for the

area have been documented during the pres-

ent era. These records correspond to species

observed during field work and documented in

the management and conservation programs of

the National Park ISA (Centrostephanus coro-

natus; CONANP, 2005) and the Reserve of the

Biosphere MAR (Brissus obesus; CONANP,

2007a). Finally, Ríos-Jara et al. (2008a) per-

formed collections of echinoderms in ISA at

different depths finding Brissus latecarinatus

in shallow-waters (< 20 m depth).

Holothuroidea: 53 references mention

holothuroids from the CMP. During the 19

century only one work was published, 19 dur-

ing the 20

century, and 33 during the 21

century. In the 19

century only one work was

published and therefore, the number of records

was low. Despite, Laetmophasma fecundum

(= Pannychia moseleyi), Psolus pauper, and

Ypsilothuria bitentaculata were first mentioned

in Ludwig (1893), it was until 1864 when Lud-

wig (1894) provided an extended description

of the species, reporting them in deep waters

from MAR; moreover, Molpadia musculus was

recorded in this work too, for the same locality.

The 20

century was the most produc-

tive in terms of the addition of new records

(28). In 1926, the Templeton Crocker expedi-

tion aboard the Zaca performed dredges in

order to obtain echinoderms along the Eastern

Pacific, and collected Holothuria (Cystipus)

inhabilis and Holothuria (Platyperona) dif-

ficilis in REV (Deichmann, 1937). From 1932

to 1938, the Allan Hancock Expedition aboard

the Velero III collected several specimens of

holothuroids and Deichmann (1941) found

that Thyone bidentata and Thyone parafu-

sus, both discovered in Tenacatita, JAL corre-

sponded to new species, and that Leptopentacta

panamica, Neothyone gibber, and Pentamera

chierchiae were new records of species for the

CMP (JAL and REV). Caso (1954) collected

for the first time Holothuria (Selenkothuria)

lubrica in Puerto Vallarta, JAL and found a

similar species which was designated as a

new species: Holothuria (Selenkothuria) por-

tovallartensis. The biggest contribution of new

records of holothuroids for the CMP (Holo-

thuria (Halodeima) kefersteinii, Holothuria

(Lessonothuria) pardalis, Holothuria (Mer-

tensiothuria) hilla, Holothuria (Mertensiothu-

ria) leucospilota, Holothuria (Semperothuria)

imitans, Holothuria (Semperothuria) languens,

Holothuria (Theelothuria) paraprinceps, Holo-

thuria (Thymiosycia) arenicola, Holothuria

(Thymiosycia) impatiens, and Isostichopus fus-

cus) was made by Deichmann (1958) which

correspond to a continuation and expansion

of the material collected by the Velero III and

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IV in JAL, as well as in ISA, MAR and REV.

The new records per work during the rest of

this century were reduced: Holothuria (Halo-

deima) inornata (Caso, 1961), Leptosynapta

albicans (Caso, 1962b), Abyssocucumis abys-

sorum (Luke, 1982), Holothuria (Stauropora)

fuscocinerea (Hickman, 1998), all of them

collected in REV and NAY. Nepote-González

(1998) conducted a research about the diver-

sity of holothuroids of MAT, finding the new

records of Chiridota rigida, Holothuria (Cys-

tipus) rigida, Labidodemas americanum, and

Pseudocnus californicus. Later, Solís-Marín

and Laguarda-Figueras (1999) performed col-

lections in several localities of the CMP (COL,

NAY, MAT) and described a new species,

Cucumaria flamma.

The addition of new records of holothuroids

to the CMP has been important during the 21

century. As part of the first program of manage-

ment and conservation of REV National Park,

CONANP (2004) documented Euapta godef-

froyi. Fuentes-Farías et al. (2005) and Honey-

Escandón et al. (2008) provided one of the most

complete checklists of echinoderms in México

and added the records of Neothyone gibbosa

and Labidodemas maccullochi for MICH, and

Euthyonidiella zacae, Holothuria (Halodeima)

atra, Holothuria (Mertensiothuria) viridiau-

rantia (= Holothuria (Mertensiothuria) hilla),

Lissothuria ornata, Neocucumis veleronis, and

Trachythyone peruana for several areas of the

Mexican Pacific. Holothuria (Mertensiothuria)

viridiaurantia is a recently described species

which was previously assigned to H. (Mer-

tensiothuria) hilla (Borrero-Pérez & Vanegas-

González, 2019). According to a preliminary

revision of the material (unpublished data)

and to Borrero-Pérez and Vanegas-González

(2019), in the CMP, H. (Mertensiothuria)

viridiaurantia inhabits in NAY, JAL, MICH

and MAT. On the other hand, based on a

taxonomical review of the material, we con-

firm that H. (Mertensiothuria) hilla inhabits

in REV, but a taxonomic confirmation of the

material from ISA and MAR is required (F.A.

Solís-Marín, personal communication, 01 April

2020). Honey-Escandón et al. (2011) described

Holothuria (Selenkothuria) carere, collected

in shallow-waters of MICH and other states

of the Mexican Pacific. Santos-Beltrán and

Salazar-Silva (2011) provided images and an

identification key of the holothuroids from

Bahía de Banderas, NAY adding the record of

Afrocucumis ovulum. Ríos-Jara et al. (2013)

described the taxonomic composition of the

echinoderms from shallow-waters in several

localities of Bahía Chamela, JAL, adding Neo-

pentamera anexigua, Pachythyone lugubris,

and Pachythyone pseudolugubris to the CMP,

and Solís-Marín, Arriaga-Ochoa, Galván-Vil-

la, and Laguarda-Figueras (2018) described

Lissothuria imbricata after the collection of

material from the same area. In recent years,

Nava-Bravo et al. (2019) elaborated the most

complete checklist of echinoderms from MICH

adding the records of Chiridota aponocrita,

Epitomapta tabogae, and Thyonella mexicana.

Checklist: The present updated checklist

includes 187 species of Echinodermata (three

Crinoidea, 35 Asteroidea, 67 Ophiuroidea, 32

Echinoidea, and 50 Holothuroidea; Table 1)

resulted from the revision of 137 literature ref-

erences (3 445 records) and the taxonomic revi-

sion of Ophiuroidea deposited at museum

collections. Additionally, we detected 57 records

in the literature (two Crinoidea, 21 Asteroidea,

14 Ophiuroidea, 15 Echinoidea, and five Holo-

thuroidea) considered as invalid and which

were removed from the updated checklist.

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TABLE 1

Checklist of valid names (in bold) and synonyms of the Echinodermata from the Central Mexican Pacific based

on literature review and museum records. All taxa are listed exactly as originally registered in the literature.

Numbers refers to literature where taxa were listed, see References section. * = New distribution records

Phylum Echinodermata Bruguière, 1791

Class Crinoidea Miller, 1821

Order Comatulida A.H. Clark, 1908

Family Antedonidae Norman, 1865

Genus Florometra A.H. Clark, 1913

Florometra serratissima (A.H. Clark, 1907) COL

132

Florometra tanneri (Hartlaub, 1895) MAR

90, 92

Antedon rhomboidea Hartlaub, 1895

Orden Hyocrinida Rasmussen, 1978

Familia Hyocrinidae Carpenter, 1884

Genus Hyocrinus Thomson, 1876

Hyocrinus foelli Roux & Pawson, 1999 COL

81, 114

Class Asteroidea de Blainville, 1830

Order Paxillosida Perrier, 1884

Family Luidiidae Sladen, 1889

Genus Luidia Forbes, 1839

Luidia bellonae Lütken, 1864 JAL, COL, MICH, REV

13, 39, 104

Luidia bellona Lütken, 1864

7, 92

Luidia (Alternaster) bellona Lütken, 1864

Luidia columbia (Gray, 1840) NAY, COL, MICH, REV

13, 15, 92, 104, 126, 135

Luidia brevispina Lütken, 1871

8, 13

Luidia columbiae (Gray, 1840)

7, 117

Luidia (Petalaster) columbia (Gray, 1840)

58, 81, 91

Petalaster Columbia Gray, 1840

Luidia foliolata Grube, 1866 JAL

Luidia superba A.H. Clark, 1917 COL, MICH

8, 10, 13, 30, 124

Luidia (Alternaster) superba A.H. Clark, 1917

58, 81, 91

Luidia tessellata Lütken, 1859 NAY, MICH, ISA

104

Luidia (Petalaster) tessellata Lütken, 1859

58, 81, 109, 110

Family Astropectinidae Gray, 1840

Genus Astropecten Gray, 1840

Astropecten armatus Gray, 1840 NAY, COL, MICH, REV

7, 8, 13, 15, 28, 30, 39 58, 81, 92, 104, 124, 135

Astropecten regalis Gray, 1840 NAY, COL, MICH

8, 13, 58, 66, 81, 104, 117, 124

Astropecten californicus Fisher, 1906 NAY, COL

13, 81, 124

Astropecten verrilli de Loriol, 1899 MICH

104

Genus Psilaster Sladen, 1885

Psilaster armatus Ludwig, 1905 MAR

86, 90, 92

Order Valvatida Perrier, 1884

Family Acanthasteridae Sladen, 1889

Genus Acanthaster Gervais, 1841

Acanthaster planci (Linnaeus, 1758) NAY, COL, MAR, ISA, MAT, REV

7, 32, 33, 41, 44, 45, 81, 107, 108, 120, 124

Acantaster planci (Linnaeus, 1758)

126

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Acantaster plansii (Linnaeus, 1758)

Acanthaster elissi (Gray, 1840)

134

Acanthaster elisii pseudoplanci Caso, 1962

Acanthaster ellisi (Gray, 1840)

43, 45

Acanthaster ellisii (Gray, 1840)

14, 15, 19, 41, 42, 72, 81, 92, 109, 110, 124, 135

Acanthaster ellisii pseudoplanci Caso, 1962

14, 15, 41, 69

Acanthaster pseudoplanci Caso, 1962

Family Asterinidae Gray, 1840

Genus Meridiastra O’Loughlin, 2002

Meridiastra modesta (Verrill, 1867) REV

81, 124

Asterina agustincasoi Caso, 1977

Genus Patiria Gray, 1840

Patiria miniata (Brandt, 1835) REV

120

Asterina miniata (Brandt, 1835)

1, 7, 15, 41, 45, 81

Patiria minata (Brandt, 1835)

126

Family Asterodiscididae Rowe, 1977

Genus Paulia Gray, 1840

Paulia horrida Gray, 1840 REV

41, 45, 91, 92, 126

Pauliella aenigma Ludwig, 1905

7, 13, 15, 77, 135

Family Asteropseidae Hotchkiss & A.M. Clark, 1976

Genus Asteropsis Müller & Troschel, 1840

Asteropsis carinifera (Lamarck, 1816) JAL, ISA, REV

41, 45, 91, 92, 126

Asteropsis spinosa (Gray, 1840)

Gymnasteria spinosa Gray, 1840

109, 110

Family Goniasteridae Forbes, 1841

Genus Hippasteria Gray, 1840

Hippasteria lepidonotus (Fisher, 1905) MAR

Cryptopeltaster lepidonotus Fisher, 1905

13, 54, 90, 92

Hippasteria pacifica Ludwig, 1905

Genus Mediaster Stimpson, 1857

Mediaster transfuga Ludwig, 1905 REV

45, 120

Family Ophidiasteridae Verrill, 1870

Genus Linckia Nardo, 1834

Linckia columbiae Gray, 1840 NAY, JAL, MAR, REV

1, 7, 8, 13, 15, 41, 43, 45, 81, 91, 92, 126, 135

Genus Narcissia Gray, 1840

Narcissia gracilis A.H. Clark, 1916 N AY

Genus Pharia Gray, 1840

Pharia pyramidata (Gray, 1840) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

7, 8, 13, 15, 39, 44, 73, 91, 92, 96, 104, 107,

113, 120, 129

Pharia piramidata (Gray, 1840)

Pharia pyramidatus (Gray, 1840)

33, 43, 45, 58, 72, 109, 110, 111, 112, 121, 124, 126

Pharia pyramidatus pyramidatus (Gray, 1840)

Pharia pyrimidata (Gray, 1840)

Genus Phataria Gray, 1840

Phataria unifascialis (Gray, 1840) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

7, 15, 22, 29, 33, 39, 42, 43, 44, 45, 58, 72,

73, 81, 84, 87, 91, 92, 96, 104, 107, 109, 110, 111, 112, 113, 120, 121, 124, 126, 129

Phataria unifacialis (Gray, 1840)

8, 13

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Family Mithrodiidae Viguier, 1878

Genus Mithrodia Gray, 1840

Mithrodia bradleyi Verrill, 1867 NAY, COL, MICH, MAR, ISA, MAT, REV

7, 15, 33, 41, 43, 44, 45, 58, 79, 81, 87, 91, 92, 104,

107, 108, 109, 110, 120, 124, 126, 129, 134

Family Oreasteridae Fisher, 1908

Genus Nidorellia Gray, 1840

Nidorellia armata (Gray, 1840) JAL, COL, MICH, MAR, ISA, MAT, REV

58, 81, 104, 107, 108, 109, 110, 129

Nidoriella armata (Gray, 1840)

7, 8, 13

Genus Pentaceraster Döderlein, 1916

Pentaceraster cumingi (Gray, 1840) NAY, JAL, COL, MICH, ISA, MAT, REV

58, 81, 91, 92, 104, 107, 109, 110, 111, 113, 124,

126

Oreaster occidentalis Verrill, 1867

7, 77

Family Solasteridae Viguier, 1878

Genus Lophaster Verrill, 1878

Lophaster furcilliger Fisher, 1905 MAR

Lophaster validus (Ludwig, 1905)

90, 91

Sarkaster validus Ludwig, 1905

Order Forcipulatida Perrier, 1884

Family Asteriidae Gray, 1840

Genus Sclerasterias Perrier, 1891

Sclerasterias heteropaes Fisher, 1924 REV

7, 13, 15, 90, 92, 126, 135

Family Heliasteridae Viguier, 1879

Genus Heliaster Gray, 1840

Heliaster helianthus (Lamarck, 1816) NAY, MICH

58, 81, 104

Heliaster microbrachius Xantus, 1860 JAL, COL

8, 13, 81

Heliaster polybrachius H.L. Clark, 1907 JAL

Family Pedicellasteridae Perrier, 1884

Genus Ampheraster Fisher, 1923

Ampheraster marianus (Ludwig, 1905) MAR

13, 55, 56, 90

Sporasterias mariana Ludwig, 1905

Order Spinulosida Perrier, 1884

Family Echinasteridae Verrill, 1867

Genus Echinaster Müller & Troschel, 1840

Subgenus Othilia Gray, 1840

Echinaster (Othilia) tenuispina Verrill, 1871 MAR

Echinaster tenuispina Verrill, 1871

Echinaster (Othilia) teniuspinus Verrill, 1871

Genus Henricia Gray, 1840

Henricia clarki Fisher, 1910 REV

7, 13, 15, 45, 54, 56, 74, 81, 90, 92, 120, 124

Henricia gracilis (Ludwig, 1905) MAR

90, 91, 92

Cribrella gracilis Ludwig, 1905

Henricia seminudus (A.H. Clark, 1916) REV

45, 81, 120, 124

Cyllaster seminuda A.H. Clark, 1916

13, 15, 34

Henricia seminuda (A.H. Clark, 1916)

Order Notomyotida Ludwig, 1910

Family Benthopectinidae Verrill, 1899

Genus Nearchaster Fisher, 1911

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Subgenus Nearchaster Fisher, 1911

Nearchaster (Nearchaster) aciculosus (Fisher, 1910) REV

Nearchaster aciculosus (Fisher, 1910)

