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Taxonomy of family Benthopectinidae (Echinodermata: Asteroidea)

in the Mexican waters of the Gulf of Mexico

Magdalena De los Palos-Peña

https://orcid.org/0000-0002-9667-8670

Francisco Alonso Solís-Marín

; https://orcid.org/0000-0001-5729-3316

Alfredo Laguarda-Figueras

1. Posgrado en Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, México;

mdelospalos@ciencias.unam.mx (*Correspondencia).

2. Laboratorio de Sistemática y Ecología de Equinodermos, Instituto de Ciencias del Mar y Limnología, Universidad

Nacional Autónoma de México, México; fasolis@cmarl.unam.mx ; laguarda@cmarl.unam.mx

Received 17-VI-2020. Corrected 05-VII-2020. Accepted 10-VIII-2020.

ABSTRACT

Introduction: The family Benthopectinidae is composed of deep-sea sea stars distributed in eight genera and

approximately 70 valid species. So far, only five species of this family have been reported for the Mexican

waters of the Gulf of Mexico. Objective: To provide an updated local taxonomy of this family. Methods:

A total of 566 specimens deposited in the Echinoderm National Collection, Instituto de Ciencias del Mar

y Limnología, Universidad Nacional Autónoma de México, and the National Museum of Natural History,

Smithsonian Institution, collected from 1952 to 2015, were taxonomically examined. Results: We present

descriptions, photographs, and an illustrated dichotomous key for Benthopecten simplex simplex, Cheiraster

(Barbadosaster) echinulatus, Cheiraster (Cheiraster) planus, Cheiraster (Christopheraster) blakei, and

Cheiraster (Christopheraster) mirabilis in the region. Conclusions: The five studied species represent 6 % of

the world biodiversity of the family and can now be identified with the illustrated key.

Key words: Taxonomic catalogue; bathymetry; distribution; dichotomous key; deep sea.

De los Palos-Peña, M., Solís-Marín, F.A., & Laguarda-

Figueras, A. (2021). Taxonomy of family

Benthopectinidae (Echinodermata: Asteroidea) in the

Mexican waters of the Gulf of Mexico. Revista de

Biología Tropical, 69(S1), 118-131. DOI 10.15517/

rbt.v69iSuppl.1.46333

Sea stars of the family Benthopectinidae

Verrill, 1899 inhabit mostly in deep-waters, with-

in this family only the genera Pontaster Sladen,

1885 and Cheiraster Studer, 1883 extend on to

the shelf in shallow-waters (Clark & Downey,

1992). Benthopectinidae includes approximately

70 valid species distributed worldwide (Lam-

bert, 2000; Mah & Blake, 2012).

The species of this family are character-

ized by having five very large pointed arms,

longitudinally flexible dorso-ventrally and are

characterized by having a single pair of muscle

bands over the arms. The disc is small, the

papular areas are restricted to the proximal

region of each arm, and the marginal plates are

alternating and overlapping (Downey, 1973;

Clark & Downey, 1992; Benavides-Serrato,

Borrero-Pérez, & Diaz-Sanchez, 2011). The

taxonomy of the family has only been reviewed

by Ludwig (1910) when describing it as the

order Notomyotida. Later, Fisher (1911a) and

Verrill (1915) made some additions and mod-

ifications on the taxonomy of the Pacific

species. Since then, it was only A.M. Clark

(1981) and A.M. Clark & Downey (1992) who

reviewed the Atlantic species. The latter works

DOI 10.15517/rbt.v69iSuppl.1.46333

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are the most important in terms of genera and

species taxonomy.

The biology and distribution of the Atlan-

tic species of Benthopectinidae has been

studied by some authors (Pain et al., 1982;

Vázquez-Bader et al., 2008). There are also

some taxonomic studies (Mortensen, 1927;

Downey, 1973; Benavides et al., 2011) and

checklists of species (Durán-González et al.,

2005; Laguarda-Figueras et al., 2005; Alvarado

& Solís-Marín, 2013). However, to date, there

have been no taxonomic studies on the species

found in the Mexican waters of the Gulf of

Mexico. The aim of this work is to contribute to

the study of deep-sea Benthopectinidae in this

area and to update the external morphological

characters with taxonomic value.

MATERIALS AND METHODS

Specimens from the five species of Ben-

thopectinidae reported for the Mexican waters

of the Gulf of Mexico deposited in the Echi-

noderm National Collection at the Instituto

de Ciencias del Mar y Limnología, Univer-

sidad Nacional Autónoma de México, Mexico

(ICML-UNAM) and the United States Nation-

al Museum of Natural History, Smithsonian

Institution, United States National Museum

of Natural History, Smithsonian Institution,

Washington, D. C. (USNM), collected from

1952 to 2015, were taxonomically analyzed

and measured using a digital caliper (TRUPER

Caldi-6MP), the following measurements were

taken in each specimen, if possible: major

(R) and minor (r) radius in order to obtain

the R/r ratio. For the taxonomic identity of

the specimens, identification guides, original

diagnosis and specialized literature were used

(Perrier, 1881; Perrier, 1894; Sladen, 1889;

Ludwig, 1910; Fisher, 1911a; Fisher, 1911b;

Verrill, 1899; Verrill, 1915; Mortensen, 1927;

H.L. Clark, 1941; Downey, 1973; A.M. Clark,

1981; Clark & Downey, 1992). Photographs of

external taxonomical characters of each species

were taken with a multifocal microscope Leica

Z16 APOA at the Laboratorio de Microscopía

y Fotografía de la Biodiversidad II (Instituto de

Biología, Universidad Nacional Autónoma de

México). A dichotomous identification key was

elaborated, for which the main morphological

characters used were: shape of the abactinal

plates; location, shape, and number of mar-

ginal spines; shape and arrangement of papular

areas; and the presence and number of spines

and spinelets.

RESULTS

A total of 566 specimens belonging to two

genera (Benthopecten Verrill, 1884, Cheiraster

Studer, 1883), three subgenera (Barbadosaster

A.M. Clark, 1981, Cheiraster, and Christo-

pheraster A.M Clark, 1981) and five species

(Benthopecten simplex simplex (Perrier, 1881)

(seven specimens), Cheiraster (Barbadosaster)

echinulatus (Perrier, 1875) (78 specimens),

Cheiraster (Cheiraster) planus Verrill, 1915

(165 specimens), Cheiraster (Christopheraster)

blakei A.M. Clark, 1981 (186 specimens), and

Cheiraster (Christopheraster) mirabilis (Per-

rier, 1881) (130 specimens)) were identified.

