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Deep-water Holothuroidea (Echinodermata) collected during the TALUD cruises off the Pacific coast of Mexico, with the description of two new species


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Research cruises aboard the R/V "E1 Puma" were organized to collect deep-water benthic and pelagic specimens off the Pacific coast of Mexico. Seventy four specimens of Holothuroidea were collected off the Pacific coast of Mexico in depths of 377-2 200 m. The collection includes representatives of 5 of the 6 orders of Holothuroidea, 3 Dendrochirotida, 2 Dactylochirotida, 2 Aspidochirotida, 4 Elasipodida and 2 Molpadiida. Apodida were not represented. Of the 13 species recognized, 11 were identified to species level and 2, belonging to the genera Ypsilocucumis Panning, 1949, and Mitsukuriella Heding and Panning, 1954, are new to science. Five species represent new geographic or depth records. A list of Mexican localities previously and newly reported for each species are plotted on distribution maps. Environmental data, i.e., depth, temperature, and dissolved oxygen measured at the bottom level during the survey are provided. When compared with other areas of the world, the reduced number of specimens collected during this survey could be linked to the limiting effect of the Pacific Mexico Oxygen Minimum Zone. An updated checklist of species of Holothuroidea recorded below 350 m depth along the Pacific coast of Mexico is also provided totaling 31 species: 13 of these occur in the California Current area, 20 in the Gulf of California, and 15(16) along the SW coast of Mexico.
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Revista Mexicana de Biodiversidad 82: 413-443, 2011
Recibido: 15 abril 2010; aceptado: 28 enero 2011
Deep-water Holothuroidea (Echinodermata) collected during the TALUD cruises
off the Pacic coast of Mexico, with the description of two new species
Holothuroidea (Echinodermata) de mar profundo recolectadas durante las campañas TALUD
frente a la costa del Pacíco mexicano, con la descripción de dos especies nuevas
Claude Massin1 and Michel E. Hendrickx2*
1Department of Recent Invertebrates, Royal Belgian Institute of Natural Sciences, Rue Vautier 29, Brussels, B-1000, Belgium.
2Unidad Académica Mazatlán, Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, PO Box 811, 82000 Mazatlán,
Sinaloa, México.
Abstract. Research cruises aboard the R/V “El Puma” were organized to collect deep-water benthic and pelagic
specimens off the Pacic coast of Mexico. Seventy four specimens of Holothuroidea were collected off the Pacic coast
of Mexico in depths of 377-2 200 m. The collection includes representatives of 5 of the 6 orders of Holothuroidea, 3
Dendrochirotida, 2 Dactylochirotida, 2 Aspidochirotida, 4 Elasipodida and 2 Molpadiida. Apodida were not represented.
Of the 13 species recognized, 11 were identied to species level and 2, belonging to the genera Ypsilocucumis Panning,
1949, and Mitsukuriella Heding and Panning, 1954, are new to science. Five species represent new geographic or depth
records. A list of Mexican localities previously and newly reported for each species are plotted on distribution maps.
Environmental data, i.e., depth, temperature, and dissolved oxygen measured at the bottom level during the survey are
provided. When compared with other areas of the world, the reduced number of specimens collected during this survey
could be linked to the limiting effect of the Pacic Mexico Oxygen Minimum Zone. An updated checklist of species of
Holothuroidea recorded below 350 m depth along the Pacic coast of Mexico is also provided totaling 31 species: 13
of these occur in the California Current area, 20 in the Gulf of California, and 15 (16) along the SW coast of Mexico.
Key words: Holothuroidea, deep-water community, continental slope, East Pacic, Mexico, new species.