7, 90, 120

Saraster insignis A.H. Clark, 1916

13, 15, 34

Class Ophiuroidea Gray, 1840

Order Euryalida Lamarck, 1816

Family Asteronychidae Ljungman, 1867

Genus Astrodia Verrill, 1899

Astrodia excavata (Lütken & Mortensen, 1899) MAR

64, 92, 106

Asteronyx excavata Lütken & Mortensen, 1899

13, 38, 50, 52, 63, 81, 88

Family Euryalidae Gray, 1840

Genus Asteroschema Örsted & Lütken in Lütken, 1856

Asteroschema sublaeve Lütken & Mortensen, 1899 MAR

50, 63, 64

Family Gorgonocephalidae Ljungman, 1867

Genus Astrocaneum Döderlein, 1911

Astrocaneum spinosum (Lyman, 1875) MAR

63, 64

Asterocaneum spinosum (Lyman, 1875)

Genus Astrodictyum Döderlein, 1927

Astrodictyum panamense (Verrill, 1867) REV

6, 45, 63, 76

Order Ophiurida Müller & Troschel, 1840, restricted sensu O’Hara, Hugall, Thuy, Stöhr, & Martynov, 2017

Family Ophiosphalmidae O’Hara, Stöhr, Hugall, Thuy, & Martynov, 2018

Genus Ophiosphalma H.L. Clark, 1941

Ophiosphalma variabile (Lütken & Mortensen, 1899) JAL, COL, MAR, REV

Ophiomusium variabile Lütken & Mortensen, 1899

45,

63, 64, 81, 88, 92, 120, 123, 124

Family Ophiomusaidae O’Hara, Stöhr, Hugall, Thuy, & Martynov, 2018

Genus Ophiomusa Hertz, 1927

Ophiomusa lymani (Wyville-Thomson, 1873) MAR

Ophiomusium lymani Wyville-Thomson, 1873

63, 64, 87

Family Ophiuridae Müller & Troschel, 1840

Genus Ophiura Lamarck, 1801

Subgenus Ophiura Lamarck, 1801

Ophiura (Ophiura) scutellata (Lütken & Mortensen, 1899) MAR

63, 64

Homalophiura scutellata (Lütken & Mortensen, 1899)

38, 52

Ophioglypha scutellata Lütken & Mortensen, 1899

Ophiura scutellata (Lütken & Mortensen, 1899)

90, 92

Genus Ophiuroglypha Hertz, 1927

Ophiuroglypha irrorata irrorata (Lyman, 1878) MAR

Ophiura (Ophiuroglypha) irrorata irrorata (Lyman, 1878)

63, 64

Ophiura irrorata (Lyman, 1878)

Family Ophiopyrgidae Perrier, 1893

Genus Gymnophiura Lütken & Mortensen, 1899

Gymnophiura mollis Lütken & Mortensen, 1899 MAR

63, 64, 90, 91, 92

Order Ophioscolecida O’Hara, Hugall, Thuy, Stöhr, & Martynov, 2017

Family Ophioscolecidae Lütken, 1869

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Genus Ophiuroconis Matsumoto, 1915

Ophiuroconis bispinosa Ziesenhenne, 1937 JAL, COL, REV

45, 63, 64, 70, 81, 92, 120, 123, 124

Order Ophiacanthida O’Hara, Hugall, Thuy, Stöhr, & Martynov, 2017

Family Ophiotomidae Paterson, 1985

Genus Ophiomitra Lyman, 1869

Ophiomitra granifera Lütken & Mortensen, 1899 NAY, MAR

63, 64, 87, 88

Ophiacantha granifera (Lütken & Mortensen, 1899)

Ophioplinthaca granifera (Lütken & Mortensen, 1899)

38, 52, 92

Ophiomitra partita Lütken & Mortensen, 1899 MAR

63, 64, 88

Ophiacantha partita (Lütken & Mortensen, 1899)

Ophioplinthaca partita (Lütken & Mortensen, 1899)

38, 52, 92

Family Ophiacanthidae Ljungman, 1867

Genus Ophiacantha Müller & Troschel, 1842

Ophiacantha costata Lütken & Mortensen, 1899 MAR

13, 38, 52, 63, 64, 83, 88, 92

Ophiacantha hirta Lütken & Mortensen, 1899 MAR

52, 63, 64, 81, 88, 92

Ophiacantha moniliformis Lütken & Mortensen, 1899 MAR, REV

38, 45, 52, 63, 64, 81 88, 92, 120, 123, 124

Ophiacantha pyriformis Ziesenhenne, 1937 REV

7, 13, 15, 45, 52, 63, 90, 135

Genus Ophiophthalmus Matsumoto, 1917 (temporary genus)

Ophiophthalmus diplasia (H.L. Clark, 1911) REV

Ophiacantha diplasia H.L. Clark, 1911

45, 63, 92

Family Ophiodermatidae Ljungman, 1867

Genus Ophioderma Müller & Troschel, 1840

Ophioderma panamense Lütken, 1859 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

7, 41, 42, 77, 79, 81, 84, 91, 92, 104,

109, 110, 111, 123, 124, 137

Ophiocryptus granulosus Nielsen, 1932

Ophioderma panamensis Lütken, 1859

45, 62, 63, 64

Ophioderma teres (Lyman, 1860) NAY, JAL, COL, MAR, REV*

11, 13, 62, 63, 64, 81, 92

Ophioderma variegatum Lütken, 1856 NAY, JAL, COL, MAR, ISA*, REV

7, 11, 13, 15, 27, 29, 39, 41, 45, 77, 79, 81, 84, 91, 92,

111, 120, 124, 134, 135

Ophioderma variegata Lütken, 1856

63, 64, 123, 137

Genus Diopederma H.L. Clark, 1913

Diopederma daniana (Verrill, 1867) JAL, MICH

63, 64, 104

Diopederma danianum (Verrill, 1867)

11, 39, 58, 81

Family Ophiomyxidae Ljungman, 1867

Genus Ophiomyxa Müller & Troschel, 1842

Ophiomyxa panamensis Lütken & Mortensen, 1899 REV

7, 63, 77, 91, 92

Family Ophiocomidae Ljungman, 1867

Genus Ophiocoma L. Agassiz, 1836

Ophiocoma aethiops Lütken, 1859 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

7, 11, 13, 15, 29, 41, 42, 43, 44, 45, 58, 59,

62, 63, 64, 79, 81, 84, 87, 91, 92, 104, 109, 110, 111, 120, 121, 122, 123, 124, 130, 134, 135

Genus Ophiocomella A.H. Clark, 1939

Ophiocomella alexandri (Lyman, 1860) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

Ophiocoma alexandri Lyman, 1860

7, 11, 13, 15, 29, 41, 42, 43, 44, 45, 49, 58, 59, 62, 63, 64, 79, 81, 84, 87, 91, 92, 104, 109, 110, 111, 120, 121,

123, 124, 130, 134

Ophiocomella schmitti A.H. Clark, 1939 NAY, COL, MAR*, REV

45, 57, 63, 64, 81, 123, 124

Ophiocomella sexradia (Duncan, 1887) MAR, REV

45, 63, 64, 92

Order Ophioleucida O’Hara, Hugall, Thuy, Stöhr, & Martynov, 2017

Family Ophiernidae O’Hara, Stöhr, Hugall, Thuy, & Martynov, 2018

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Genus Ophiernus Lyman, 1878

Ophiernus adspersus annectens Lütken & Mortensen, 1899 MAR

63, 64, 89

Ophiernus adspersus adspersus Lyman, 1883

Ophiernus polyporum Lütken & Mortensen, 1899

38, 52

Ophiernus polyporus Lütken & Mortensen, 1899

Order Amphilepidida O’Hara, Hugall, Thuy, Stöhr, & Martynov, 2017

Family Ophiolepididae Ljungman, 1867

Genus Ophiolepis Müller & Troschel, 1840

Ophiolepis crassa Nielsen, 1932 COL*, MAR, ISA*, MAT

63, 64, 92

Ophiolepis pacifica Lütken, 1856 NAY, JAL, MICH*, MAR, MAT, REV*

62, 63, 64, 81

Ophiozona pacifica (Lütken, 1856)

11, 13, 29, 92

Ophiolepis plateia Ziesenhenne, 1940 JAL, ISA*

13, 52, 63, 64, 81, 115, 136

Ophiolepis variegata Lütken, 1856 NAY, JAL, COL, MAR

11, 39, 63, 64, 81, 87, 92, 123, 124

Family Ophionereididae Ljungman, 1867

Genus Ophionereis Lütken, 1859

Ophionereis annulata (Le Conte, 1851) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

7, 13, 15, 45, 58, 59, 61, 62, 63, 64,

81, 91, 92, 104, 109, 110, 111, 120, 123, 124, 130, 135

Ophiocoma annulata (Le Conte, 1851)

Ophionereis dictiota Ziesenhenne, 1940

Ophionereis dictyota Ziesenhenne, 1940

11, 45, 104, 123, 124, 134

Ophionereis perplexa Ziesenhenne, 1940 JAL, MAT

Family Amphiuridae Ljungman, 1867

Genus Amphiura Forbes, 1843

Amphiura seminuda Lütken & Mortensen, 1899 NAY, REV

28, 63, 92

Amphiura serpentina Lütken & Mortensen, 1899 MAR

52, 63, 64, 92, 131

Amphiura serpentina var. a Lütken & Mortensen, 1899

Subgenus Amphiura (Amphiura) Forbes, 1843

Amphiura (Amphiura) assimilis Lütken & Mortensen, 1899 MAR

Amphiodia assimilis (Lütken & Mortensen, 1899)

63, 64, 90

Amphiura assimilis Lütken & Mortensen, 1899

Genus Amphichondrius Nielsen, 1932

Amphichondrius granulatus (Lütken & Mortensen, 1899) N AY

Amphichondrius unamexici Caso, 1983

25, 28, 90, 92

Amphichondrius laevis Ziesenhenne, 1940 JAL

52, 63, 64, 81, 87, 115, 136

Genus Amphiodia Verrill, 1899

Amphiodia occidentalis (Lyman, 1860) NAY

63, 64, 81

Amphiodia platyspina Nielsen, 1932 N AY

63, 64, 81

Amphiodia psara H.L. Clark, 1935 NAY*

Amphiodia sculptilis Ziesenhenne, 1940 JAL

52, 63, 64, 81, 136

Amphiodia tabogae Nielsen, 1932 NAY*

Amphiodia violacea (Lütken, 1856) N AY

63, 64, 81

Subgenus Amphispina Nielsen, 1932

Amphiodia (Amphispina) digitata Nielsen, 1932 NAY

63, 64, 81

Amphiodia (Amphispina) urtica (Lyman, 1860) NAY, MAR

63, 64, 81

Genus Amphipholis Ljungman, 1866

Amphipholis elevata Nielsen, 1932 MAR

63, 64, 92

Amphipholis pugetana (Lyman, 1860) NAY, REV*

63, 64, 81

Amphipholis squamata (Delle Chiaje, 1828) NAY, JAL, COL, MAT, ISA*

59, 63, 64

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Genus Dougaloplus A.M. Clark, 1970

Dougaloplus notacanthus (Lütken & Mortensen, 1899) MAR

63, 64, 92

Amphiura notacantha Lütken & Mortensen, 1899

52, 81, 88

Genus Microphiopholis Turner, 1985

Microphiopholis geminata (Le Conte, 1851) NAY*

Microphiopholis platydisca (Nielsen, 1932) NAY, COL

63, 64

Amphipholis platydisca Nielsen, 1932

Microphiopholis puntarenae (Lütken, 1856) MAR

63, 64

Amphipholis puntarenae (Lütken, 1856)

Genus Ophiocnida Lyman, 1865

Ophiocnida hispida (Le Conte, 1851) NAY*, JAL, COL, MAR, MAT

59, 62, 63, 64, 92

Genus Ophiophragmus Lyman, 1865

Ophiophragmus lonchophorus Ziesenhenne, 1940 NAY, JAL

13, 52, 63, 64, 90, 136

Ophiophragmus marginatus (Lütken, 1856) NAY, JAL

28, 63, 64, 81, 111

Ophiophragmus papillatus Ziesenhenne, 1940 NAY, JAL, MICH, MAT, REV*

64, 104

Ophiophragmus paucispinus Nielsen, 1932 MAR*

Genus Ophiostigma Lütken, 1856

Ophiostigma tenue Lütken, 1856 JAL, MAR, ISA, MAT

35, 63, 64, 92

Family Ophiactidae Matsumoto, 1915

Genus Ophiactis Lütken, 1856

Ophiactis savignyi (Müller & Troschel, 1842) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

7, 11, 13, 23, 29, 45, 58, 59,

62, 63, 64, 81, 91, 104, 111, 120, 121, 122, 123, 124, 130, 135

Ophiactis simplex (Le Conte, 1851) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

58, 59, 62, 63, 64, 81, 91, 92, 104, 111

Genus Hemipholis Lyman, 1865

Hemipholis gracilis Verrill, 1867 NAY, JAL

63, 64, 71

Family Ophiopholidae O‘Hara, Stöhr, Hugall, Thuy, & Martynov, 2018

Genus Ophiopholis Müller & Troschel, 1842

Ophiopholis bakeri McClendon, 1909 REV

63, 92

Family Ophiotrichidae Ljungman, 1867

Genus Ophiothrix Müller & Troschel, 1840

Ophiothrix galapagensis Lütken & Mortensen, 1899 MAR, REV

7, 13, 15, 45, 63, 64, 91, 92, 135

Subgenus Ophiothrix A.M. Clark, 1966

Ophiothrix (Ophiothrix) rudis Lyman, 1874 NAY, JAL, MICH, MAR, MAT, REV

62, 63, 64

Ophiothrix rudis Lyman, 1874

58, 81, 92, 104

Ophiotrix rudis Lyman, 1874

Ophiothrix (Ophiothrix) spiculata Le Conte, 1851 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

62, 63, 64, 104

Ophiothrix spiculata Le Conte, 1851

11, 13, 29, 39, 58, 59, 81, 87, 91, 92, 111, 123, 124

Genus Ophiothela Verrill, 1867

Ophiothela mirabilis Verrill, 1867 NAY, JAL, COL, MICH, ISA, MAT

58, 59, 62, 63, 64, 81, 104, 109, 110, 111

Ophiothrix mirabilis (Verrill, 1867)

Class Echinoidea Leske, 1778

Order Cidaroida Claus, 1880

Family Cidaridae Gray, 1825

Genus Eucidaris Pomel, 1883

Eucidaris thouarsii (L. Agassiz & Desor, 1846) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

7, 9, 13, 15, 21, 32, 33,

39, 41, 42, 43, 44, 45, 59, 65, 72, 73, 75, 77, 81, 84, 87, 92, 104, 107, 108, 109, 110, 111, 112, 113, 120, 121, 124, 126, 129, 134, 135

Cidaris Thouarsii L. Agassiz & Desor, 1846

Eucidaris Thouarsii (L. Agassiz & Desor, 1846)

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Eucidaris thourarsii (L. Agassiz & Desor, 1846)

Eucidaris thowarsii (L. Agassiz & Desor, 1846)

Genus Hesperocidaris Mortensen, 1928

Hesperocidaris asteriscus H.L. Clark, 1948 JAL, MICH, ISA, REV

7, 13, 15, 21, 45, 58, 92, 104, 111, 113, 126

Hesperocidaris perplexa (H.L. Clark, 1907) REV

7, 13, 15, 21, 45, 65, 81, 92, 97, 120, 124, 135

Tretocidaris perplexa H.L. Clark, 1907

36, 51

Order Diadematoida Duncan, 1889

Family Diadematidae Gray, 1855

Genus Astropyga Gray, 1825

Astropyga pulvinata (Lamarck, 1816) JAL, COL, MICH, MAR, MAT, REV

9, 21, 33, 41, 44, 45, 58, 65, 73, 81, 87, 91, 92, 104,

107, 113

Astropygia pulvinata (Lamarck, 1816)