Based on the material examined from both

collections, the species with the highest num-

ber of records was C. (Cheiraster) planus,

while B. simplex simplex and C. (Christopher-

aster) blakei were those with the least number

of records.

Dichotomous key to the genera, subgenera, and species of the family

Benthopectinidae of the Mexican waters of the Gulf of Mexico

1. Regular interradial suture between the marginal plates, without odd interradial plates (Fig. 1A-I)

(genus Cheiraster) ..............................................................................2

1’. Interradial suture between marginal plates absent, one odd interradial plate in each series, the superomarginal inserted

into the disc (Fig. 1A-II) (genus Benthopecten). A single very large spine on the odd interradial plate, paired supero-

marginals, also with a single spine and a few spinelets; papular areas distally bifurcated (Fig. 1B-I) and the most distal

pores aligned with the second or third superomarginal; adambulacral plates with five to seven furrow spines and two

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(on some plates one) subambulacral spines; oral plates with five or six furrow spines and three or four suborals; few

and prominent numerous actinal pectinate pedicellariae (Fig. 1C-I)……………………………………..Benthopecten

simplex simplex

2. Double papular areas, two lateral areas at each base of the arm proximally fused in large specimens (> 100 mm)

forming a bilobed V-shaped area (Fig. 1B-IV), pores small and interstitial. Arrangement of the dorsal superomarginal

plates framing the paxillary area in abactinal view .....................................................3

2’. Papular areas start in a single large primary middle pore (Fig. 1B-II, 1B-III) ..................................4

3. Parapaxiliform abactinal plates, not markedly convex, the most proximal crowded and highly variable in size, most of

the larger plates ornamented with a central spine of more than 2 mm long surrounded by small spinules. Papular areas

loosely joined and fused proximally only in large specimens, numerous pores, very small (subgenus Christopheraster).

Disc spines moderately long centrally, but not more than ½ of R, spaced and frequently extending to papular areas,

gradually decrease in size until reaching that of a spinelet. Marginal spines similar in size .............Cheiraster

(Christopheraster) blakei

3’. Few relatively very large spines, rarely more than 15, restricted to the center of the disc, without gradually decreasing

in size to that of a paxillary spinelet; the spine of the fourth superomarginal plate is conspicuously enlarged, mainly

widened at base and fourth inferomarginal plate reduced and sometimes without any spine.............Cheiraster

(Christopheraster) mirabilis

4. Irregular papular areas, restricted to the arm base and distally bilobed (except in small specimens, < 50 mm), lobes

may be unequal (Fig. 1B-II) (subgenus Barbadosaster). Proximal abactinal plates polygonal in shape, stacked and not

markedly convex, ornamented with numerous spinules forming two concentric rings around a central spinelet on the

larger plates. Superomarginal plates wide, adambulacral plates with seven to nine furrow spines, two subambulacral

spines, prominent pectinate pedicellariae (both types in 14 % of specimens) (Fig. 1C-III and Fig. 1C-IV), mainly in

the actinal areas and less frequently between the inferomarginal plates .....Cheiraster (Barbadosaster) echinulatus

4’. Papular areas start in a primary middle pore and develop in a “U”-shaped double series (Fig. 1B-III). Abactinal plates

parapaxiliform, the ones on the disc markedly convex and stacked together, on the disc and at the base of the arms

ornamented with a central spinelet between small spinules. Adambulacral plates with a single subambulacral spine

(subgenus Cheiraster), abactinal plates rounded with a group of spinules forming a single ring around a central spine,

superomarginal spines longer than the corresponding inferomarginal ones at least in the proximal half of the disc.

Rounded adambulacral plates with a series of six to nine furrow spines, pectinate pedicellariae (Fig. 1C-II) present in

some actinal plates and frequently in the inferomarginal plates ..................Cheiraster (Cheiraster) planus

Systematics:

Family Benthopectinidae Verrill, 1899

Genus Benthopecten Verrill, 1884

Benthopecten simplex simplex (Perrier, 1881)

(Fig. 2A, 2B, 2C, 2D)

Archaster simplex Perrier, 1881: 28.

Pararchaster armatus Sladen, 1889: 19, pls.

1, 4.

Pararchaster simplex.- Perrier, 1894: 253-256.

Benthopecten simplex.- Ludwig, 1910: 451,

464; Fisher, 1911a: 143; Verrill, 1915: 122;

Downey, 1973: 40, pl. 12; Pain et al., 1982:

195.

Benthopecten armatus.- Farran, 1913: 2;

Mortensen, 1927: 74, Fig. 41.

Benthopecten spinosus.- H.L. Clark, 1941: 26

Benthopecten chardyi Sibuet, 1975: 238, 289,

Fig. 3, pl. 1.

Benthopecten simplex simplex.- A.M. Clark,

1981: 130; A.M. Clark & Downey, 1992: 121,

pl. 30.

Diagnosis (modified from A.M. Clark,

1981): R = 150 mm, R/r = 7.7-7.9. Multiple

spinules in most of the proximal abactinal

plates, two or three but sometimes 10 when

they are accompanied by a spinelet or a bigger

spine, especially in specimens with average R

> 50 mm. Spines of the primary radial plates

tend to be more developed than the ones on

the primary interradial plates. Papular areas

distally bifurcating, the distalmost pores level

with the first three superomarginals or can

extend to the fourth in bigger specimens. There

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is an odd interradial superomarginal plate (Fig.

2A) with a single very large spine, the other

superomarginals also with a single spine and a

few spinelets. Adambulacral plates with five to

seven furrow spines and a single subambulacral

spine (Fig. 2D). Oral plates with five or six

furrow spines and three or four suboral spines.

Pedicellariae highly modified, usually present

in the actinal area, between the inferomargin-

als, and some abactinals (Fig. 2B).

Material examined: USNM E13210,

one specimen, Tamaulipas (24°52’12” N &

94°19’47” W), 3 713 m; USNM E13219, five

specimens, Veracruz (19°32’24” N & 95°4’11”

W), 2 185 m; USNM E30697, one speci-

men, Baltimore (38°2’60” N & 73°27’0” W),

1 916 m.

Geographic and bathymetric distribution:

Cape Cod; Gulf of Mexico, Tamaulipas, and

Veracruz; Colombia and Guyana basins and

from south and southwest of Iceland, Rockall

Trough; Kattegat, Denmark; south of the Gulf

of Guinea; 1 175 to 3 713 m (A.M. Clark &

Downey, 1992; Durán-González et al., 2005).

Remarks: According to the original

description (Perrier, 1881), the adambulacral

plates usually have five to seven furrow spines,

but in all the revised specimens there are five.

Fig. 1. A. Diagnostic morphological characters of species in the genera Cheiraster and Benthopecten: Cheiraster: subequal

superomarginal plates (I). Benthopecten: presence of an odd interradial plate (II). B. Distribution of papular pores in

Benthopecten simplex simplex (I), Cheiraster (Barbadosaster) echinulatus (II), Cheiraster (Cheiraster) planus (III),

and Cheiraster (Christopheraster) blakei and Cheiraster (Christopheraster) mirabilis (IV). C. Types of pedicellariae in

Benthopecten simplex simplex (I), Cheiraster (Cheiraster) planus (II) and in Cheiraster (Christopheraster) blakei and

Cheiraster (Christopheraster) mirabilis (III), Cheiraster (Barbadosaster) echinulatus presents type III and may or not

present type IV.

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Genus Cheiraster Studer, 1883

Subgenus Barbadosaster A.M. Clark, 1981

Cheiraster (Barbadosaster) echinulatus

(Perrier, 1875)

(Fig. 2E, 2F, 2G, 2H)

Archaster echinulatus Perrier, 1875: 348; Per-

rier, 1884: 263, pl. 10.

Cheiraster mirabilis.- Perrier, 1894: 276, pl.

20.

Cheiraster vincenti Perrier, 1894: 275.

Cheiraster echinulatus.- Perrier, 1894: 278;

Verrill, 1915: 129; Downey, 1973: 42, pl. 13.

Pontaster oligoporus Perrier, 1894: 293.

Luidiaster vincenti.- Ludwig, 1910: 425.

Pectinaster echinulatus.- Ludwig, 1910: 449.

Pectinaster mixtus Verrill, 1915: 140, pl. 6, 15,

17; Downey, 1973: 43, pl. 14.

Pectinaster vincenti.- Verrill, 1915: 139.

Pectinaster oligoporus.- Verrill, 1915: 147.

Luidiaster mixtus.- H.L. Clark, 1941: 29.

Cheiraster (Barbadosaster) echinulatus.- A.M.

Clark, 1981: 112, Figs. 3A-F, 4A-H; A.M.

Clark & Downey, 1992: 131, pl. 32B-E; Bena-

vides-Serrato et al., 2011: 143.

Diagnosis (modified from Downey, 1973):

R = up to 70 mm, average R/r = 4.5-5.7. Little

rigid and relatively short arms, abactinal plates

polygonal and paxiliform, each one bears from

eight to 25 granuliform spinelets and a small

central spine. Each papular area has three to

eight pores at the base of each arm without

any arrangement (Fig. 2E), but in some big

specimens the distal pores form two distal

lobes. Superomarginal plates are very large

and rectangular so that they invade the disc;

as a result, the paxilar area on the arms is very

narrow (Fig. 2G). Except for the first two inter-

radial plates, the superomarginal plates have

only one erect spine and they may have other

small spinelets around. The inferomarginal

plates have only one spine and about eight

small spines. Adambulacral plates with seven

to nine furrow spines which form a prolonged

angle less than 90° (Fig. 2H). Two subambula-

cral spines, very prominent pedicellariae on the

actinal plates (Fig. 2F), sometimes between the

adambulacral plates.

Material examined: USNM E12581,

one specimen, Quintana Roo (21°9’36” N &

86°25’48” W), 29 m; USNM E12778, one

specimen, Campeche (23°0’0” N & 86°47’59”

W), 482 m; USNM E12699, two specimens,

Campeche (24°18’0” N & 87°49’47” W), 375

m; USNM E23485, one specimen, Bahamas

(25°30’0” N & 79°20’59” W), 311 m; ICML-

UNAM 8033, four specimens, Campeche

(22°24’34” N & 91°34’86” W), 539 m;

ICML-UNAM 8458, four specimens, Yuca-

tán (24°16’60” N & 88°12’57” W), 455 m;

ICML-UNAM 8468, five specimens, Quin-

tana Roo (23°46’20” N & 87°06’64” W),

618 m; ICML-UNAM 8472, one specimen,

Quintana Roo (23°48’38” N & 87°08’16” W),

613 m; ICML-UNAM 8680, three specimens,

Campeche (22°15’87” N & 91°44’89” W), 250

m; ICML-UNAM 8725, two specimens, Yuca-

tán (23°10’83” N & 89°59’71” W), 460 m;

ICML-UNAM 8728, one specimen, Yucatán

(23°14’95” N & 89°59’53” W), 536 m; ICML-

UNAM 5068, 53 specimens, Quintana Roo

(23°22’09” N & 88°06’09” W), 122 m.

Geographic and bathymetric distribution:

Gulf of Mexico, Campeche, Yucatán, Quintana

Roo; Florida strait, south to Nicaragua and

Venezuela, east of Leeward Islands, Cuba,

Curaçao, San Vicente, Colombia; 29 to 5 062

m (Clark & Downey, 1992; Benavides-Serrato

et al., 2011).

Remarks: The bathymetric range of this

species is extended to its shallower limit, since

it was previously reported from 130 to 5 062

m (Clark & Downey, 1992; Benavides-Serrato

et al., 2011), and in the present work it has

been found at 29 m depth. There are two types

of pectinate pedicellariae, the first ones are

mounted on the surface of the actinal plates

with a rounded margin, and the second ones

are located between two actinal plates, half

of the pedicellariae in one plate and the other

half on the other, with irregular margins. These

pedicellariae are not exclusive of the species,

but it is important to note that both types may

be observed in the same specimen, unlike the

other four species that generally have only one

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Fig. 2. A-D. Benthopecten simplex simplex (USNM E30697, E13219). A. Abactinal view and odd interradial plate with

conspicuous spine, rectangle shows the spine. B. Actinal view and detail of oral plates; arrow shows pedicellariae. C.

Detail of marginal spines, rectangle shows spine. D. Adambulacral spines, rectangle shows furrow spines and suboral spine

insertion. E-H. Cheiraster (Barbadosaster) echinulatus (USNM E23485, E12699). E. Detail of superomarginal plates,

rectangle shows plate. F. Adambulacral spines, rectangle shows furrow and subambulacral spines. G. Abactinal view and

detail of irregular shape of papular areas, rectangle shows superomarginal plates. H. Actinal view and detail of oral plates;

arrow shows pedicellariae.

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type of pedicellariae. This was observed only in

14 % of the reviewed specimens.

Subgenus Cheiraster Studer, 1883

Cheiraster (Cheiraster) planus Verrill, 1915

(Fig. 3A, 3B, 3C, 3D)

Cheiraster planus Verrill, 1915: 133, pl. 18.