126

Genus Centrostephanus Peters, 1855

Centrostephanus coronatus (Verrill, 1867) JAL, COL, MICH, MAR, ISA, MAT

42, 43, 44, 72, 73, 81, 96, 107, 109, 110, 111, 112,

113, 124, 129

Genus Diadema Gray, 1825

Diadema mexicanum A. Agassiz, 1863 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

1, 5, 7, 13, 15, 21, 22, 29, 31, 33, 41,

42, 43, 44, 45, 53, 58, 72, 73, 74, 75, 79, 81, 84, 91, 92, 96, 98, 104, 107, 108, 109, 110, 111, 112, 113, 120, 121, 124, 126, 129, 134

Centrechinus mexicanus (A. Agassiz, 1863)

7, 9, 40, 65, 77, 135

Diadema mexicanus A. Agassiz, 1863

Order Arbacioida Gregory, 1900

Family Arbaciidae Gray, 1855

Genus Arbacia Gray, 1835

Arbacia stellata (Blainville, 1825; ?Gmelin, 1788) NAY, JAL, MAR, ISA

Arbacia incisa (A. Agassiz, 1863)

9, 13, 21, 42, 43, 81, 84, 109, 110, 111, 113

Order Camarodonta Jackson, 1912

Family Echinometridae Gray, 1855

Genus Echinometra Gray, 1825

Echinometra insularis H.L. Clark, 1912 REV

37, 51, 100

Echinometra oblonga (Blainville, 1825) REV

1, 7, 13, 15, 21, 31, 37, 40, 41, 45, 74, 79, 81, 91, 92, 95, 120, 124, 126, 134, 135

Echinometra mathei oblonga (Blainville, 1825)

Echinometra vanbrunti A. Agassiz, 1863 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

1, 7, 15, 31, 33, 41, 42, 43, 44,

45, 58, 73, 79, 81, 84, 87, 91, 92, 104, 107, 108, 109, 110, 111, 113, 120, 121, 124, 126, 129, 134

Echinometra van brunti A. Agassiz, 1863

13, 21, 22, 29

Echinometra Van brunti A. Agassiz, 1863

2, 9

Echinometra van-brunti A. Agassiz, 1863

Echinometra VanBrunti A. Agassiz, 1863

Echinometra vanbruntii A. Agassiz, 1863

Heliocidaris stenopora H.L. Clark, 1912

Genus Heterocentrotus Brandt, 1835

Heterocentrotus mamillatus (Linnaeus, 1758) REV

7, 32, 108

Family Toxopneustidae Troschel, 1872

Genus Toxopneustes L. Agassiz, 1841

Toxopneustes roseus (A. Agassiz, 1863) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

7, 13, 15, 21, 22, 29, 33, 40, 41, 42,

43, 44, 45, 58, 65, 73, 79, 81, 87, 91, 92, 96, 104, 107, 109, 110, 111, 112, 113, 121, 124, 126, 129

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Genus Tripneustes L. Agassiz, 1841

Tripneustes depressus A. Agassiz, 1863 JAL, COL, MICH, MAR, ISA, MAT, REV

1, 7, 13, 15, 18, 20, 21, 31, 33, 40, 41, 42,

43, 44, 45, 65, 73, 75, 79, 80, 81, 91, 92, 99, 104, 107, 108, 109, 110, 111, 113, 120, 121, 124, 126, 129, 130, 134, 135

Tripneustes gratilla (Linnaeus, 1758) REV

Order Cassiduloida Claus, 1880

Family Cassidulidae L. Agassiz & Desor, 1847

Genus Rhyncholampas A. Agassiz, 1869

Rhyncholampas pacificus (A. Agassiz, 1863) MAR, REV

Cassidulus pacifica (A. Agassiz, 1863)

7, 91, 92

Cassidulus pacificus (A. Agassiz, 1863)

13, 15, 26, 40, 101

Order Clypeasteroida A. Agassiz, 1872

Family Clypeasteridae L. Agassiz, 1835

Genus Clypeaster Lamarck, 1801

Clypeaster europacificus H.L. Clark, 1914 REV

7, 13, 15, 24, 40, 51, 81, 92, 120, 124, 135

Clypeaster (Alexandria) europacificus H.L. Clark, 1914

102

Clypeaster ochrus H.L. Clark, 1914 MAR, ISA, REV

13, 24, 39, 81, 92, 93, 120, 124, 126

Clypeaster (Stolonoclypus) ochrus H.L. Clark, 1914

Clypeaster rotundus (A. Agassiz, 1863) JAL, MAR, ISA, REV

7, 15, 24, 40, 43, 77, 92

Clypeaster (Stolonoclypus) rotundus A. Agassiz, 1863

91, 102

Clypeaster speciosus Verrill, 1870 NAY, COL, REV

7, 13, 15, 24, 39, 40, 41, 45, 77, 92, 126

Clypeaster (Stolonoclypus) speciosus Verrill, 1870

91, 102

Family Dendrasteridae Lambert, 1900

Genus Dendraster L. Agassiz in Agassiz & Desor, 1847

Dendraster excentricus (Eschscholtz, 1831) ISA

Family Mellitidae Stefanini, 1912

Genus Mellita L. Agassiz, 1841

Mellita notabilis H.L. Clark, 1947 MICH

58, 67, 81, 104

Genus Encope L. Agassiz, 1840

Encope micropora L. Agassiz, 1841 JAL, MICH

Encope fragilis H.L. Clark, 1948

Encope micropora fragilis H.L. Clark, 1948

Encope perspectiva L. Agassiz, 1841

40, 58, 81, 104

Encope wetmorei A.H. Clark, 1946

58, 81, 104

Encope micropora insularis H.L. Clark, 1948 REV

24, 102

Encope insularis H.L. Clark, 1948

13, 15, 40, 51

Genus Lanthonia Coppard, 2016

Lanthonia longifissa (Michelin, 1858) COL

Mellita longifissa Michelin, 1858

24, 81, 124

Order Holasteroida Durham & Melville, 1957

Family Plexechinidae Mooi & David, 1996

Genus Plexechinus A. Agassiz, 1898

Plexechinus cinctus A. Agassiz, 1898 MAR

3, 4, 13, 26, 51, 90, 92

Order Spatangoida L. Agassiz, 1840

Family Schizasteridae Lambert, 1905

Genus Brisaster Gray, 1855

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Brisaster townsendi (A. Agassiz, 1898) MAR

Briaster townsendi (A. Agassiz, 1898)

Brisaster towsendi (A. Agassiz, 1898)

13, 26

Schizaster townsendi A. Agassiz, 1898

Schizaster Townsendi A. Agassiz, 1898

3, 4

Family Prenasteridae Lambert, 1905

Genus Agassizia Valenciennes, 1846

Agassizia scrobiculata Valenciennes, 1846 N AY

26, 81

Family Brissidae Gray, 1855

Genus Brissopsis L. Agassiz, 1840

Brissopsis pacifica (A. Agassiz, 1898) REV

7, 13, 15, 26, 40, 91, 92

Genus Brissus Gray, 1825

Brissus latecarinatus (Leske, 1778) ISA

109, 110

Brissus obesus Verrill, 1867 MAR

Genus Meoma Gray, 1851

Meoma ventricosa grandis Gray, 1851 JAL, REV

81, 91, 93, 120

Meoma grandis Gray, 1851

7, 26, 40, 77, 92, 103

Meoma ventricosa (Lamarck, 1816)

45, 124, 126

Family Loveniidae Lambert, 1905

Genus Lovenia Desor in Agassiz & Desor, 1847

Lovenia cordiformis A. Agassiz, 1872 REV

13, 15, 26, 40, 41, 45, 77, 81, 92, 93, 124

Lovenia cardiformis A. Agassiz, 1872

Lovenia (Lovenia) cordiformis A. Agassiz, 1872

Class Holothuroidea Selenka, 1867

Order Dendrochirotida Grube, 1840

Family Psolidae Burmeister, 1837

Genus Lissothuria Verrill, 1867

Lissothuria imbricata Solís-Marín, Arriaga-Ochoa, Galván-Villa, & Laguarda-Figueras, 2018 JAL

125

Lissothuria ornata Verrill, 1867 JAL, MICH, MAT

104, 111, 113, 125, 127

Thyonepsolus beebei Deichmann, 1937

Genus Psolus Oken, 1815

Psolus pauper Ludwig, 1893 MAR

85, 94

Family Sclerodactylidae Panning, 1949

Genus Afrocucumis Deichmann, 1944

Afrocucumis ovulum (Selenka, 1867) NAY, JAL, MICH, MAT

104, 113, 116, 127

Genus Neothyone Deichmann, 1941

Neothyone gibber (Selenka, 1867) NAY, JAL, MICH, MAT, REV

7, 44, 47, 81, 84, 90, 104, 105, 111, 112, 113, 128

Thyone gibber (Selenka, 1867)

13, 15

Neothyone gibbosa Deichmann, 1941 JAL, MICH

58, 81, 104, 111, 113

Genus Pachythyone Deichmann, 1941

Pachythyone lugubris (Deichmann, 1939) JAL

111, 113

Pachythyone pseudolugubris Deichmann, 1941 JAL, MICH, MAT

104, 111, 113, 127

Genus Neopentamera Deichmann, 1941

Neopentamera anexigua Deichmann, 1941 JAL

111, 113

Genus Euthyonidiella Heding & Panning, 1954

Euthyonidiella zacae (Deichmann, 1938) REV

120

Phyllophorus zacae Deichmann, 1938

45, 81, 124

236

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Family Phyllophoridae Östergren, 1907

Genus Pentamera Ayres, 1852

Pentamera chierchiae (Ludwig, 1886) JAL, MICH, MAT, REV

Pentamera chierchia (Ludwig, 1886)

7, 13, 15, 45, 47, 81, 91, 92, 104, 111, 113, 124, 126, 128

Genus Thyone Oken, 1815

Thyone bidentata Deichmann, 1941 JAL

13, 47, 113, 119

Thyone parafusus Deichmann, 1941 JAL

13, 47, 90, 92, 119, 130

Family Cucumariidae Ludwig, 1894

Genus Abyssocucumis Heding, 1942

Abyssocucumis abyssorum (Théel, 1886) NAY

Genus Cucumaria de Blainville, 1830

Cucumaria flamma Solís-Marín & Laguarda-Figueras, 1999 NAY, JAL, COL, MICH, MAT

44, 58, 73, 81, 104, 107, 111, 113,

118, 121, 124, 128, 129

Pattalus mollis Selenka, 1868

44, 105

Genus Neocucumis Deichmann, 1944

Neocucumis veleronis (Deichmann, 1941) JAL, COL, MICH, MAT

81, 104, 111, 113, 124, 127

Genus Pseudocnus Panning, 1949

Pseudocnus californicus (Semper, 1868) NAY, JAL, COL, MICH, MAT

44, 58, 81, 104, 105, 111, 113, 116, 124, 128

Genus Trachythyone Studer, 1876

Trachythyone peruana (Semper, 1868) NAY, JAL

81, 113

Genus Thyonella Verrill, 1872

Thyonella mexicana (Deichmann, 1941) MICH

104

Genus Leptopentacta H.L. Clark, 1938

Leptopentacta panamica Deichmann, 1941 JAL

13, 47, 90

Family Ypsilothuriidae Heding, 1942

Genus Ypsilothuria Perrier, 1886

Ypsilothuria bitentaculata (Ludwig, 1893) MAR

92, 94

Shpaerothuria bitentaculata Ludwig, 1893

Order Holothuriida Miller, Kerr, Paulay, Reich, Wilson, Carvajal & Rouse, 2017

Family Holothuriidae Burmeister, 1837

Genus Holothuria Linnaeus, 1767

Subgenus Cystipus Haacke, 1880

Holothuria (Cystipus) inhabilis Selenka, 1867 REV

91, 126

Holothuria inhabilis Selenka, 1867

7, 46, 92, 130

Holothuria parinhabilis Cherbonnier, 1951

Jaegerothuria inhabilis (Selenka, 1867)

Holothuria (Cystipus) rigida (Selenka, 1867) NAY, JAL, COL, MICH, MAT

44, 58, 81, 104, 105, 113, 124

Holothuria rigida (Selenka, 1867)

44, 84, 111

Subgenus Halodeima Pearson, 1914

Holothuria (Halodeima) atra Jaeger, 1833 NAY, JAL, ISA

Holothuria atra Jaeger, 1833

109, 110

Holothuria (Halodeima) inornata Semper, 1868 NAY, JAL, MICH, ISA, MAT, REV

104, 107, 113, 116, 119, 120, 126, 127

Holothuria inornata Semper, 1868

13, 15, 111, 112, 129

Holothuria (Halodeima) kefersteinii (Selenka, 1867) NAY, JAL, MICH, MAR, ISA, MAT, REV

104, 107

Holothuria kefersteiini (Selenka, 1867)

Holothuria kefersteini (Selenka, 1867)

44, 45, 84, 92

Holothuria kefersteinii (Selenka, 1867)

129

Holothuria kefersteni (Selenka, 1867)

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Holothuria (Halodeima) kefersteini (Selenka, 1867)

58, 81, 91, 105, 109, 110, 116, 119, 120, 124, 126

Holothuria (Paraholothuria) riojai Caso, 1963

Ludwigothuria kefersteini (Selenka, 1867)

48, 77

Subgenus Lessonothuria Deichmann, 1958

Holothuria (Lessonothuria) pardalis Selenka, 1867 JAL

91, 113

Holothuria pardalis Selenka, 1867

13, 90

Lessonothuria pardalis (Selenka, 1867)

Subgenus Mertensiothuria Deichmann, 1958

Holothuria (Mertensiothuria) hilla Lesson, 1830 MAR, ISA, REV

120, 126

Brandtothuria gyrifer (Selenka, 1867)

Holothuria gyrifer (Selenka, 1867)

13, 16

Holothuria hilla Lesson, 1830

45, 111

Holothuria (Thymiosycia) hilla Lesson, 1830

81, 109, 110, 124

Holothuria (Mertensiothuria) leucospilota (Brandt, 1835) COL, MAR, REV

81, 91, 120, 124, 126

Holothuria leucospilota (Brandt, 1835)

7, 13, 15, 31, 41, 45, 79, 90, 92

Mertensiothuria leucospilota (Brandt, 1835)

Holothuria (Mertensiothuria) viridiaurantia Borrero-Pérez & Vanegas-González, 2019 NAY, JAL, MICH, MAT

Holothuria hilla Lesson, 1830

111, 129

Holothuria (Mertensiothuria) hilla Lesson, 1830

104, 113, 116, 127

Holothuria (Thymiosycia) hilla Lesson, 1830

Subgenus Platyperona Rowe, 1969

Holothuria (Platyperona) difficilis Semper, 1868 NAY, JAL, MICH, MAR, MAT, REV

81, 91, 104, 116, 120, 124, 126, 127

Holothuria difficilis Semper, 1868

7, 13, 15, 45, 46, 112, 129, 130

Microthele difficilis (Semper, 1868)

Subgenus Selenkothuria Deichmann, 1958

Holothuria (Selenkothuria) carere Honey-Escandón, Solís-Marín, & Laguarda-Figueras, 2011 MICH