Pectinaster gracilis Verrill, 1915: 145, pl. 6,

14, 15.

Cheiraster mirabilis.- Downey, 1973: 41, pl.

13.

Cheiraster (Cheiraster) planus.- A.M. Clark

1981: 116, Fig. 5A, Fig. 5B; A.M. Clark &

Downey, 1992: 128, pl. 31A-C; Benavides-

Serrato et al., 2011: 140.

Diagnosis (modified from Verrill, 1915;

Clark & Downey, 1992): R = up to 120 mm

approximately, R/r = 6.1-8.9. Disc consider-

ably broad and arms are very long and slender,

abactinal plates are uniform in size, rounded

and convex with a group of spinules. On the

proximal plates the average number of spinules

is 10, forming a ring surrounding a central

spinelet (Fig. 3A). Papular areas start with one

big single median pore and develop into two

double U-shaped series, being irregular only

in the biggest specimens. The plates of the

papular areas next to the madreporite and in

the central part of the disc are larger than the

rest and have numerous spines. Marginal spines

are well developed, conical and broad at the

base. Superomarginal spines are distinctively

larger than the corresponding inferomarginals

(Fig. 3C), the rest of the plate is covered with

fine spinelets and there is only one accessory

spine. Adambulacral plates have six to nine

rounded spines and one conical subambulacral

spine mounted on a convex protuberance (Fig.

3D). Pectinate pedicellariae present on the

actinal plates and on the inferomarginal plates

(Fig. 3B).

Material examined: USNM E13215, seven

specimens, Veracruz (21°30’0” N & 96°55’12”

W), 1238 m; USNM E23587, six specimens,

Quintana Roo (18°30’0” N & 87°36’35” W),

715 m; USNM E41881, 19 specimens, Louisi-

ana, USA (27°23’60” N & 93°18’36” W), 732

m; ICML-UNAM 5024, one specimen, Quin-

tana Roo (23°22’0” N & 88°06’0” W), 86.6 m;

ICML-UNAM 5025, one specimen, Quintana

Roo (22°23’02” N & 87°15’04” W) depth

not recorded; ICML-UNAM 7980, four speci-

mens, Veracruz (18°46’03” N & 94°25’96” W),

386 m; ICML-UNAM 7979, three specimens,

Veracruz (18°47’0.43” N & 94°32’8.57” W),

380 m; ICML-UNAM 8029, four specimens,

Campeche (22°24’15” N & 91°35’00” W),

548 m; ICML-UNAM 8033, seven specimens,

Campeche (22°24’34” N & 91°34’86” W),

539 m; ICML-UNAM 8036, three specimens,

Campeche (22°24’12” N & 91°35’75” W),

548 m; ICML-UNAM 9022, one specimen,

Tamaulipas (24°55’58” N & 96°30’50” W),

700 m; ICML-UNAM 9026, 12 specimens,

Tamaulipas (24°56’05” N & 96°29’18” W),

760 m; ICML-UNAM 9028, 24 specimens,

Tamaulipas (24°56’48” N & 96°24’59” W),

821 m; ICML-UNAM 9193, nine specimens,

Veracruz (18°57’107” N & 94°20’0” W), 671.9

m; ICML-UNAM 9196, six specimens, Vera-

cruz (18°59’04” N & 94°19’09” W), 815

m; ICML-UNAM 9218, six specimens, Vera-

cruz (19°06’33” N & 94°06’38” W), 867

m; ICML-UNAM 9222, one specimen, Vera-

cruz (18°57’06” N & 94°06’61” W), 524 m;

ICML-UNAM 9247, one specimen, Tabasco

(19°08’81” N & 93°27’48” W), 677 m; ICML-

UNAM 9260, one specimen, Tabasco (19°40’0”

N & 92°45’13” W), 770 m; ICML-UNAM

8791, one specimen, Campeche (22°29’38”

N & 90°43’66” W), 130 m; ICML-UNAM

9639, one specimen, Yucatán (23°36’02” N &

89°34’51” W), 435 m; ICML-UNAM 9707,

one specimen, Quintana Roo (23°38’49”N &

87°04’84” W), 572 m; ICML-UNAM 9716,

two specimens, Quintana Roo (23°40’58” N

& 87°04’84” W), 642 m; ICML-UNAM 9746,

eight specimens, Quintana Roo (23°26’70”

N & 86°53’0” W), 633 m; ICML-UNAM

9826, two specimens, Campeche (22°20’18”

N & 91°41’46” W), 428 m; ICML-UNAM

9850, one specimen, Campeche (22°26’00”

N & 91°26’86” W), 546 m; ICML-UNAM

9900, two specimens, Tabasco (19°33’82” N &

93°01’46” W), 664 m; ICML-UNAM 11551,

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one specimen, Tamaulipas (24°55’0” N &

96°29’07” W), 766 m; ICML-UNAM 11555,

three specimens, Tamaulipas (24°54’08”

N & 96°31’40” W), 683 m; ICML-UNAM

11563, four specimens, Tamaulipas (24°55’0”

N & 96°29’07” W), 765 m; ICML-UNAM

11617, one specimen, Veracruz (18°53’90”

N & 94°08’37” W), 557 m; ICML-UNAM

11637, two specimens, Tabasco (19°08’99”

N & 93°27’91” W), 690 m; ICML-UNAM

11644, five specimens, Tabasco (19°13’57”

N & 93°54’88” W), 746 m; ICML-UNAM

11690, one specimen, Tabasco (19°04’61”

N & 94°04’49” W), 828 m; ICML-UNAM

11700, one specimen, Tabasco (19°02’96”

N & 94°05’33” W), 764 m; ICML-UNAM

11708, one specimen, Tabasco (18°54’99”

N & 93°51’69” W), 419 m; ICML-UNAM

11741, two specimens, Tabasco (18°58’0”

N & 94°07’57” W), 710 m; ICML-UNAM

12343, one specimen, Yucatán (23°22’45”

N & 89°59’28” W), 582 m; ICML-UNAM

12386, four specimens, Yucatán (23°18’26”

N & 89°56’60” W), 410 m; ICML-UNAM

12376, five specimens, Yucatán (23°13’76” N

& 89°58’23” W), 392 m.

Geographic and bathymetric distribu-

tion: Gulf of Mexico, Tamaulipas, Veracruz,

Tabasco, Campeche, Yucatán, Quintana Roo;

Jamaica, Cuba, Honduras, Trinidad and Toba-

go, Nicaragua, Colombia, Las Antillas and

Guyana; 86.6 to 1 339 m (Durán-González et

al., 2005; Benavides-Serrato et al., 2011).

Remarks: The bathymetric range of this

species is extended to its shallower limit, since

it was previously reported from 226 to 1 339

(Alvarado & Solís-Marín, 2013) and in the

present work it has been found at 86.6 m depth.

Subgenus Christopheraster A.M. Clark, 1981

Cheiraster (Christopheraster) blakei

A.M. Clark, 1981

(Fig. 3E, 3F, 3G, 3H)

Cheiraster mirabilis.- Verrill, 1915: 124, pl. 14.

Cheiraster echinulatus.- Verrill, 1915: 131, pl.

14, 19.

Cheiraster coronatus.- H.L. Clark, 1941: 26.

Luidiaster enoplus.- H.L. Clark, 1941: 28.

Cheiraster enoplus.- Downey, 1973: 42, pl. 14.

Cheiraster (Christopheraster) blakei A.M.

Clark, 1981: 113, figs. 1b, 2c-f; A.M. Clark &

Downey, 1992: 132, pl. 32A, 33D, E; Bena-

vides-Serrato et al., 2011: 145.

Diagnosis (modified from A.M. Clark,

1981): R = up to 130 mm, R/r = 6.0-9.8/1 for

large specimens, and 4.5-6.5/1 in specimens of

R = 30-60 mm. Arms very long and become

attenuated in larger specimens. Abactinal plates

convex, polygonal, and very variable in size,

the primary radial and interradial ones are

conspicuous and have 25 small spinules, 10

on average; the larger plates develop a long

spine and form a non-isolated group (Fig. 3E).

Papular areas on each arm ill-defined, exten-

sive, and distally bilobed with numerous pores

that extend up to the third superomarginal

plate. Superomarginal plates have only one

spine (Fig. 3G), the larger ones are on the fifth

and sixth plates; correspondent inferomarginal

spines similar or slightly larger, with a small

accessory spine below it. Actinal areas small

and triangular in shape. Each adambulacral

plate has seven to nine furrow spines, forming

a 90° angle, but it can be obtuse (Fig. 3H). A

single subambulacral spine and usually small

spinules in the actinal view of the plate. Pec-

tinate pedicellariae develop in the adradial

actinal plates, on the adjacent margins of the

adambulacral plates, and sometimes in the

abactinal surface (Fig. 3F).

Material examined: USNM E23428, one

specimen, Yucatán (21°16’48” N & 86°12’35”

W), 412 m; USNM E23427, Bahamas (27°

48’36” N & 78°49’47” W), 824 m; USNM

E23566, six specimens, Campeche (23°55’48”

N & 87°31’48” W), 274 m; USNM E23509,

four specimens, Campeche (23°58’12” N

& 87°29’23” W), 366 m; USNM E23601,

six specimens, Quintana Roo (18°30’0” N

& 87°36’35” W), 715 m; USNM E38622,

two specimens, Quintana Roo (18°49’48” N

& 87°31’11” W), 695 m; USNM E23427,

eight specimens, Bahamas (27°48’36” N &

78°49’47” W), 824 m; ICML-UNAM 8169,

three specimens, Yucatán (22°24’7” N &

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Fig. 3. A-D. Cheiraster (Cheiraster) planus (USNM E41881, ICML-UNAM 11708). A. Abactinal view and detail of

abactinal plates. B. Actinal view and detail of oral plates; arrow shows pedicellariae. C. Marginal spines, rectangle shows

superomarginals longer than inferomarginales. D. Adambulacral spines, rectangle shows furrow spines and suboral spine

insertion. E-H. Cheiraster (Christopheraster) blakei (USNM E23427, ICML-UNAM 8778). E. Detail of superomarginal

plates, rectangle shows furrow spines and subambulacral spine. F. Adambulacral spines. G. Abactinal view, rectangle and

arrow show the group of non-isolated spines that decrease in size towards the arms. H. Actinal view; detail of oral plates,

arrow shows pedicellariae. I-L. Cheiraster (Christopheraster) mirabilis (USNM E23416). I. Abactinal view, rectangles

show abactinal plates and isolated group of spines at the center of the disc, papular areas. J. Actinal view and detail of oral

plates; arrow shows pedicellariae. K. Detail of superomarginals, rectangle shows the spine of the fourth superomarginal plate

enlarged. L. Detail of adambulacral plates, rectangle shows the spine of the fourth inferomarginal plate reduced.

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91°30’7” W), 407 m; ICML-UNAM 8778, one

specimen, Yucatán (22°46’16” N & 90°46’88”

W), 735 m; ICML-UNAM 12318, one speci-

men, Yucatán (23°33” N & 89°40” W), 358 m;

ICML-UNAM 12385, 62 specimens, Yucatán

(23°18’26” N & 89°56’60” W), 410 m; ICML-

UNAM 12375, 92 specimens, Yucatán (23°13’

74” N & 89°58’23” W), 392 m.

Geographic and bathymetric distribution:

Gulf of Mexico, Campeche, Yucatán; Bahamas,

southeast Florida, Cuba, Belize, Colombia,

Honduras, Jamaica, north of Brazil; 250 to 1

030 m (being more abundant from 500 to 800

m) (Benavides-Serrato et al., 2011; Solís-Marín

et al., 2014).

Cheiraster (Christopheraster) mirabilis

(Perrier, 1881)

(Fig. 3I, 3J, 3K, 3L)

Archaster mirabilis Perrier, 1881: 27.

Cheiraster coronatus.- Perrier, 1894: 271; Lud-

wig, 1910: 455.

Cheiraster mirabilis.- Verrill, 1915: 124.

Cheiraster enoplus Verrill, 1915: 135, pl. 18.

Cheiraster (Christopheraster) mirabilis.- A.M.

Clark, 1981: 112, Fig. 2A, 2B; A.M. Clark &

Downey, 1992: 133; Benavides-Serrato et al.,

2011: 147.

Diagnosis (modified from Verrill, 1915):

R = up to 185 mm, R/r = 7.5-8.8/1 for large

specimens (R > 70 mm), R/r = 6-7.5/1 for R

= 30-60 mm. Arms very long and thin. Center

of the disc with a dozen long spines (Fig. 3I).

Superomarginal plates somewhat small, imbri-

cated and with a convex proximal edge, surface

covered with numerous fine spinules. Papular

pores form two symmetric lateral groups that

extend up to the disc. Superomarginal plates

have a marginal long, conical spine, the one of

the fourth plate abruptly elongated (Fig. 3K),

and the first three reduced. Inferomarginal

plates covered with long but fine spinules, in

the first three plates there are four to five spines

that increase in size, the one in the fourth plate

is reduced (Fig. 3L), the rest of the spines

longer than the corresponding superomargin-

als; some of these plates have fasciculate

rudimentary pedicellariae. Adambulacral plates

with 10-12 furrow spines and one large, conical

subambulacral spine (Fig. 3L). Prominent oral

plates with about eleven spines, two or three

pectinate pedicellariae between the interactinal

plates (Fig. 3J).