82, 104

Holothuria (Selenkothuria) lubrica Selenka, 1867 NAY, JAL, MICH, MAR, ISA, REV

58, 81, 91, 104, 109, 110, 113, 116, 120,

124, 126

Holothuria lubrica Selenka, 1867

7, 12, 13, 15, 45, 58, 92, 111, 112, 121

Holothuria pseudolubrica Cherbonnier, 1951

Selenkothuria lubrica (Selenka, 1867)

41, 48

Holothuria (Selenkothuria) portovallartensis Caso, 1954 NAY, JAL, MICH, REV

58, 81, 91, 104, 116, 119, 126

Holothuria portovallartensis Caso, 1954

12, 13, 92

Selenkothuria portovallartensis (Caso, 1954)

Subgenus Semperothuria Deichmann, 1958

Holothuria (Semperothuria) imitans Ludwig, 1875 NAY, JAL, COL, MICH, ISA, MAT, REV

81, 91, 104, 105, 109, 110,

116, 120, 124, 126

Holothuria imitans Ludwig, 1875

7, 13, 15, 44, 45, 92, 129

Holothuria imitans var. polymorpha Caso, 1962

Semperothuria imitans (Ludwig, 1875)

48, 77

Holothuria (Semperothuria) languens Selenka, 1867 JAL, MICH, MAR

81, 104

Holothuria languens Selenka, 1867

Semperothuria languens (Selenka, 1867)

Subgenus Stauropora Rowe, 1969

Holothuria (Stauropora) fuscocinerea Jaeger, 1833 JAL, MICH, MAR, ISA, REV

Holothuria fuscocinerea Jaeger, 1833

45, 73, 78

Holothuria fuscoscinerea Jaeger, 1833

Holothuria (Mertensiothuria) fuscocinerea Jaeger, 1833

81, 109, 110, 124

Holothuria (Stauropora) pluricuriosa Deichmann, 1937

120, 126

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Subgenus Theelothuria Deichmann, 1958

Holothuria (Theelothuria) paraprinceps Deichmann, 1937 REV

126

Holothuria paraprinceps Deichmann, 1937

Theelothuria paraprinceps (Deichmann, 1937)

48, 77

Subgenus Thymiosycia Pearson, 1914

Holothuria (Thymiosycia) arenicola Semper, 1868 NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

44, 58, 81, 91, 104,

105, 107, 109, 110, 113, 116, 119, 120, 124, 126

Brandtothuria arenicola (Semper, 1868)

42, 48, 77

Holothuria arenicola Semper, 1868

7, 15, 44, 45, 73, 84, 92, 111, 121, 129

Holothuria (Thymiosycia) impatiens (Forskål, 1775) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

58, 81, 91, 104,

105, 107, 109, 110, 113, 116, 120, 124

Brandtothuria impatiens (Forskål, 1775)

42, 48, 77

Holothuria impatiens (Forskål, 1775)

7, 15, 43, 44, 45, 58, 84, 92, 111, 112, 124, 129

Genus Labidodemas Selenka, 1867

Labidodemas americanum Deichmann, 1938 NAY, JAL, COL, MICH, MAT, REV

44, 45, 81, 104, 105, 113, 116, 120, 124, 126,

129

Labidodemas americanus Deichmann, 1938

Labidodemas maccullochi (Deichmann, 1958) JAL, COL, MICH

111, 113

Holothuria (Irenothuria) maccullochi (Deichmann, 1958)

Labidodemas macullochi (Deichmann, 1958)

81, 124

Order Synallactida Miller, Kerr, Paulay, Reich, Wilson, Carvajal, & Rouse, 2017

Family Stichopodidae Haeckel, 1896

Genus Isostichopus Deichmann, 1958

Isostichopus fuscus (Ludwig, 1875) NAY, JAL, COL, MICH, MAR, ISA, MAT, REV

7, 42, 43, 44, 45, 48, 58, 72, 73, 81, 84,

91, 92, 104, 105, 107, 109, 110, 111, 112, 113, 119, 120, 121, 124, 126, 129, 133

Isostichopus fuscis (Ludwig, 1875)

Stichopus fuscus Ludwig, 1875

15, 17, 78

Order Elasipodida Théel, 1882

Family Laetmogonidae Ekman, 1926

Genus Pannychia Théel, 1882

Pannychia moseleyi Théel, 1882 JAL, COL, MAR

92, 94

Laetmophasma fecundum Ludwig, 1893

Order Molpadida Haeckel, 1896

Family Molpadiidae Müller, 1850

Genus Molpadia Cuvier, 1817

Molpadia musculus Risso, 1826 MAR

Ankyroderma danielsseni Théel, 1886

Order Apodida Brandt, 1835

Family Chiridotidae Östergren, 1898

Genus Chiridota Eschscholtz, 1829

Chiridota aponocrita A.H. Clark, 1920 MICH

104

Chiridota rigida Semper, 1867 NAY, MICH, MAT

44, 81, 104, 105

Family Synaptidae Burmeister, 1837

Genus Epitomapta Heding, 1928

Epitomapta tabogae Heding, 1928 MICH

104

Genus Euapta Östergren, 1898

239

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Euapta godeffroyi (Semper, 1868) NAY, JAL, COL, MICH, MAR, ISA, REV

41, 43, 45, 60, 73, 81, 109, 110, 113

Genus Leptosynapta Verrill, 1867

Leptosynapta albicans (Selenka, 1867) REV

7, 15, 90, 94

COL = Colima; ISA = Isabel Island; JAL = Jalisco; MAR = Marías Islands; MAT = Marietas Islands; MICH = Michoacán;

NAY = Nayarit; REV = Revillagigedo Islands.

Invalid and doubtful records: The

records considered as invalid (52) or doubtful

(five) are based on: 1) erroneous or doubtful

distributions, 2) incorrect taxonomic identifica-

tions, or 3) entries in faunal list not backed up

by records in the primary literature.

There are some records considered invalid

because of inconsistencies in their geographi-

cal distributions. The ophiuroids Ophiocomella

ophiactoides and Astrophyton muricatum, and

the holothuroid Isostichopus badionotus have

been reported in the CMP (Honey-Escandón et

al., 2008; Solís-Marín et al., 2016b; CONANP,

2019), yet the species distribution range is

exclusively for the Caribbean and the Gulf

of México (Hendler, Miller, Pawson, & Kier,

1995; Solís-Marín et al., 2013a), meanwhile

the echinoids Echinothrix calamaris and Echi-

nothrix diadema inhabits the Indian Ocean, but

have been reported in REV (Sotelo-Casas &

Rodríguez-Troncoso, 2014). The holothuroid

Pentamera chiloensis is reported in Barra

de Navidad, JAL (Rodríguez-Troncoso et al.,

2013); this species distributes only in South

America (Perú and Chile; Solís-Marín et al.,

2013a). Records previously mentioned corre-

spond to erroneous records and must be invalid

(R.C. Sotelo-Casas and F.A. Solís-Marín, per-

sonal communication, 01 March 2020).

The crinoid Hyocrinus foelli (Solís-Marín

et al., 2013a; Solís-Marín et al., 2013b; Solís-

Marín, Laguarda-Figueras, & Honey-Escan-

dón, 2014) and the holothuroid Psychronaetes

hanseni (Maluf, 1991; Bautista-Romero et al.,

1994; Massin & Hendrickx, 2011) have been

reported in Clarion Island, REV, but according

to their original descriptions (Pawson, 1983;

Roux & Pawson, 1999) and the validation of

the geographical coordinates, all the records

corresponds to the Clarion-Clipperton Fracture

Zone which is located approximately 1 200 km

far away from REV. The echinoids Aporoci-

daris milleri and Kamptosoma asterias were

reported in NAY (station 96, 2 988-3 001 m

depth) by Parker (1963), yet the validation of

the geographic coordinates revealed that this

station is located in the entrance of the Gulf

of California, and far away from the maritime

zone of NAY. Moreover, K. asterias is distrib-

uted in the Central Pacific Ocean, Chile and

the Antarctica (Mooi, Constable, Lockhart, &

Pearse, 2004).

Furthermore, some records had inconsis-

tencies in their taxonomic identifications. Oph-

ionereis albomaculata (ophiuroid) has records

in ISA and MAR (Caso, 1961; Maluf, 1991;

Maluf & Brusca, 2005; Granja-Fernández et

al., 2015a; Granja-Fernández et al., 2017);

during our revision of museum collections, we

taxonomically reviewed all the material known

for O. albomaculata (Galápagos Islands and

Panamá), and none corresponded to México.

The material identified as O. albomaculata

from México must relate to another species

of Ophionereis. The collected material of the

holothuroid Holothuria (Lessonothuria) par-

dalis from REV (Solís-Marín et al., 2013a;

Sotelo-Casas & Rodríguez-Troncoso, 2014),

was previously identified as H. (Lessenothuria)

pardalis, yet it corresponds to a new species

in process of description (F.A. Solís-Marín,

personal communication, 01 April 2020). It

is important to mention that in the CMP, the

species only distributes in Tenacatita, JAL

(Deichmann, 1958).

Most of the entries are based on unbacked

up records of primary references. Bautista-

Romero et al. (1994) constructed a checklist of

echinoderms from REV according to previous

records (Fisher, 1911; Ziesenhenne, 1937; Caso,

1962b; Maluf, 1988), and reported: Florometra

serratissima (crinoid); Amphiaster insignis,

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Astropecten verrilli, Echinaster (Othilia) tenui-

spina, Heliaster kubiniji, Henricia aspera,

Henricia asthenactis, Henricia leviuscula,

Henricia polyacantha, Leiaster teres, Luidia

foliolata, Luidia phragma, Mediaster aequa-

lis, Narcissia gracilis, Odontaster crassus,

Tethyaster canaliculatus, and Thrissacanthias

penicillatus (asteroids); Amphiodia (Amphispi-

na) urtica, Amphipholis squamata, Amphiura

arcystata, Astrocaneum spinosum, Diopederma

daniana, Ophiolepis crassa, Ophionereis eury-

brachiplax, Ophiopaepale diplax and Ophiura

luetkenii (ophiuroids); and Arbacia stellata (as

Arbacia incisa), Centrostephanus coronatus,

Encope grandis, Encope micropora, Lytechinus

pictus, and Strongylocentrotus fragilis (echi-

noids). We consulted the previous literature

and we did not find these records on them or

in other literature references, therefore these

records are invalid.

Based on literature records, Lophaster

furcilliger (asteroid), Astrocaneum spinosum

(ophiuroid), and Centrostephanus coronatus

(echinoid) have been recorded in the pro-

gram of management and conservation of

REV (CONANP, 2019). The only records of

L. furcilliger in the CMP correspond to MAR

(Ludwig, 1905; Maluf, 1988; Maluf, 1991),

therefore their inclusion in REV is incorrect.

Despite it is very probable that A. spinosum

and C. coronatus distributes in REV, we did not

find any record in publications to back it up.

Other species reported in a program of manage-

ment and conservation (ISA; CONANP, 2005)

are the echinoids Clypeaster europacificus and

Lovenia cordiformis, but again no primary ref-

erences supporting these records were found,

and therefore are considered as invalid records.

Finally, based on a revision of literature,

the asteroids Paulia horrida and Luidia bel-

lonae were reported in MAR by Maluf (1988)

and Maluf and Brusca (2005), respectively;

nevertheless, we did not find any record of

these species for MAR but for REV. Similarly,

the asteroid Cnemidaster nudus has only one

mention for COL (Solís-Marín et al., 2016b),

but it corresponds to an erroneous record (F.A.

Solís-Marín, personal communication, 01 April

2020). For these reasons, we consider both

records as invalid.

The following records were considered

as doubtful. The ophiuroid Amphiodia peri-

ercta inhabits in the North Pacific Ocean (from

Aleutian Islands, Alaska to central California;

Lambert & Austin, 2007), yet it has a report in

Clarion Island, REV (Maluf, 1988; Bautista-

Romero et al., 1994). Due to the northern

affinity of A. periercta, its distribution in REV

is unlikely. The record of Hymenaster pentago-

nalis in REV was obtained by the Expedition

Ocean Exploration Trust/Nautilus Live 2017

(CONANP, 2019), yet this species distributes

only in Hawaii (Mah, 2020).

The echinoids Caenocentrotus gibbosus

(Clark, 1925; Grant & Hertlein, 1938; Maluf,

1988; Maluf,1991; Bautista-Romero et al.,

1994) and Lytechinus semituberculatus (Clark,

1925; Clark,1948; Mortensen, 1943b; Caso,

1962b; Maluf, 1988; Bautista-Romero et al.,

1994) have many mentions in Clarion Island,

REV, yet in all these references the locality is

treated as doubtful. It is important to empha-

size that both species inhabit in South America

(Galápagos Islands, Perú, and Chile; Grant

& Hertlein, 1938; Solís-Marín et al., 2013a),

thus their distribution in México is unlikely.

Similarly, the holothuroid Holothuria (Selenko-

thuria) theeli is recorded from REV as doubtful

(Hertlein, 1963; Maluf, 1991; Maluf & Brusca,

2005). In the Eastern Pacific, H. (Selenkothu-

ria) theeli inhabits in lower latitudes (i.e. Costa

Rica-Perú; Solís-Marín et al., 2013a) and in

México it has been collected only in Guerrero

(Honey-Escandón et al., 2008). For these rea-

sons, we determined these records as doubtful.

New records: Following review of muse-

um material, we add 16 new records of Ophiu-

roidea to different states (four NAY, one COL,

and one MICH) and islands (two MAR, four

ISA, and four REV) from the CMP (appendix

1). All the species were previously recorded

in areas from the CMP (Table 1), except-

ing Amphiodia psara, Amphiodia tabogae,

Microphiopholis geminata, and Ophiophrag-

mus paucispinus, which represent new records

for the CMP.

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Amphiodia psara has previous records in

USA (California; Clark, 1935) and México

(Gulf of California and Guerrero; Maluf, 1988;

Maluf & Brusca, 2005; Granja-Fernández et

al., 2015b). After the taxonomic revision of

A. psara in different museum collections, we

did not find material from Guerrero, but for

the Pacific of Baja California and Baja Cali-

fornia Sur, the Gulf of California (Ángel de la

Guarda Island, Tiburón Island, Consag Rock,

and Sonora), and NAY. The records from Baja

California, Baja California Sur and NAY are

new, being Punta Mita, Nayarit, the southern-

most area of its geographical distribution.

Amphiodia tabogae, M. geminata, and O.

paucispinus distribute along the Eastern Pacific

(México to Galápagos Islands; Solís-Marín et

al., 2013a). Specifically, in México, the three

species inhabit the Gulf of California; A. tabo-

gae has been reported also in Guerrero (Granja-

Fernández et al., 2015b). The finding of these

species in Punta Mita, NAY (A. tabogae and

M. geminata), and in MAR (O. paucispinus),

represents new records in the CMP, and fills

the distribution gap along the Mexican Pacific.

Diversity and distribution: The diversity

of the Echinodermata from the CMP is repre-

sented by 187 species of echinoderms belong-

ing to 27 orders, 63 families, 110 genera, and

16 subgenera (Table 2). Compared to the total

number of echinoderm species worldwide (7

437 species; WoRMS, 2020), the CMP shelters

2.5 %. The orders with the highest number

of species in the CMP were Amphilepidida

(Ophiuroidea, 40 species), Holothuriida and

Dendrochirotida (Holothuroidea, 21 species,

respectively), Ophiacanthida (Ophiuroidea, 16

species), and Valvatida (Asteroidea, 15 spe-

cies), meanwhile the best represented families

were Amphiuridae (Ophiuroidea, 26 species),

and Holothuriidae (Holothuroidea, 21 spe-

cies) (Table 1). On the other hand, the orders

Ophiurida (45 species), Aspidochirotida (22

species), and Dendrochirotida (22 species)

were the best represented in the Southern

Mexican Pacific (SMP; Granja-Fernández et al.

2015b). Differences in the numbers of recorded

orders between the CMP and the SMP are due

to recent changes in the systematics of the

Ophiuroidea (O´Hara, Stöhr, Hugall, Thuy &

Martynov, 2018) in which most of the mem-

bers previously included in the order Ophiu-

rida, now are in Amphilepidida, and in the

Holothuroidea, where most of the members of

Aspidochirotida were changed to Holothuriida

(Miller et al., 2017). If we compare both faunas

in the same classification system, similar trends

are shown, suggesting that both areas possess

similar faunas in terms of orders and families

of echinoderms.

In the CMP, the class Ophiuroidea was

the most numerous with 67 species, followed

by Holothuroidea (50), Asteroidea (35), Echi-

noidea (32), and Crinoidea (three) (Table 2).

In general, the number of species of Holo-

thuroidea, Asteroidea and Echinoidea were

similar between the CMP and the SMP, but

Ophiuroidea was represented by more spe-

cies in the CMP (67) than to the SMP (46);

meanwhile, the class Crinoidea had not records

in the SMP (Granja-Fernández et al., 2015b).

Differences on the Ophiuroidea are related

to the recent sampling effort directed in the

TABLE 2

Taxonomic categories of echinoderms from the Central Mexican Pacific

Class Orders Families Genera Subgenera Species

Crinoidea 2 2 2 0 3

Asteroidea 5 16 24 2 35

Ophiuroidea 6 20 35 4 67

Echinoidea 8 14 23 0 32

Holothuroidea 6 11 26 10 50

TOTAL 27 63 110 16 187

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CMP (Granja-Fernández et al., 2017), and the

extensive revision of samples from museum

collections (reflected in the new records of

species in this work). Nine species of Crinoi-

dea from México to Perú (Solís-Marín et al.,

2013a) and at least seven in the northern part

of the American continent (Lambert & Austin,

2007) are reported in the literature therefore,

finding crinoids in the CMP and SMP is likely.

Despite the above, a null to a scarce number of

species of Crinoidea are reported in both areas.

We consider that the explanation of this pattern

(and in general of the Eastern Pacific) is not

due to the natural biogeographic distribution of

the Crinoidea in the Eastern Pacific, yet to the

scarce prospection in deep waters (the natural

environment of the class in the Eastern Pacific),

as well as to the low interest and lack of taxo-

nomical specialists in the class.

As previously mentioned, the CMP is

represented by 187 species of echinoderms; of

those, 122 inhabit in ~ 640 km of the coastline

of the studied states, and 142 in an extensive

maritime area of the islands (~ 155 000 km

The SMP has not extensive islands such as

the CMP, therefore if only their coastlines are

compared, the latter possesses a lower number

of echinoderms species than the SMP (162 spe-

cies; Granja-Fernández et al., 2015b). In order

to have a better comprehension of the number

of species between both areas, a standardiza-

tion per area is required, therefore considering

the whole CMP (~ 640 km) has 5.2 species/

km, and the SMP (~ 1 200 km) possesses 7.4

species/km. The collecting effort in more areas,

habitats, and depths could explain higher biodi-

versity of echinoderms in the coastline of CMP,

but further collections of material and biogeo-

graphic analysis are required in order to better

understand their distribution pattern.

In terms of spatial distribution, the coast

of the state of JAL presented the highest num-

ber of species (84), followed by the coast of

NAY (74), MICH (63), and COL (55); among

the islands, REV showed the major number

of species (85) followed by MAR (81), MAT

(48), and ISA (44) (Table 3). Compared to the

most complete and recent works of echino-

derms by states (Honey-Escandón et al., 2008;

Rodríguez-Troncoso et al., 2013; Solís-Marín

et al., 2016b; Granja-Fernández et al., 2017;

Ríos-Jara et al., 2017; Nava-Bravo et al.,

2019) and islands (CONANP, 2005; CONANP,

2007a; CONANP, 2007b; CONANP, 2019;

Honey-Escandón et al., 2008; Solís-Marín et

al., 2013a; Granja-Fernández et al., 2017), this

work increases the number of recorded species

in 27 % for NAY, 49 % for JAL, 1.5 % for

MICH, 70 % for MAR, 64 % for ISA, 48 % for

MAT, 38 % for REV, and finally 45 % for the

entire CMP region. We found the same number

of species for COL as its most recent account of

echinoderms (Solís-Marín et al., 2016b). Dif-

ferences in the numbers of species among pre-

vious studies and the present one is attributed

to a more intense literature review, the valida-

tion of geographical coordinates, the correction

of records, and the addition of new ones.

TABLE 3

Number of species of echinoderms from the Central Mexican Pacific

NAY JAL COL MICH MAR ISA MAT REV

Crinoidea 0 0 2 0 1 0 0 0

Asteroidea 13 10 13 11 12 8 6 19

Ophiuroidea 33 27 18 12 39 13 16 25

Echinoidea 7 12 9 10 14 12 7 21

Holothuroidea 21 35 13 30 15 11 19 20

TOTAL 74 84 55 63 81 44 48 85

COL = Colima; ISA = Isabel Island; JAL = Jalisco; MAR = Marías Islands; MAT = Marietas Islands; MICH = Michoacán;

NAY = Nayarit; REV = Revillagigedo Islands.

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JAL, MAR and REV were the areas with

the highest number of species and have been

prospected in several shallow and deep-water

localities/stations since late 1800´s by foreign

(Albatross, Velero, and Zaca; e.g. Agassiz,

1898; Clark, 1940; Deichmann, 1958) and

national expeditions (e.g. Caso, 1943b; Caso,

1961; Caso, 1962b). In contrast, the state

of COL, as well as ISA and MAT islands

have the lowest number of species, and their

poor prospections initiated in the late 1900´s

(e.g. Caso, 1961; Caso, 1980), and have been

mostly performed during the 2000´s, strictly

in shallow-waters (e.g. Chávez-Dagostino et

al., 2000; Ríos-Jara et al., 2008a, Ríos-Jara

et al., 2013; Sotelo-Casas et al., 2018). On

the other hand, MAR and REV (the unique

oceanic islands of the CMP) display unique

characteristics such as particular oceanographic

and topographic conditions, high productiv-

ity, mesophotic reefs, great depths, among

others, which promote high levels of general

marine biodiversity and endemism (CONANP,

2007a; CONANP, 2019), explaining their high-

est number of echinoderms in the region. In

conclusion, the obtained data shows that the

number of species of echinoderms in the CMP

region is mostly associated with a sampling

effort, yet specific conditions of particular

areas (i.e. MAR and REV) can be influencing

their high biodiversity, but more collections of

material and further biogeographic analysis are

required in order to sustain this statement.

Final considerations: The CMP is located

in an oceanographic transitional zone, therefore

fauna from the North and South regions of the

Mexican Pacific (Arriaga-Cabrera et al., 1998),

the Eastern Pacific (e.g. Solís-Marín et al.,

2013a), and even of higher latitudes (e.g. Lam-

bert & Austin, 2007), can inhabit in the region.

In this sense, MAR and REV are important as

they are considered a stepping stone to import

organisms from the Indo-Pacific species into

the Tropical Eastern Pacific (Reyes-Bonilla &

López-Pérez, 1998), as well as in the connec-

tion for dispersal species to and from northern

(Gulf of California) and the mainland CMP

(López-Pérez et al., 2015).

All the islands considered in the pres-

ent work possess their own programs of

management and conservation (CONANP,

2005; CONANP, 2007a; CONANP, 2007b;

CONANP, 2019) in which detailed descrip-

tions of geographical, physical, demographi-

cal, social and environmental characteristics

(among others) are provided. Despite the above,

we detected that all these programs under-

estimated the number of species of echino-

derms since they reported 16 species for ISAB

(CONANP, 2005), 24 for MAR (CONANP,

2007a), 25 for MAT (CONANP, 2007b), and 52

for REV (CONANP, 2019). The inclusion of all

the species reported in the present work on the

lists of these natural protected areas will further

emphasize the importance of the islands.

Due to the importance of the islands of

the CMP, most of the research of echinoderms

in the region have been conducted on them

(e.g. Ríos-Jara et al., 2008a; Sotelo-Casas et

al., 2018) relegating the mainland of the states

of NAY, JAL, COL, and MICH, which remain

almost unstudied and which have a high poten-

tial of finding numerous species of echino-

derms since they have complex environments

(extensive bays, coastal lagoons, estuaries,

mangroves and coral reefs; Arriaga-Cabrera

et al., 1998). Regardless the differences on the

diversity of echinoderms and exploration of the

mainland and islands from the CMP, all of them

are important due to 34 species collected in

the region have been described as new species

(five Asteroidea, 16 Ophiuroidea, five Echinoi-

dea, and eight Holothuroidea; Table 4). This

highlights the potential of the CMP in finding

more new species. Although the present manu-

script provides the most-completed checklist of

the echinoderms in the CMP, we encourage to

intensify research in shallow and deep-waters

and across different habitats of all the islands

and mainland of the states of the region in order

to better understand its biodiversity and distri-

butional patterns. Finally, the preservation of

the marine fauna of the entire CMP is key in the

maintenance of the entire Mexican and Tropi-

cal Pacific; therefore, we encourage ensuring

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its effective protection through inventories of

marine and coastal species.

Ethical statement: 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

followed all pertinent ethical and legal proce-

dures and requirements. All financial sources

are fully and clearly stated in the acknowledge-

ments section. A signed document has been

filed in the journal archives.

TABLE 4

Described species and subspecies of Echinodermata based on material collected in the Central Mexican Pacific

Class Species Locality Reference

Asteroidea

Ampheraster marianus

Marías Islands Ludwig, 1905

Astropecten regalis

San Blas, Nayarit Gray, 1840

Henricia gracilis

Marías Islands Ludwig, 1905

Henricia seminudus

Revillagigedo Islands Clark, 1916

Psilaster armatus

Marías Islands Ludwig, 1905

Ophiuroidea

Amphichondrius laevis

Tenacatita Bay, Jalisco Ziesenhenne, 1940

Amphiodia sculptilis

Tenacatita Bay, Jalisco Ziesenhenne, 1940

Amphiura serpentina

Marías Islands Lütken & Mortensen, 1899

Astrodia excavata

Marías Islands Lütken & Mortensen, 1899

Dougaloplus notacanthus

Marías Islands Lütken & Mortensen, 1899

Ophiacantha costata

Marías Islands Lütken & Mortensen, 1899

Ophiacantha hirta

Marías Islands Lütken & Mortensen, 1899

Ophiacantha moniliformis

Marías Islands Lütken & Mortensen, 1899

Ophiacantha pyriformis

Revillagigedo Islands Ziesenhenne, 1937

Ophiernus adspersus annectens

Marías Islands Lütken & Mortensen, 1899

Ophiolepis plateia

Tenacatita Bay, Jalisco Ziesenhenne, 1940

Ophiomitra granifera

Marías Islands Lütken & Mortensen, 1899

Ophiomitra partita

Marías Islands Lütken & Mortensen, 1899

Ophiophragmus lonchophorus

Tenacatita Bay, Jalisco Ziesenhenne, 1940

Ophiosphalma variabile

Marías Islands Lütken & Mortensen, 1899

Ophiura (Ophiura) scutellata

Marías Islands Lütken & Mortensen, 1899

Echinoidea

Brisaster townsendi

Marías Islands Agassiz, 1898

Echinometra insularis

Revillagigedo Islands Clark, 1912

Encope micropora insularis

Revillagigedo Islands Clark, 1948

Hesperocidaris perplexa

Revillagigedo Islands Clark, 1907

Plexechinus cinctus

Marías Islands Agassiz, 1898

Holothuroidea

Cucumaria flamma

Colima, Nayarit, Marietas Islands Solís-Marín & Laguarda-

Figueras, 1999

Lissothuria imbricata

Jalisco Solís-Marín et al., 2018

Holothuria (Selenkothuria) carere

Michoacán Honey-Escandón et al., 2011

Holothuria (Selenkothuria)

portovallartensis

Puerto Vallarta, Jalisco Caso, 1954

Psolus pauper

Marías Islands Ludwig, 1894

Thyone bidentata

Tenacatita Bay, Jalisco Deichmann, 1941

Thyone parafusus

Tenacatita Bay, Jalisco Deichmann, 1941

Ypsilothuria bitentaculata

Marías Islands Ludwig, 1894

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ACKNOWLEDGMENTS

We are very grateful to Alicia Durán,

Alejandra Martínez-Melo, and Pedro Medina

for providing valuable literature. We wish to

express our sincere gratitude to Alicia Durán

(ICML), Penny Benson and Adam Baldinger

(MCZ), Gordon Hendler and Cathy Groves

(LACM) for providing their support during

collection´s visits. This work was supported

by a scholarship from CONACYT 336853,

and an Ernst Mayr Grant (MCZ), and is part

of the post-doc of RGF at UdG supported by

PRODEP (511-6/2019.-12278). We thank four

anonymous reviewers and editor for comment-

ing and improving the manuscript.

RESUMEN

Equinodermos (Echinodermata) del

Pacífico Central mexicano

Introducción: Los equinodermos del Pacífico Cen-

tral mexicano poseen un alto interés científico, pero ante-

rior al presente trabajo existía un vacío en la información

básica como listados de especies incompletos con inconsis-

tencias sistemáticas y de distribución espacial. Objetivo:

Proporcionar una revisión histórica y un listado de especies

actualizado con la riqueza completa de equinodermos para

cada estado e islas de la región. Métodos: Se elaboró un

listado de especies basado en una búsqueda exhaustiva

de Echinodermata, y fue complementada con la revisión

taxonómica de Ophiuroidea de colecciones científicas.

Todas las coordenadas geográficas de los registros fueron

validadas. Resultados: La región posee 187 especies de

Echinodermata: tres Crinoidea, 35 Asteroidea, 67 Ophiu-

roidea, 32 Echinoidea, y 50 Holothuroidea. Detectamos

52 registros en la literatura considerados como inválidos

y cinco como dudosos. Proveemos 16 nuevos registros de

Ophiuroidea para diversos estados e islas; de estos, cuatro

son nuevos registros para la región. Jalisco presentó el

mayor número de especies (84), seguido por las costas

de Nayarit (74), Michoacán (63), y Colima (55); entre las

islas, Revillagigedo presentó el mayor número de especies

(85) seguido por Marías (81), Marietas (48), e Isabel (44).

Conclusiones: Los números de especies de la región están

mayormente relacionados tanto con el esfuerzo de mues-

treo como con características ambientales que promueven

una alta biodiversidad. El Pacífico Central mexicano es

una región oceanográfica con condiciones mixtas del norte

y sur del Pacífico mexicano, y por lo tanto con una impor-

tancia biogeográfica reflejada en la riqueza de la región.

Palabras clave: Echinodermata; biodiversidad; listado de

especies; revisión histórica; nuevos registros.