Material examined: USNM E50647, one

specimen, Quintana Roo (22°41’24” N &

86°40’48” W), 411 m; USNM E23416, three

specimens, Bahamas (25°58’48” N & 78°12’0”

W), 658 m; ICML-UNAM 8162, three speci-

mens, Yucatán (22°28’08” N & 91°12’95”

W), 416 m; ICML-UNAM 8714, one speci-

men, Yucatán (23°17’28” N & 89°56’73” W),

388 m; ICML-UNAM 9624, five specimens,

Campeche (22°24’42” N & 91°30’35” W),

364 m; ICML-UNAM 9641, one specimen,

Yucatán (23°30’98”N & 89°49’42” W), 422

m; ICML-UNAM 9661, four specimens, Yuca-

tán (24°14’89” N & 88°12’59” W), 423 m;

ICML-UNAM 9686, one specimen, Quintana

Roo (24°21’45” N & 87°37’92” W), 812 m;

ICML-UNAM 9691, one specimen, Quintana

Roo (24°24’57” N & 87°38’22” W), 820 m;

ICML-UNAM 9747, three specimens, Quin-

tana Roo (23°26’70” N & 86°53’07” W),

633 m; ICML-UNAM 9769, one specimen,

Quintana Roo, (23°35’46” N & 86°50’08”

W), 806 m; ICML-UNAM 9825, 39 speci-

mens, Campeche (22°20’18” N & 91°41’46”

W), 428 m; ICML-UNAM 8759, one speci-

men, Campeche (22°46’23” N & 90°45’79”

W), 728 m; ICML-UNAM 8773, three speci-

mens, Campeche (22°33’ N & 90°48’ W),

346 m; ICML-UNAM 11690, one specimen,

Tabasco (19°04’61” N & 94°04’49” W),

828 m; ICML-UNAM 9826, one specimen,

Campeche (22°20’18” N & 91°41’46” W),

428 m; ICML-UNAM 12386, three speci-

mens, Yucatán (23°18’26” N & 89°56’60” W),

410 m; ICML-UNAM 12376, nine specimens,

Yucatán (23°13’76” N & 89°58’23” W), 392

m; ICML-UNAM 12385, 19 specimens, Yuca-

tán (23°18’26” N & 89°56’60” W), 410 m;

ICML-UNAM 12375, 30 specimens, Yucatán

(23°13’74” N & 89°58’23” W), 392 m.

Geographic and bathymetric distribution:

Gulf of Mexico, Tabasco, Campeche, Yucatán,

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Quintana Roo; southeast Florida, Cuba, Jamai-

ca; 342 to 1 470 m, being more abundant from

400 to 700 m depth (Clark & Downey, 1992;

Durán-González et al., 2005; Benavides-Serra-

to et al., 2011).

Remarks: The bathymetric range of this

species is extended to its shallower limit, since

it was previously reported at 380 m (Alvarado

& Solís-Marín, 2013), and in the present work

it has been found at 342 m depth.

DISCUSSION

The family Benthopectinidae corresponds

to a group of organisms that occur mainly in

deep waters and whose species richness con-

sists of eight genera, and approximately 70

valid species (Mah & Blake, 2012). This family

is the only one included in the order Notomy-

otida described by Ludwig (1910). Mah and

Foltz (2011) suggested a phylogenetic hypoth-

esis from Valvatacea and Paxillosida orders,

in which the family Benthopectinidae was

supported as a sister group to Pseudarchast-

eridae Sladen, 1889 and was included within

the order Paxillosida. This was supported by

Mah and Blake (2012) since they have shown

close morphological resemblance to the Goni-

asteridae Forbes, 1841 family, as both families

occur primarily in deep-sea or high-latitude

and correspond to widely distributed taxa.

Also, Linchangco et al. (2017) placed the order

Paxillosida as sister to Cheiraster sp., show-

ing results similar to those presented by Mah

and Foltz (2011), since they did not use mor-

phological characters either. However, in this

study, the Benthopectinidae family was consid-

ered within the order Notomyotida, following

the classification proposed by A.M. Clark &

Downey (1992), until a phylogenetic recon-

struction method is realized including mor-

phological characters. So far, there have been

no more detailed studies that are only focused

on this taxon. The only previous taxonomic

work was carried out by Clark (1981) for the

Atlantic species where the author redefined

the genera Cheiraster and Pectinaster Perrier,

1885, reduced the genus Luidiaster Studer,

1883 to a subgenus of Cheiraster, described

two more subgenera for Cheiraster, and three

new species. The subdivision within the gen-

era may be useful in some cases to identify

certain morphological similarity between the

species. However, the characters used for the

designation of subgenera within the genera in

this family cannot be applied in all cases, since

the characters related to ontogeny such as the

early or late development of certain spines, are

highly variable, and therefore it is suggested

that a complete family review that includes

molecular methods and the use of morphologi-

cal characters including ossicle morphology is

realized to redefine the species and eliminate

confusing or unnecessary subdivisions.

Five species of Benthopectinidae are

reported for the Mexican waters of the Gulf

of Mexico, which represents 6 % of the world

diversity of this family. Since the species of

this family are distributed in eight genera

(Mah & Blake, 2012), the two found in these

waters represent 25 % of the total genera of

the family. Regarding the taxonomy of the

species, Cheiraster (Christopheraster) mira-

bilis and Cheiraster (Christopheraster) blakei

share morphological characteristics such as the

shape of the papular areas of different ontog-

eny, this was described by Clark (1981) and

showed drawings of the different arrangement

that the species of the genus present, and that

the development of the pores throughout the

growth is very particular in these two species.

The change in the pattern of pore distribution

in both species is not very marked, but it can

be appreciated in some of the reviewed speci-

mens. Based on the review of the specimens

from the Gulf of Mexico, we determined that,

to differentiate both species, the two most

important characters are: 1) the number and

arrangement of spines on the disc, which are

abundant and present in the entire disc, gradu-

ally decreasing in size towards the arms in C.