REFERENCES

Numbers in brackets refer to literature where the taxa were

recorded (Table 1)

Adem, J., Cobo, E., Blásquez, L., Miranda, F., Villalobos,

A., Herrera, T., . . . Vázquez, L. (1960). La Isla Soco-

rro. Archipiélago de Revillagigedo. Monografías del

Instituto de Geofísica. Universidad Nacional Autóno-

ma de México, 2, 1-234. [1]

Agassiz, A. (1872). Revision of the Echini. Part I. Illustra-

ted catalogue of the Museum of Comparative Zoölogy

at Harvard College, 7, 1-242. [2]

Agassiz, A. (1898). Reports on the dredging operations off

the West coast of Central America to the Galapagos,

to the West coast of Mexico, and in the Gulf of Cali-

fornia, in charge of Alexander Agassiz, carried on

by the U.S. Fish Commission Steamer “Albatross”,

during 1891, Lieut. Commander Z.L. Tanner, U.S.N.,

commanding. XXIII. Preliminary report on the Echi-

ni. Bulletin of the Museum of Comparative Zoölogy at

Harvard College, 32(5), 71-86. [3]

Agassiz, A. (1904). Reports on an exploration off the West

coast of Mexico, Central and South America and off

the Galapagos Islands, in charge of Alexander Agas-

siz by the U.S. Fish Commission Steamer “Alba-

tross”, during 1891, lieut. Commander Z.L. Tanner,

U.S.N commanding. XXXII. The Panamic deep sea

Echini. Memoirs of the Museum of Comparative Zoo-

logy at Harvard College, 31, 1-243. [4]

Alvarado, J.J., Reyes-Bonilla, H., & Benítez-Villalobos, F.

(2015). Diadema mexicanum, erizo de mar clave en

los arrecifes coralinos del Pacífico Tropical Oriental:

lo que sabemos y perspectivas futuras (Diadema-

toida: Diadematidae). Revista de Biología Tropical,

63(2), 135-157. [5]

Arriaga-Cabrera, L., Vázquez-Domínguez, E., González-

Cano, J., Jiménez-Rosenberg, R., Muñoz-López, E.,

& Aguilar-Sierra, V. (1998). Regiones marinas prio-

ritarias de México. México: Comisión Nacional para

el Conocimiento y uso de la Biodiversidad.

Ayala-Bocos, A., Reyes-Bonilla, H., Herrero-Pérezrul,

M.D., Walther-Mendoza, M., & Castañeda-Fernán-

dez de Lara, V. (2011). New records and range

extensions of Astrodictyum panamense (Ophiuroidea:

Gorgonocephalidae) in the eastern Pacific Ocean.

Marine Biodiversity Records, 4(46), 1-3. [6]

Bautista-Romero, J., Reyes-Bonilla, H., Lluch-Cota, D.B.,

& Lluch-Cota S.E (1994). Aspectos generales de la

fauna marina. In A. Ortega-Rubio & A. Castellanos-

Vera (Eds.), La Isla Socorro, Reserva de la Biós-

fera Archipiélago de Revillagigedo, México (pp.

247-275). La Paz, México: Centro de Investigaciones

Biológicas del Noroeste S.C. [7]

246

Revista de Biología Tropical, ISSN electrónico: 2215-2075 Vol. 69(S1): 219-253, March 2021 (Published Mar. 30, 2021)

Borrero-Pérez, G.H., & Vanegas-González, M.J. (2019).

Holothuria (Mertensiothuria) viridiaurantia sp. nov.

(Holothuriida, Holothuriidae), a new sea cucumber

from the Eastern Pacific Ocean revealed by morpho-

logy and DNA barcoding. ZooKeys, 893, 1-19.

Caso, M.E. (1943). Contribución al conocimiento de los

astéridos de México (Master’s thesis). Universidad

Nacional Autónoma de México, México. [8]

Caso, M.E. (1948a). Contribución al conocimiento de los

equinoideos de México. II. Algunas especies de equi-

noideos litorales. Anales del Instituto de Biología,

Universidad Nacional Autónoma de México, 19(1),

183-231. [9]

Caso, M.E. (1948b). Datos históricos y estado actual de los

conocimientos de la fauna de astéridos de México.

Sociedad Mexicana de Historia Natural, 1, 21-32.

[10]

Caso, M.E. (1951). Contribución al conocimiento de los

ofiuroideos de México. I. Algunas especies de ofiu-

roideos litorales. Anales del Instituto de Biología,

Universidad Nacional Autónoma de México, 22(1),

219-312. [11]

Caso, M.E. (1954). Contribución al conocimiento de los

holoturoideos de México. Algunas especies de holo-

turoideos litorales y descripción de una nueva especie

Holothuria portovallartensis. Anales del Instituto de

Biología, Universidad Nacional Autónoma de Méxi-

co, 25, 417-422. [12]

Caso, M.E. (1961). Los Equinodermos de México (Doc-

toral thesis). Universidad Nacional Autónoma de

México, México. [13]

Caso, M.E. (1962a). Estudios sobre Astéridos de México.

Observaciones sobre especies pacíficas del género

Acanthaster y descripción de una subespecie nueva,

Acanthaster ellisii pseudoplanci. Anales del Instituto

de Biología, Universidad Nacional Autónoma de

México, 32(1-2), 313-331. [14]

Caso, M.E. (1962b). Estudios sobre equinodermos de

México. Contribución al conocimiento de los equino-

dermos de las Islas Revillagigedo. Anales del Institu-

to de Biología, Universidad Nacional Autónoma de

México, 33(1-2), 293-330. [15]

Caso, M.E. (1965). Estudios sobre equinodermos de Méxi-

co. Contribución al conocimiento de los holoturoi-

deos de Zihuatanejo y de la Isla de Ixtapa (primera

parte). Anales del Instituto de Biología, Universidad

Nacional Autónoma de México, 36, 253-291. [16]

Caso, M.E. (1966). Contribución al estudio de los holo-

turoideos de México. Morfología interna y ecología

de Stichopus fuscus Ludwig. Anales del Instituto de

Biología, Universidad Nacional Autónoma de Méxi-

co, 37, 175-182. [17]

Caso, M.E. (1974a). Contribución al estudio de los equinoi-

deos de México. Morfología de Tripneustes depressus

Agassiz y estudio comparativo entre T. ventricosus

y T. depressus. Anales del Instituto de Biología,

Universidad Nacional Autónoma de México, 1(1),

25-40. [18]

Caso, M.E. (1974b). Morfología externa de Acanthaster

planci (Linnaeus). Symposium on Indian Ocean and

Adjacent Seas. Journal of the Marine Biological

Association of India, 16(1), 83-93. [19]

Caso, M.E. (1978a). Ciencia y técnica de los equinodermos

en relación con el hombre. Primera parte. Aspecto

científico. Anales del Centro de Ciencias del Mar y

Limnolología, Universidad Nacional Autónoma de

México, 5(1), 255-286. [20]

Caso, M.E. (1978b). Los equinoideos del Pacífico de Méxi-

co. Parte primera. Órdenes Cidaroidea y Aulodonta.

Parte segunda. Órdenes Stiridonta y Camarodonta.

Publicaciones Especiales, Centro de Ciencias del

Mar y Limnología, Universidad Nacional Autónoma

de México, 1, 1-244. [21]

Caso, M.E. (1979a). Los equinodermos de la Bahía de

Mazatlán, Sinaloa. Anales del Centro de Ciencias del

Mar y Limnología, Universidad Nacional Autónoma

de México, Publicación Especial, 6(1), 197-368. [22]

Caso, M.E. (1979b). Los equinodermos (Asteroidea,

Ophiuroidea y Echinoidea) de la Laguna de Térmi-

nos, Campeche. Centro de Ciencias del Mar y Lim-

nología, Universidad Nacional Autónoma de México,

Publicación Especial, 3, 1-186. [23]

Caso, M.E. (1980). Los equinoideos del Pacífico de

México. Parte tercera. Orden Clypeasteroida. Publi-

caciones Especiales, Centro de Ciencias del Mar

y Limnología, Universidad Nacional Autónoma de

México, 4, 1-252. [24]

Caso, M.E. (1983a). Especies del género Amphichondrius

de la costa del Pacífico Americano. Descripción

de una nueva especie del género Amphichondrius.

Amphichondrius unamexici sp. nov. (Ophiuroidea).

Anales del Instituto de Ciencias del Mar y Limno-

logía, Universidad Nacional Autónoma de México,

10(1), 209-230. [25]

Caso, M.E. (1983b). Los equinoideos del Pacífico de Méxi-

co. Parte cuarta. Órdenes Cassiduloida y Spatangoi-

da. Publicaciones Especiales, Centro de Ciencias del

Mar y Limnología, Universidad Nacional Autónoma

de México, 6, 1-200. [26]

Caso, M.E. (1986a). Descripción de una nueva especie de

ofiuroideo de la Bahía de Mazatlán, Sin. Ophioderma

sodipallaresi sp. nov. y comparación con Ophioder-

ma variegatum Lütken. Anales del Instituto de Cien-

cias del Mar y Limnología, Universidad Nacional

Autónoma de México, 13(2), 223-248. [27]

247

Revista de Biología Tropical, ISSN electrónico: 2215-2075, Vol. 69(S1): 219-253, March 2021 (Published Mar. 30, 2021)

Caso, M.E. (1986b). Los equinodermos del Golfo de

California colectados en las campañas SIPCO I-II-III

a bordo del B/O “El Puma”. Anales del Instituto de

Ciencias del Mar y Limnología, Universidad Nacio-

nal Autónoma de México, 13(1), 91-184. [28]

Caso, M.E. (1992). Los equinodermos (Asteroideos, ofiu-

roideos y equinoideos) de la Bahía de Mazatlán,

Sinaloa. Anales del Centro de Ciencias del Mar y

Limnología, Universidad Nacional Autónoma de

México, Publicación Especial, 11, 1-214. [29]

Caso, M.E. (1994). Estudio morfológico, taxonómico,

ecológico y distribución geográfica de los asteroideos

colectados durante las campañas oceanográficas Cor-

tés 1, 2, 3. Anales del Instituto de Ciencias del Mar

y Limnología, Universidad Nacional Autónoma de

México, Publicación Especial, 12, 1-111. [30]

Chacón-Torres, A., Villaroel-Melo, M., Arceo-Zataray,

M., Carrillo-Orea, M.P., Corona-García J.M, Cha-

varría-Guillén, M.P., . . . Villegas-Sandoval, R.E.

(1993). Reconocimiento ecológico de la Isla Socorro,

Archipiélago Revillagigedo, Colima, México. Revista

Trimestral de Ciencia, Arte y Cultura, Universidad

Michoacana, 9, 11-30. [31]

Chan, G.L. (1974). Report on biological observations

of the Revillagigedos expeditions. Kentfield, USA:

NAUI Bio-marine Exploration Seminar, College of

Marin. [32]

Chávez-Dagostino, R.M., Nepote-González, A.C., Solís-

Marín, F.A., & Medina-Rosas, P. (2000). Listado

preliminar sobre equinoideos y asteroideos (Echi-

nodermata: Echinoidea y Asteroidea) de las Islas

Marietas, Nayarit, México. Mexico, 2(1), 69-72. [33]

Clark, A.H. (1916). Six new starfishes from the Gulf of

California and adjacent waters. Proceedings of the

Biological Society of Washington, 29, 51-62. [34]

Clark A.M. (1970). Notes on the family Amphiuridae

(Ophiuroidea). Bulletin of the British Museum (Natu-

ral History) Zoology, 19(1), 1-81. [35]

Clark, H.L. (1907). The Cidaridae. Bulletin of the Museum

of Comparative Zoölogy, 51(7), 165-230. [36]

Clark, H.L. (1912). Hawaiian and other Pacific Echini.

The Pedinidae, Phymosomatidae, Stomopneustidae,

Echinidae, Temnopleuridae, Strongylocentrotidae,

and Echinometridae. Memoirs of the Museum of

Comparative Zoölogy at Harvard College, 34(4),

209-383. [37]

Clark, H.L. (1915). Catalogue of recent ophiurans: based

on the collection of the Museum of Comparative

Zoölogy. Memoirs of the Museum of Comparative

Zoölogy at Harvard College, 25(4), 164-376. [38]

Clark, H.L. (1925). A catalogue of the recent sea-urchins

(Echinoidea) in the collection of the British Museum

(Natural History). London, United Kingdom: The

British Museum (Natural History).

Clark, H.L. (1935). Some new echinoderms from Califor-

nia. The Annals and Magazine of Natural History,

10(15), 120-129.

Clark, H.L. (1940). XXI. Notes on Echinoderms from

the West Coast of Central America. Eastern Pacific

Expeditions of the New York Zoological Society.

Zoologica, 25(22), 331-352. [39]

Clark, H.L. (1948). A report on the Echini of the warmer

Eastern Pacific, based on the collections of the Velero

III. Allan Hancock Pacific Expeditions, 8(5), 225-

360. [40]

CONANP. (2004). Programa de conservación y manejo

Reserva de la Biosfera Archipiélago de Revillagige-

do, México. Ciudad de México, México: Comisión

Nacional de Áreas Naturales Protegidas, Secretaría

de Medio Ambiente y Recursos Naturales. [41]

CONANP. (2005). Programa de conservación y manejo

del Parque Nacional Isla Isabel, México. Ciudad

de México, México: Comisión Nacional de Áreas

Naturales Protegidas, Secretaría de Medio Ambiente

y Recursos Naturales. [42]

CONANP. (2007a). Programa de conservación y manejo,

Reserva de la Biosfera Islas Marías, México. Ciudad

de México, México: Comisión Nacional de Áreas

Naturales Protegidas, Secretaría de Medio Ambiente

y Recursos Naturales. [43]

CONANP. (2007b). Programa de conservación y manejo,

Parque Nacional Islas Marietas, México. Ciudad

de México, México: Comisión Nacional de Áreas

Naturales Protegidas, Secretaría de Medio Ambiente

y Recursos Naturales. [44]

CONANP. (2019). Programa de manejo, Parque Nacional

Revillagigedo. Ciudad de México, México: Comisión

Nacional de Áreas Naturales Protegidas, Secretaría

de Medio Ambiente y Recursos Naturales. [45]

Deichmann, E. (1937). The Templeton Crocker Expedition.

IX. Holothurians from the Gulf of California, the

West Coast of Lower California and Clarion Island.

Zoologica, 22, 161-176. [46]

Deichmann, E. (1941). The Holothuroidea collected by

the Velero III during the years 1932 to 1938. Part I,

Dendrochirota. Allan Hancock Pacific Expeditions,

8, 61-195. [47]

Deichmann, E. (1958). The Holothuroidea collected by the

Velero III and IV during the years 1932 to 1954. Part

248

Revista de Biología Tropical, ISSN electrónico: 2215-2075 Vol. 69(S1): 219-253, March 2021 (Published Mar. 30, 2021)

II Aspidochirota. Allan Hancock Pacific Expeditions,

11(2), 253-348. [48]

Devaney, D.M. (1970). Studies on Ophiocomid brittlestars.

I. A new genus (Clarkcoma) of Ophiocominae with

a reevaluation of the genus Ophiocoma. Smithsonian

Contributions to Zoology, 51, 1-41. [49]

Döderlein, L. (1911). Beiträge zur Naturgeschichte Osta-

siens. Über Japanische und andere Euryalae. Abhand-

lugen der Bayerischen Akademie der Wissencaften.

Mathemaatisch-Naturwissenschaftliche Abteilung,

2(5), 1-123. [50]

Downey, M.E. (1968). Catalog of recent Echinoid type

specimens in the U.S. National Museum Smithsonian

Institution and the Museum of Comparative Zoology

Harvard University. United States National Museum

Bulletin, 264, 1-99. [51]

Downey, M.E. (1969). Catalog of recent ophiuroid type

specimens in major collections in the United States.

United States National Museum Bulletin, 293, 1-239.

[52]

Espino-Barr, E., Cibrián-Rodríguez, R., & García-Boa,

A. (1996). Estructura y densidad de la población del

erizo tinta Diadema mexicanum en el litoral rocoso

del estado de Colima. Ciencia Pesquera, 12, 60-67.

[53]

Fisher, W.K. (1911). Asteroidea of the North Pacific and

adjacent waters. Part 1. Phanerozonia and Spinulo-

sa. Smithsonian Institution, United States National

Museum Bulletin, 76, 1-419. [54]

Fisher, W.K. (1928a). Asteroidea of the North Pacific and

adjacent waters. Part 2. Forcipulata (part). Smith-

sonian Institution, United States National Museum

Bulletin, 76, 1-245. [55]

Fisher, W.K. (1928b). Asteroidea of the North Pacific

and adjacent waters. Part 3. Forcipulata (conclu-

ded). Smithsonian Institution, United States National

Museum Bulletin, 76, 1-356. [56]

Frontana-Uribe, S.C., Hernández-Alcántara, P., & Solís-

Weiss, V. (2000). Ophiocomella schmitti (Echinoder-

mata: Ophiuroidea) from Socorro Island, Mexican

Pacific. Revista de Biología Tropical, 49(2), 766. [57]

Fuentes-Farías, A.M., Villarroel-Melo, M.D., & Solís-

Marín, F.A. (2005). Los invertebrados marinos.

Anexo 4.20 y 4.21. Listado de especies de equino-

dermos del Estado de Michoacán. In L.E. Villase-

ñor-Gómez (Ed.), La biodiversidad en Michoacán,

Estudio de Estado (pp. 85-88). México: Comisión

Nacional para el Conocimiento y Uso de la Biodiver-

sidad, Secretaría de Urbanismo y Medio Ambiente,

Universidad Michoacana de San Nicolás de Hidalgo.

[58]

García-Hernández, V.C., Reyes-Bonilla, H., Balart, E.F.,

Ríos-Jara, E., Lluch-Cota, S.E., & Serviere-Zarago-

za, E. (2014). Comparison of ecological diversity

and species composition of macroalgae, benthic

macroinvertebrates, and fish assemblages between

two tropical rocky reefs. Revista de Biología Marina

y Oceanografía, 49, 477-491. [59]

Granja-Fernández, R., Antonio-Pérez, V., & López-Pérez,

R. A. (2013a). Euapta godeffroyi (Holothuroidea:

Synaptidae): Filling the distribution gap between

Mexico and Costa Rica, eastern tropical Pacific.

Hidrobiológica, 23(1), 130-132. [60]

Granja-Fernández, R., Hernández-Moreno, P., & Bastida-

Zavala, R. (2013b). First record of the association

between Malmgreniella cf. variegata (Polychaeta,

Polynoidae) and Ophionereis annulata (Echinoder-

mata, Ophionereididae) in the Mexican Pacific. Sym-

biosis, 60, 85-90. [61]

Granja-Fernández, R., Herrero-Pérezrul, M.D., López-

Pérez, R.A., Hernández, L., Rodríguez-Zaragoza,

F.A., Jones, R.W., & Pineda-López, R. (2014).

Ophiuroidea (Echinodermata) from coral reefs in the

Mexican Pacific. Zookeys, 406, 101-145. [62]

Granja-Fernández, R., Herrero-Pérezrul, M.D., López-

Pérez, R.A., Hernández-Morales, A., & Rangel-Solís,

P.D. (2015a). A literature review of the Ophiuroidea

(Echinodermata) from the Pacific coast of Mexico.

Revista de Biología Tropical, 63(2), 37-47. [63]

Granja-Fernández, R., Solís-Marín, F.A., Benítez-Villalo-

bos, F., Herrero-Pérezrul, M.D., & López-Pérez, A.

(2015b). Checklist of echinoderms (Echinodermata)

from the Southern Mexican Pacific: a historical

review. Revista de Biología Tropical, 63(2), 87-114.

Granja-Fernández, R., Rodríguez-Troncoso, A.P., Herrero-

Pérezrul, M.D., Sotelo-Casas, R.C., Flores-Ortega,

J.R., Godínez-Domínguez, E., . . . Cupul-Magaña,

A.L. (2017). Ophiuroidea (Echinodermata) from the

Central Mexican Pacific: an updated checklist inclu-

ding new distribution records. Marine Biodiversity,

47, 167-177. [64]

Grant, U.S., & Hertlein, L.G. (1938). The West American

Cenozoic Echinoidea. Publications of the University

at Los Angeles in Mathematical and Physical Scien-

ces, 2, 1-225. [65]

Gray, J.E. (1840). XXII. A synopsis of the genera and spe-

cies of the class Hypostoma (Asterias, Linnaeus). The

Annals and Magazine of Natural History, 6, 175-184.

[66]

Harold, A.S., & Telford, M. (1990). Systematics, phylo-

geny and biogeography of the genus Mellita (Echi-

noidea: Clypeasteroida). Journal of Natural History,

24(4), 987-1026. [67]

249

Revista de Biología Tropical, ISSN electrónico: 2215-2075, Vol. 69(S1): 219-253, March 2021 (Published Mar. 30, 2021)

Hartlaub C. (1895). Reports on the dredging operations off

the west coast of Central America to the Galapagos,

to the west coast of Mexico, and in the Gulf of Cali-

fornia, in charge of Alexander Agassiz, carried on

by the U. S. Fish Commission steamer “Albatross”,

during 1891, Lieut. Commander Z. L. Tanner, U.

S. N., commanding. 13. Die Comatuliden. Bulletin

of the Museum of Comparative Zoölogy at Harvard

College, 27, 129-152. [68]

Haszprunar, G., & Spies, M. (2014). An integrative

approach to the taxonomy of the crown-of-thorns

starfish species group (Asteroidea: Acanthaster): A

review of names and comparison to recent molecular

data. Zootaxa, 3841, 271-284. [69]

Hendler, G. (1996). Taxonomic atlas of The Benthic Fauna

of the Santa Maria Basin and Western Santa Bar-

bara Channel. Class Ophiuroidea (Vol. 14). Santa

Barbara, USA: Santa Barbara Museum of Natural

History. [70]

Hendler, G. (2011). New light on the nomenclature, taxo-

nomy, and biology of Hemipholis species (Echino-

dermata: Ophiurodea: Ophiactidae). Zootaxa, 3048,

44-52. [71]

Hendler, G., Miller, J.E., Pawson, D.L., & Kier, P.M.

(1995). Sea stars, sea urchins, and allies. Echino-

derms of Florida and the Caribbean. Washington,

USA: Smithsonian Institution.

Hermosillo-Nuñez, B.B., Rodríguez-Zaragoza, F.A., Ortiz,

M., Galván-Villa, C., Cupul-Magaña, A., & Ríos-

Jara, E. (2015). Effect of habitat structure on the most

frequent echinoderm species inhabiting coral reef

communities at Isla Isabel National Park (Mexico).

Community Ecology, 16(1), 125-134. [72]

Hermosillo-Nuñez, B.B., Rodríguez-Zaragoza, F.A., Ortiz,

M., Calderon-Aguilera, L.E., Cupul-Magaña, A.L.

(2016). Influence of the coral reef assemblages on

the spatial distribution of echinoderms in a gradient

of human impacts along the tropical Mexican Pacific.

Biodiversity and Conservation, 25, 2137-2152. [73]

Herrero-Pérezrul, M.D., Reyes-Bonilla, H., González-

Azcárraga A., Cintra-Buenrostro, C.E., & Rojas-

Sierra, A. (2007). Equinodermos. In G.D. Danemann

& E. Ezcurra (Eds.), Bahía de los Ángeles: recursos

naturales y comunidad, Línea base 2007 (pp. 339-

361). México: Secretaría de Medio Ambiente y

Recursos Naturales, Instituto Nacional de Ecología,

Pronatura Noroeste A.C., San Diego Natural History

Museum. [74]

Herrero-Pérezrul, M.D., Ramírez-Ortiz, G., Rosales-

Estrada, M., & Reyes-Bonilla, H. (2015). Densidad

poblacional y distribución espacial de erizos de

mar (Echinodermata: Echinoidea) en Isla Socorro,

Archipiélago de Revillagigedo, México. Revista de

Biología Tropical, 63(2), 221-232. [75]

Herrero-Pérezrul, M.D., Reyes-Bonilla, H., & Granja-Fer-

nández, R. (2017). Effects of environmental factors

on the abundances of the basket stars Astrocaneum

spinosum and Astrodictyum panamense (Ophiuroi-

dea: Gorgonocephalidae) in the northern Gulf of

California, Mexico. Marine Biology Research, 13(2),

210-219. [76]

Hertlein, L.G. (1963). Contribution to the biogeography of

Cocos Island, including a bibliography. Proceedings

of the California Academy of Sciences, 32(8), 219-

289. [77]

Hickman, C.P. (1998). A field guide to sea stars and other

echinoderms of Galápagos. Virginia, USA: Sugar

Spring Press. [78]

Holguín-Quiñones, O.E. (1994). Comunidades bentónicas

marinas. In A. Ortega-Rubio & A. Castellanos-Vera

(Eds.), La Isla Socorro, Reserva de la Biósfera Archi-

piélago de Revillagigedo, México (pp. 225-245). La

Paz, México: Centro de Investigaciones Biológicas

del Noroeste, S.C. [79]

Holguin-Quiñones, O.E., & Michel-Morfín, J.E. (2006).

Population structure and accompanying biota of the

snail Turbo (Callopoma) funiculosus (Gastropoda:

Turbinidae), on Socorro Island, Revillagigedo Archi-

pielago, Mexico. Revista de Biología Tropical, 54(4),

1079-1084. [80]

Honey-Escandón, M., Solís-Marín F.A., & Laguarda-

Figueras A. (2008). Equinodermos (Echinodermata)

del Pacífico Mexicano. Revista de Biología Tropical,

56(3), 57-73. [81]

Honey-Escandón M., Solís-Marín, F.A., & Laguarda-

Figueras, A. (2011). Holothuria (Selenkothuria) care-

re, a new species of sea cucumber (Echinodermata:

Holothuroidea) from the Mexican Pacific. Zootaxa,

2922, 27-33. [82]

Koehler, R. (1907). Révision de la Collection des Ophiures

du Museum D´Histoire Naturelle de Paris. Bulletin

Scientifique de la France et de la Belgique, 61, 279-

370. [83]

Kroh, A., & Mooi, R. (2020). World Echinoidea Database.

Retrieved from http://www.marinespecies.org/echi-

noidea on 2020-06-13

Lambert, P., & Austin, W. (2007). Brittle stars, sea urchins

and feather stars of British Columbia, Southeast

Alaska and Puget Sound. Victoria, Canada: Royal BC

Museum Handbook.

López-Pérez, A., Cupul-Magaña, A., Ahumada-Sempoal,

M., Medina-Rosas, P., Reyes-Bonilla, H., Herrero-

Pérezrul, M.D., . . . Lara-Hernández, J. (2015). The

coral communities of the Islas Marias archipelago,

Mexico: structure and biogeographic relevance to the

Eastern Pacific. Marine Ecology, 37, 679-690.

250

Revista de Biología Tropical, ISSN electrónico: 2215-2075 Vol. 69(S1): 219-253, March 2021 (Published Mar. 30, 2021)

López-Uriarte, E., Ríos-Jara, E., Galván-Villa, C.M., Juá-

rez-Carrillo, E., Enciso-Padilla, I., Robles-Jarero,

E.G., & Pérez-Peña, M. (2009). Macroinvertebrados

bénticos del litoral somero de Punta La Rosada,

Bahía Chamela, Jalisco. Scientia-CUCBA, 11(1-2),

57-68. [84]

Ludwig, H. (1893). Vorläufiger Bericht über die erbeuteten

Holothurien. Reports on the Dredging Operations off

the West Coast of Central America to the Galapagos,

etc., by the U. S. Fish Commission Steamer “Alba-

tross”. IV. Bulletin of the Museum of Comparative

Zoology at Harvard College, 24(4), 105-114.

Ludwig, H. (1894). Reports on an exploration off the west

coasts of Mexico, central and South America, and

off the Galapagos Islands, in charge of Alexander

Agassiz, by the U.S. Fish Commission Steamer

“Albatross” during 1891, Lieut. Commander Z.L.

Tanner, U.S.N., commanding. XII. The Holothurioi-

dea. Memoirs of the Museum of Comparative Zoölogy

at Harvard College, 17(3), 1-183. [85]

Ludwig, H. (1905). VII. Asteroidea. Reports on an explora-

tion off the West Coast of Mexico, Central and South

America, and off the Galapagos islands in charge of

Alexander Agassiz, by the U.S. Fish Commission

steamer “Albatross”, during 1891, lieut. Commander

Z.L. Tanner, U.Sn., commanding. XXXV. Reports

on the scientific results of the expedition to the Tro-

pical Pacific, in charge of Alexander Agassiz on the

U.S. Fish Commission Steamer “Albatross”, from

August 1899, to March, 1900, commander Jefferson

F. Moser, U.S.N., commanding. Memoirs of the

Museum of Comparative Zoölogy at Harvard Colle-

ge, 32, 1-292. [86]

Luke, S.R. (1982). Catalog of the benthic invertebrate

collections of the SCRIPPS Institution of Oceano-

graphy, Echinodermata. California, USA: SCRIPPS

Institution of Oceanography Reference Series, no.

82-5, University of California. [87]

Lütken, C.F., & Mortensen, T. (1899). The Ophiuridae.

Memoirs of the Museum of Comparative Zoology,

23, 93-208. [88]

Madsen, F.J. (1977). The Ophioleucidae (Ophiuroidea).

Galathea Report. Scientific Results of the Danish

Deep-Sea Expedition Round the world 1950-52, 14,

109-122. [89]

Mah, C.L. (2020). World Asteroidea Database. Retrieved

from http://www.marinespecies.org/asteroidea

Maluf, L.Y. (1988). Composition and distribution of the

Central Eastern Pacific echinoderms. Natural History

Museum of Los Angeles County. Technical Report, 2,

1-242. [90]

Maluf, L.Y. (1991). Echinoderm fauna of the Galápagos

Islands. In M.J. James (Ed.), Galápagos marine

invertebrates: taxonomy, biogeography, and evolu-

tion in Darwin’s Islands (pp. 345-367). New York,

USA: Plenum press. [91]

Maluf, L.Y., & Brusca, R.C. (2005). Echinodermata. In

M.E. Hendrickx, R.C. Brusca, & L.T. Findley (Eds.),

A distributional checklist of the macrofauna of the

Gulf of California, Mexico. Part I. Invertebrates

(pp. 327-343). Tucson, USA: Arizona-Sonora Desert

Museum. [92]

Martínez-Melo, A., Solís-Marín, F.A., Buitrón-Sánchez,

B.E., & Laguarda-Figueras, A. (2015). Taxonomía y

biogeografía ecológica de los equinoideos irregulares

(Echinoidea: Irregularia) de México. Revista de Bio-

logía Tropical, 63(2), 59-75. [93]

Massin, C., & Hendrickx, M.E. (2011). Deep-water

Holothuroidea (Echinodermata) collected during the

TALUD cruises off the Pacific coast of Mexico, with

the description of two new species. Revista Mexicana

de Biodiversidad, 82, 413-443. [94]

McCartney, M.A., Keller, G., & Lessios, H.A. (2000).

Dispersal barriers in tropical oceans and speciation in

Atlantic and eastern Pacific sea urchins of the genus

Echinometra. Molecular Ecology, 9, 1391-1400. [95]

Messing, C. (2020). World List of Crinoidea. Crinoidea.

Retrieved from http://www.marinespecies.org/aphia.

php?p=taxdetails&id=123081 on 2020-06-13

Miller, A.K., Kerr, A.M., Paulay, G., Reich, M., Wilson,

N.G., Carvajal, J.I., & Rouse, G.W. (2017). Mole-

cular phylogeny of extant Holothuroidea (Echino-

dermata). Molecular Phylogenetics and Evolution,

111, 110-131.