(Christopheraster) blakei, and which are all

similar in size and restricted to the center of the

disc in C. (Christopheraster) mirabilis; and 2)

the superomarginal spines, which are sub-equal

in size in C. (Christopheraster) blakei, while

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in C. (Christopheraster) mirabilis the spine of

the fourth superomarginal plate is much more

developed than the rest. It is important to note

that the last characteristic presents variation;

while in the Bahamas specimens the spine of

the fourth superomarginal plate is markedly

larger than the rest (up to 15 mm as reported

by A.M. Clark & Downey, 1992), in the Gulf

of Mexico specimens, the spine is larger but

not almost twice as large, as reported by

Clark (1981).

In Cheiraster (Barbadosaster) echinula-

tus, one of the diagnostic characteristics of the

species is the rectangular shape of the supero-

marginal plates that considerably invade the

disc. The variation of shape of these plates is

mainly that they tend to invade the disc, but,

especially in small specimens, the plates are

only slightly longer than they are wide, which

means the rectangular shape is not as obvious

as Perrier (1875) described. In the case of Chei-

raster (Cheiraster) planus, the main character-

istic is the marginal spines, since the ones in the

superomarginal plates are larger in size than the

corresponding inferomarginal ones. Although

this species was described by Verrill (1915)

from a single specimen, it is the most common

species of the family in the Gulf of Mexico.

Finally, Benthopecten simplex simplex, is a spe-

cies whose type locality is the Gulf of Mexico,

but a subspecies (Benthopecten simplex char-

dyi Sibuet, 1975) has been reported in the Gulf

of Guinea and Gabon (Clark & Downey, 1992).

This subspecies is still considered valid, since

it differs from B. simplex simplex by a delayed

development of the disc spines and a late

appearance of the second subambulacral spine,

in addition to a greater number of adambulacral

plates corresponding to the first 10 inferomar-

ginal ones. According to Clark (1981), these

can be considered a combination of extremes in

the characters subject to variation in B. simplex

simplex. In other words, they could simply be

two highly variable characteristics. However,

the subspecies has not been validated or refuted

since there are no records in the rest of the

Atlantic of an intermediate form or specimens

that may suggest that these characters are sim-

ply subject to great variation.

A more detailed study may find that the

diversity of the family is much greater and that

there are many cryptic or overlooked species

due to lack of definition between one species

and another, as is the example of the synonym

of B. simplex simplex: Benthopecten armatus

(Sladen, 1889), synonymized with Benthopec-

ten spinosus Verrill, 1884 by some authors,

such as Grieg (1921) which is in fact a valid

species, and treated as valid by other authors

like Mortensen (1927) being finally synony-

mized with B. simplex simplex by Clark (1981).

The present study represents the first updated

compilation of the species of this family pres-

ent in the Gulf of Mexico, as well as the first

work providing external morphological char-

acters of taxonomic importance to distinguish

them. It is the first time that a dichotomous key

for these species in the Mexican waters of the

Gulf of Mexico is presented.

Ethical statement: authors declare that

they all agree with this publication and made

significant contributions; that there is no con-

flict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

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.

ACKNOWLEDGMENTS

We would like to thank to Ma. Esther

Diupotex Chong and Alicia Durán González

for their technical support at the ENC, ICML,

UNAM. To Susana Guzmán Gómez for her

technical support with the multifocal photogra-

phy. To CONACYT for the scholarship (num-

ber 929010) and to Dave Pawson, William

Moser (USNM) and the Smithsonian staff for

their support. Finally, the authors would like

to thank M.G. Lovegrove and A.A. Caballero-

Ochoa for the valuable comments on the manu-

script’s English and scientific content.

130

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RESUMEN

Descripción taxonómica de las especies de la familia

Benthopectinidae (Echinodermata: Asteroidea)

de las aguas mexicanas del Golfo de México

Introducción: La familia Benthopectinidae está

compuesta por estrellas de mar de profundidad distribuidas

en ocho géneros y 70 especies válidas aproximadamen-

te. Hasta el momento, se han reportado cinco especies

de esta familia para las aguas mexicanas del Golfo de

México. Objetivo: Realizar una actualización local de

la taxonomía de esta familia. Métodos: Se examinaron

un total de 566 especímenes depositados en la Colección

Nacional de Equinodermos, Instituto de Ciencias del Mar

y Limnología, Universidad Nacional Autónoma de México

y el Museo Nacional de Historia Natural, Smithsonian

Institution, recolectados de 1952 a 2015. Resultados: Pre-

sentamos descripciones, fotografías y una clave dicotómica

ilustrada para Benthopecten simplex simplex, Cheiraster

(Barbadosaster) echinulatus, Cheiraster (Cheiraster) pla-

nus, Cheiraster (Christopheraster) blakei, y Cheiraster

(Christopheraster) mirabilis en la región. Conclusiones:

Las cinco especies estudiadas representan el 6 % de la

diversidad mundial de la familia y ahora pueden ser identi-

ficadas con la clave ilustrada proporcionada.

Palabras clave: Catálogo taxonómico; batimetría; distri-

bución; clave dicotómica; mar profundo.

REFERENCES

Alvarado, J.J., & Solís-Marín, F.A. (2013). Echinoderm

Research and Diversity in Latin America. Berlin,

Heidelberg: Springer.

Benavides-Serrato, M.G., Borrero-Pérez, G.H., & Diaz-

Sanchez, C.M. (2011). Equinodermos del Caribe

colombiano I: Crinoidea, Asteroidea y Ophiuroidea.

Santa Marta, Colombia: Serie de Publicaciones Espe-

ciales de Invemar.

Clark, A.M. (1981). Notes on Atlantic and other Aste-

roidea. 1. Family Benthopectinidae. Bulletin of the

British Museum (Natural History), Zoology series,

41(3), 91-135.

Clark, A.M., & Downey, M.E. (1992). Starfishes of the

Atlantic. London, United Kingdom: Chapman &

Hall.

Clark, H.L. (1941). The echinoderms (other than holothu-

rians). Reports on the scientific results of the Atlantis

expeditions to the West Indies, under the joint auspi-

ces of the University of Havana and Harvard Univer-

sity. Memorias de la Sociedad Cubana de Historia

Natural “Felipe Poey”, 15(1), 1-154.

Downey, M.E. (1973). Starfishes from the Caribbean and

the Gulf of Mexico. Smithsonian Contributions to

Zoology, 126, 1-158.

Durán-González A., Laguarda-Figueras, A., Solís-Marín,

F.A., Buitrón-Sánchez, B.E., Ahearn, C.G., & Torres-

Vega, J. (2005). Equinodermos (Echinodermata) de

las aguas mexicanas del Golfo de México. Revista de

Biología Tropical, 53(3), 53-68.

Farran, G.P. (1913). The deep-water Asteroidea, Ophiu-

roidea and Echinoidea of the West coast of Ireland.