Miranda, A., Ambriz, G., Valencia, D., Sánchez, M., &

Szekely, A. (2011). Programa de Manejo del san-

tuario de las Islas de la Bahía de Chamela: Islas La

Pajarera, Cocinas, Mamut, Colorada, San Pedro,

San Agustín, San Andrés y Negrita, e islotes Los Ane-

gados, Novillas, Mosca y Submarino, Jalisco, Méxi-

co. Ciudad de México, México: Comisión Nacional

de Áreas Naturales Protegidas. [96]

Mooi, R., Constable H., Lockhart, S., & Pearse, J. (2004).

Echinothurioid phylogeny and the phylogenetic sig-

nificance of Kamptosoma (Echinoidea: Echinoder-

mata). Deep-Sea Research II, 51, 1903-1919.

Mortensen, T. (1928). A monograph of the Echinoidea I.

Cidaroidea. Copenhagen, Denmark: C.A. Rietzel

publisher, Oxford University press. [97]

Mortensen, T. (1940). A monograph of the Echinoidea. III.

1. Aulodonta. With additions to Vol. II (Lepidocen-

troida and Stirodonta). Copenhagen, Denmark: C.A.

Reitzel publisher. [98]

251

Revista de Biología Tropical, ISSN electrónico: 2215-2075, Vol. 69(S1): 219-253, March 2021 (Published Mar. 30, 2021)

Mortensen, T. (1943a). A monograph of the Echinoidea. III.

2. Camarodonta. I. Orthopsidae, Glyphocyphidae,

Temnopleuridae and Toxopneustidae. Copenhagen,

Denmark: C.A. Reitzel publisher. [99]

Mortensen, T. (1943b). A monograph of the Echinoidea.

III. 3. Camarodonta. II. Echinidae, Strongylocentro-

tidae, Parasaleniidae, Echinometridae. Copenhagen,

Denmark: C.A. Reitzel publisher. [100]

Mortensen, T. (1948a). A monograph of the Echinoidea.

IV. 1. Holectypoida, Cassiduloida. Copenhagen, Den-

mark: C.A. Reitzel publisher. [101]

Mortensen, T. (1948b). A monograph of the Echinoidea.

IV. 2. Clypeastroida. Clypeastridae, Arachnoididae,

Fibulariidae, Laganidae and Scutellidae. Copenha-

gen, Denmark: C.A. Reitzel publisher. [102]

Mortensen, T. (1951). A monograph of the Echinoidea. V.

2. Spatangoida II. Amphisternata, II. Spatangidae,

Loveniidae, Pericosmidae, Schizasteridae, Brissidae.

Copenhagen, Denmark: C.A. Reitzel publisher. [103]

Nava-Bravo, H.H., Solís-Marín, F.A., Fuentes-Farías, A.L.,

Cruz-Barraza, J.A., Bastida-Zavala, J.R., Laguarda-

Figueras, A., … Conejeros-Vargas, C.A. (2019).

Invertebrados marinos. Apéndice 35. In A. Cruz-

Angón, K.C. Nájera-Cordero, & E.D. Melgarejo

(Coords.), La biodiversidad en Michoacán. Estudio

de Estado 2, vol. II (pp. 291-306). México: Comisión

Nacional para el Conocimiento y Uso de la Biodiver-

sidad. [104]

Nepote-González, A.C. (1998). Holoturias (Echinoderma-

ta: Holothuroidea) de las islas Marietas, Bahía de

Banderas, Jalisco-Nayarit, México (Bachelor thesis).

Universidad de Guadalajara, México. [105]

O´Hara, T.D., Stöhr, S., Hugall, A.F., Thuy, B., & Marty-

nov, A. (2018). Morphological diagnoses of higher

taxa in Ophiuroidea (Echinodermata) in support of a

new classification. European Journal of Taxonomy,

416, 1-35.

Okanishi, M., & Fujita, T. (2014). A taxonomic review of

the genus Astrodia (Echinodermata: Ophiuroidea:

Asteronychidae). Journal of the Marine Biological

Association of the United Kingdom, 94(1), 187-201.

[106]

Padilla-Pérez, M.S., Rodríguez-Troncoso A.P., Sotelo-

Casas R.C., & Cupul-Magaña A.L. (2017). Equi-

nodermos del Parque Nacional Islas Marietas:

Generalidades, importancia e identificación visual

como herramienta para su protección. Áreas Natura-

les Protegidas Scripta, 3(2), 51-92. [107]

Parker, R.H. 1963. Zoogeography and ecology of some

macro-invertebrates, particularly mollusks, in the

Gulf of California and continental slope off Mexico.

Massachusetts, USA: Woods Hole.

Pawson, D. (1983). Psychronaetes hanseni, a new genus

and species of Elasipodan sea cucumber from the

Eastern Central Pacific (Echinodermata: Holothu-

roidea). Proceedings of the Biological Society of

Washington, 96(1), 154-159.

Reyes-Bonilla, H. (1995). Asteroidea and Echinoidea

(Echinodermata) of Isla San Benedicto, Revillagi-

gedo Archipelago, México. Revista de Investigación

Científica Sección Ciencias del mar, UABCS, 6(1),

29-38. [108]

Reyes-Bonilla, H., & López-Pérez, A. (1998). Biogeografía

de los corales pétreos (Scleractinia) del Pacífico de

México. Ciencias Marinas, 24(2), 211-224.

Ríos-Jara, E., Galván-Villa, C.M., & Solís-Marín, F.A.

(2008a). Equinodermos del Parque Nacional Isla

Isabel, Nayarit, México. Revista Mexicana de Biodi-

versidad, 79, 131-141. [109]

Ríos-Jara, E., López-Uriarte, E., Pérez-Peña, M., Enciso-

Padilla, I., Arreola-Robles, J.L., Hermosillo-Gon-

zález, A., & Galván-Villa, C.M. (2008b). Listados

taxonómicos de la biota marina del Parque Nacional

Isla Isabel (invertebrados, peces y macroalgas).

Zapopan, México: Universidad de Guadalajara. [110]

Ríos-Jara, E., Galván-Villa, C.M., Rodríguez-Zaragoza,

F.A., López-Uriarte, E., Bastida-Izaguirre, D., &

Solís-Marín, F.A. (2013). Los equinodermos (Echino-

dermata) de bahía Chamela, Jalisco, México. Revista

Mexicana de Biodiversidad, 84, 263-279. [111]

Ríos-Jara, E., Juárez-Carrillo, E., & Galván-Villa, C.M.

(2017) Invertebrados marinos. In CONABIO (Ed.),

La biodiversidad en Jalisco. Estudio de Estado (Vol.

2, pp. 251-269). México: Comisión Nacional para el

Conocimiento y Uso de la Biodiversidad, Secretaría

de Medio Ambiente y Desarrollo Territorial. [112]

Rodríguez-Troncoso, A.P., Sotelo-Casas, R.C., Galván-

Villa, C.M., Godínez-Domínguez, E., Flores-Ortega,

J.R., del Castillo-Fernández, K.M., . . .Solís-Marín,

F.A. (2013). Equinodermos de la Costa Sur de Jalisco

y Colima. In E. Godínez-Domínguez & M.C. Franco-

Gordo (Eds.), Inventario de biodiversidad de la costa

Sur de Jalisco y Colima, (Vol. 2 pp. 46-60). México:

Universidad de Guadalajara. [113]

Roux, M. (2004). New hyocrinid crinoids (Echinodermata)

from submersible investigations in the Pacific Ocean.

Pacific Science, 58(4), 597-613. [114]

Roux, M., & Pawson, D.L. (1999). Two new Pacific Ocean

species of Hyocrinid crinoids (Echinodermata), with

comments on presumed giant-dwarf gradients related

to seamounts and abyssal plains. Pacific Science,

53(3), 289-298.

Rowe, F.W.E., & Pawson, D.L. (1977). A catalogue of echi-

noderm type-specimens in the Australian Museum,

252

Revista de Biología Tropical, ISSN electrónico: 2215-2075 Vol. 69(S1): 219-253, March 2021 (Published Mar. 30, 2021)

Sydney. Records of The Australian Museum, 30(14),

337-364. [115]

Santos-Beltrán, C., & Salazar-Silva, P. (2011). Holotu-

roideos (Echinodermata: Holothuroidea) de playas

rocosas, zona norte de Bahía de Banderas, Nayarit,

México. Ciencia y Mar, 15(45), 3-11. [116]

Sladen, W.P. (1889). Report on the Asteroidea collected by

the H. M. S. Challenger during the years 1873-1876.

The Voyage of H. M. S. Challenger, Zoology, 30,

1-893. [117]

Solís-Marín, F.A., & Laguarda-Figueras, A. (1999). Cucu-

maria flamma, a new species of sea cucumber from

the central eastern Pacific (Echinodermata: Holothu-

roidea). Proceedings of the Biological Society of

Washington, 112(4), 778-786. [118]

Solís-Marín, F.A., Arriaga-Ochoa, J.A., Laguarda-Figue-

ras, A., Frontana-Uribe, S.C., & Durán-González,

A. (2009). Holoturoideos (Echinodermata: Holothu-

roidea) del Golfo de California. México: Comisión

Nacional para el Conocimiento y Uso de la Biodi-

versidad, Instituto de Ciencias del Mar y Limnología,

UNAM. [119]

Solís-Marín, F.A., Alvarado, J.J., Abreu-Pérez, M., Aguile-

ra, O., Alió, J., Bacallado-Aránega, J.J., … Williams,

S.M. (2013a). Appendix. In J.J. Alvarado Barrientos

& F.A. Solís-Marín (Eds.), Echinoderm Research

and diversity in Latin America (pp. 543-658). Berlin,

Germany: Springer-Verlag. [120]

Solís-Marín, F.A., Honey-Escandón, M.B.I., Herrero-

Pérezrul, M.D., Benítez-Villalobos, F., Díaz-Martí-

nez, J.P., Buitrón-Sánchez, B.E., … Durán-González,

A. (2013b). The echinoderms of Mexico: Biodiver-

sity, distribution and current state of knowledge. In

J.J. Alvarado & F.A. Solís-Marín (Eds.), Echinoderm

Research and diversity in Latin America (pp. 11-65).

Berlin, Germany: Springer-Verlag. [121]

Solís-Marín, F.A., Laguarda-Figueras, A., & Honey-Escan-

dón, M. (2014). Biodiversidad de equinodermos

(Echinodermata) en México. Revista Mexicana de

Biodiversidad, 85, S441-S449. [122]

Solís-Marín, F.A., Durán-González, A., & Hernández-

Herrejón, L.A. (2016a). Ofiuros (Ophiuroidea). In

CONABIO (Ed.), La Biodiversidad en Colima. Estu-

dio de Estado (pp. 305-308). México: Comisión

Nacional para el Conocimiento y Uso de la Biodi-

versidad. [123]

Solís-Marín, F.A., Laguarda-Figueras, A., & Durán-Gon-

zález, A. (2016b). Estrellas, erizos y pepinos de

mar (Echinodermata). Apéndice 1. Lista de equino-

dermos. In CONABIO (Ed.), La Biodiversidad en

Colima. Estudio de Estado (pp. 295-304, 203-205).

México: Comisión Nacional para el Conocimiento y

Uso de la Biodiversidad. [124]

Solís-Marín, F.A., Arriaga-Ochoa, J.A., Galván-Villa,

C.M., & Laguarda-Figueras, A. (2018). Lissothuria

imbricata sp. nov., a new species of sea cucumber

(Echinodermata: Holothuroidea) from the central-

eastern Pacific Ocean. Zootaxa, 4438(2), 373-380.

[125]

Sotelo-Casas, R.C., & Rodríguez-Troncoso, A.P. (2014).

Los equinodermos: invertebrados esenciales dentro

de la estructura de la comunidad coralina. In J.L.

Cifuentes-Lemus & F.G. Cupul-Magaña (Coords.),

Temas sobre investigaciones costeras (pp. 56-81).

México: Universidad de Guadalajara. [126]

Sotelo-Casas, R.C., Cupul-Magaña, A.L., Solís-Marín,

F.A., & Rodríguez-Troncoso, A.P. (2015). New

records of sea cucumbers (Echinodermata: Holothu-

roidea) at Islas Marietas, Central Mexican Pacific.

Marine Biological Association of the United King-

dom, 8, 1-8. [127]

Sotelo-Casas, R.C., Cupul-Magaña, A.L., Solís-Marín, F.A.,

& Rodríguez-Troncoso, A.P. (2016). Recruitment pat-

terns of 2 sea cucumber species in a Central Mexican

Pacific coral reef community. Revista Mexicana de

Biodiversidad, 87, 86-91. [128]

Sotelo-Casas, R.C., Cupul-Magaña, A.L., Rodríguez-

Zaragoza, F.A., Solís-Marín, F.A., & Rodríguez-

Troncoso, A.P. (2018). Structural and environmental

effects on an assemblage of echinoderms associated

with a coral community. Marine Biodiversity, 48,

1401-1411. [129]

Spalding, M.D., Fox, H.E., Allen, G.R., Davidson, N.,

Ferdaña, Z.A., Finlayson, M., . . . Robertson, J.

(2007). Marine ecoregions of the world: A biore-

gionalization of coastal and shelf areas. BioScience,

57(7), 573-583.

Steinbeck, J., & Ricketts, E.F. (1941). Sea of Cortez. A

leisurely journal of travel and research. New York,

USA: Viking Press. [130]

Stöhr, S., O´Hara, T., & Thuy, B. (2020). World Ophiuroi-

dea database. Retrieved from http://www.marinespe-

cies.org/ophiuroidea

Tommasi, L.R. (1976). Ophiuroidea collected in the Peru-

Chile trench by the USNS “Eltanin” during Cruise

III. Papéis Avulsos de Zoologia, 29(8), 281-318.

[131]

Valdés de Anda, M., Hendrickx, M.E., & Solís-Marín,

F.A. (2018). New records of Florometra serratissima

(A.H. Clark, 1907) (Echinodermata: Crinoidea) from

the Gulf of California and the Pacific Coast of Mexi-

co. Cashiers de Biologie Marine, 59, 167-173. [132]

Vergara-Chen, C., Guerra, Z., & Collado, G.N. (2015). El

pepino de mar, Isostichopus fuscus, recurso marino

253

Revista de Biología Tropical, ISSN electrónico: 2215-2075, Vol. 69(S1): 219-253, March 2021 (Published Mar. 30, 2021)

en peligro con altas necesidades de manejo. Tecno-

ciencia, 17(2), 21-41. [133]

Villalobos, A.F. (1992). La Isla Socorro. Notas acerca

del aspecto hidrobiológico de la parte sur de la Isla.

(tomando de Monografías del Instituto de Geofísica

2: 155-180). Hidrobiológica, 2(1-2), 65-91. [134]

WoRMS (2020). World Register of Marine Species.

Retrieved from http://www.marinespecies.org/aphia.

php?p=search

Ziesenhenne, F.C. (1937). The Templeton Crocker Expedi-

tion. X. Echinoderms from the West Coast of Lower

California, the Gulf of California and Clarion Island.

Zoologica, New York Zoological Society, 22(15),

209-239. [135]

Ziesenhenne, F.C. (1940). New Ophiurans of the Allan

Hancock Pacific Expeditions. Allan Hancock Pacific

Expeditions, 8(2), 9-52. [136]

Ziesenhenne, F.C. (1955). A review of the genus Ophio-

derma Müller and Troschel. In Essays in the Natural

Sciences in honor of Captain Allan Hancock, on the

occasion of his birthday, July 26, 1955 (pp. 185-201).

Los Angeles, USA: University of Southern California

Press. [137]

See Digital Appendix at: / Ver Apéndice digital en: revistas.ucr.ac.cr