Scientific Investigations, 1912, Fisheries Branch,

12(6), 1-66.

Fisher, W.K. (1911a). Two new genera of starfishes. The

Annals and Magazine of Natural History, 8(7), 89-92.

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

adjacent waters. Part 1. Phanerozonia and Spinulo-

sa. Bulletin of the United States National Museum,

Smithsonian Institution, 76, 1-419.

Forbes, E. (1841). A history of British starfishes and

other animals of the class Echinodermata. London,

England: John Van Voorst.

Grieg, JA., (1921). Echinodermata. Report of the scientific

results of the Michael Sars North Atlantic deep-sea

expedition 1910, 3(2): 1-47.

Laguarda-Figueras A., Solís-Marín, F.A., Durán-González,

A., Ahearn, C.G., Buitrón-Sánchez, B.E., & Torres-

Vega, J. (2005). Equinodermos (Echinodermata) del

Caribe Mexicano. Revista de Biología Tropical,

53(3), 109-122.

Lambert, P. (2000). Sea Stars of British Columbia,

Southeast Alaska, and Puget Sound. Canada: UBC

Press, Royal British Columbia Museum.

Linchangco, G.V., Foltz, D.W, Reid, R., Williams, J., Nod-

zak, C., Kerr, A.M, Miller, A., Hunter, R., Wilson,

N.G., Nielsen, W.J., Mah, C.L., Rouse, G.W, Wray,

G.A., & Janies, D.A. (2017). The phylogeny of extant

starfish (Asteroidea: Echinodermata) including Xylo-

plax, based on comparative transcriptomics. Molecu-

lar Phylogenetics and Evolution, 115, 161-170.

Ludwig, H. (1910). Notomyota, eine neue Ordnung der

Seesterne. Sitzungsberichte der Königlich Preussis-

chen Akademie der Wissenschaften, 23, 435-466.

Mah, C., & Foltz, D. (2011). Molecular phylogeny of the

Valvatacea (Asteroidea: Echinodermata). Zoological

Journal of the Linnean Society, 161(4), 769-788.

Mah, C., & Blake, D.B. (2012). Global diversity and phylo-

geny of the Asteroidea (Echinodermata). PloS ONE,

7(4), e35644.

Mortensen, T. (1927). Handbook of the echinoderms of the

British Isles. Great Britain: Oxford University Press.

Pain S.L., Tyler, P.A., & Gage, J.D. (1982). The reproduc-

tive biology of the deep-sea asteroids Benthopecten

simplex (Perrier), Pectinaster filholi Perrier, and Pon-

taster tenuispinus Düben & Koren (Phanerozonia:

131

Revista de Biología Tropical, ISSN electrónico: 2215-2075, Vol. 69(S1): 118-131, March 2021 (Published Mar. 30, 2021)

Benthopectinidae) from the rockall trough. Journal

of Experimental Marine Biology and Ecology, 65(2),

195-211.

Perrier, E. (1875). Révision de la collection de stellérides

du Museum d’Histoire naturelle de Paris. Paris:

Reinwald.

Perrier, E. (1881). Description sommaire des espèces nou-

velles d’Astéries. Reports on the results of dredging

under the supervision on Alexander Agassiz, in the

Gulf of Mexico, 1877-78, by the United States coast

survey steamer “Blake”, Lieut.-Commander C.D.

Sigsbee, U.S.N., commanding, and in the Caribbean

Sea, 1878-79, by the U.S.C.S.S. “Blake”, commander

J.R. Bartlett, U.S.N., commanding. Bulletin of the

Museum of Comparative Zoölogy at Harvard Colle-

ge, 9(1), 1-31.

Perrier, E. (1885). Premiére note Préliminaire des les

Echinodermes, recueillis durant les campagnes de

dragages sous-marines du Travailleur et du Talisman.

Annales des sciences naturelles, Zoologie, 22(8),

1-72.

Perrier, E. (1894). Échinodermes. Expéditions Scientifi-

ques du Travailleur et du Talisman Pendant les années

1880, 1881, 1882, 1883. Paris: G. Masson Éditeur.

Sibuet, M. (1975). Astérides abyssales de l’Atlantique sud.

(Résultats de la campagne Walda, juin-juillet-aout

1971). Bulletin du Muséum National d’Historie Natu-

relle, 199, 281-296.

Sladen, W.P. (1885). The Asteroidea. Report on the scienti-

fic results of the voyage of H.M.S. Challenger during

the years 1873-76 under the command of Captain

George S. Nares, R.N., F.R.S. and the late Captain

Frank Tourle Thomson, R.N. prepared under the

superintendence of the late Sir C. Wyville Thomson,

Knt., F.R.S., %c. regius professor of Natural History

in the University of Edinburgh director of the civilian

scientific staff on board and now of John Murray one

of the naturalists of the expedition. Narrative, 1(2),

607-617.

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

by H.M.S. “Challenger” during the years 1873-1876.

Zoology, 30, 1-893.

Solís-Marín F.A., Laguarda-Figueras, A., Durán-González,

A., Vázquez-Bader, A.R., & Gracia, A. (2014). Bio-

diversidad de los equinodermos (Echinodermata) del

mar profundo mexicano. In A. Low-Pfeng, & E.M.

Peters (Eds.), La frontera final: el océano profundo

(215-253). México: INECC-SEMARNAT.

Studer, T. (1883). Über der Asteriden, welche während

der Reise S.M.S. Gazelle um die Erde gesammelt

wurden. Sitzungs-berichte der Gesellschaft Natur-

forschender Freunde zu Berlin, 8, 128-132.

Vázquez-Bader, A.R., Laguarda-Figueras, A., Gracia, A.,

Solís-Marín, F.A., Celaya-Hernández, E.V., & Durán-

González, A. (2008). Seasonal changes in the density

and species composition of the epifaunal echinoderms

recorded from the southwestern Gulf of Mexico.

Revista de Biología Tropical, 56(3), 297-310.

Verrill, A.E. (1884). Notice of the remarkable marine fauna

occupying the Outer Banks off the Southern Coast of

New England, No. 9. American Journal of Science,

s3-28(165), 213-220.

Verrill, A.E. (1899). IV. Revision of certain genera and

species of starfishes with descriptions of New Forms.

Transactions of the Connecticut Academy of Arts and

Science, 10(4), 145-234.

Verrill, A.E. (1915). Report on the starfishes of the West

Indies, Florida, and Brazil, including those obtained

by the Bahama expedition from the University of

Iowa in 1893. Bulletins from the laboratories of Natu-

ral History, 7(1), 1-232.