Resumen. Una serie de campañas oceanográcas fue organizada a bordo del B/O “El Puma”, frente a las costas del
Pacíco mexicano con el propósito de recolectar ejemplares de la fauna bentónica y pelágica de aguas profundas. La
recolección incluyó representantes de 5 de los 6 órdenes de Holothuroidea, i.e., 3 Dendrochirotida, 2 Dactylochirotida,
2 Aspidochirotida, 4 Elasipodida y 2 Molpadiida. Los Apodida no están representados. De las 13 especies capturadas
por debajo de los 350 m de profundidad (377-2 200 m), 11 fueron identicadas hasta especie y 2 pertenecientes a los
géneros Ypsilocucumis Panning, 1949, y Mitsukuriella Heding y Panning, 1954, son nuevas para la ciencia. El material
examinado comprende 74 ejemplares. Las localidades previas y nuevas registradas para cada especie recolectada están
compiladas para el Pacíco mexicano en mapas de distribución. Se proporciona información acerca de las condiciones
de captura de cada especie (temperatura y oxígeno disuelto). Comparativamente con otras áreas del mundo, el número
reducido de organismos recolectados durante el estudio podría estar relacionado con la presencia de una zona del
mínimo de oxígeno a lo largo del Pacíco mexicano. Se anexa una lista actualizada de las especies de Holothuroidea
registradas en profundidades mayores a 350 m frente a la costa del Pacíco mexicano. En total, 31 especies están
registradas: 13 en el área de la corriente de California, 20 en el golfo de California y 15 (16) a lo largo de la costa SO
de México.
Palabras clave: Holothuroidea, comunidad de aguas profundas, talud continental, nuevas especies, Pacíco este,
Deep-sea macroinvertebrate communities are
characterized by high diversity values (Hessler and
Sanders, 1967; Sanders and Hessler, 1969; Grassle, 1989;
Smith et al., 1998). In areas where the oxygen-minimum
zone (OMZ) intercepts the continental slope, anoxic and
severely hypoxic benthic fringes are species-poor, but in
even deeper water the hypoxic zone is species-rich. In the
OMZ, depth and dissolved oxygen concentration are the
most important factors affecting deep water community
composition (Rosenberg et al., 1983; Levin and Gage,
1998; Rogers, 2000; Hendrickx, 2001; Levin et al., 2001;
669.indd 1 14/06/2011 03:42:43 p.m.
414 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
McClain, 2004; Méndez, 2006; Zamorano et al., 2006) and
species size (Rex and Etter, 1998; McClain and Rex, 2001;
Olabarria and Thurston, 2003, 2004).
Study of Mexican echinoderms in the East Pacic
started with the contributions of A.E. Verrill, who studied
material collected along the west coast of America,
including specimens from the area of La Paz (Verrill,
1868), Cabo San Lucas and the Gulf of California (Verrill,
1870, 1871b). He also reviewed large collections obtained
during surveys covering much larger geographic areas (e.g.,
Verrill, 1867, 1871a), and described several new taxa. The
Holothuroidea collected by the “Albatross” (west coast of
America, from Peru to California, including Mexico) were
studied by Ludwig (1893, 1894) and H.L. Clark (1913,
1920, 1923). Ludwig’s monograph (1894) is by far the
most important contribution for deep-water Holothuroidea
in the region, with a total of 21 species described that
occur below 350 m depth. Several other contributions
complement the study of deep-water Holothuroidea in the
central eastern Pacic, with the description of 14 species:
Müller (1850) described 1 new species, Mitsukuri (1912)
1, H.L. Clark (1913, 1920, 1923) 5, Deichmann (1938a) 1,
Cherbonnier (1941) 1, Belyaev (1971) 2, Hansen (1975)
2, and Pawson (1983) 1. Other contributions, essentially
by Deichmann (1937, 1938a, 1938b, 1938c), increased the
number of deep-water species known to the region.
Several papers, principally based on literature, present
compilation lists of holothurians from the central eastern
Pacic (see Table 2). Parker (1964) listed material collected
in the Gulf of California and in several other sites along the
west coast of Mexico, including 12 species of deep-water
Holothuroidea. Bergen (1980) listed material collected in
Southern California that includes 5 deep-water species.
Maluf (1988) compiled thoroughly all the information
on Central eastern Pacic echinoderms (list of species,
depth distribution, geographical range). According to
Maluf (1988), there were 55 species of Holothuroidea
occurring below 350 m depth. Two deep-water species
cited in the literature and of dubious status were also cited
by Maluf (1988: 167) in an annex but were not included
in her general analysis: Achlyonice ecalcarea Théel, 1879
(off Baja California, 1598 m) and Bathyplotes hancocki
Domantay, 1961 (Southern California and Gulf of
California). Another species, Penagione leander Pawson
and Foell, 1986, an abyssal benthopelagic sea cucumber,
was omitted by Maluf (1988) and therefore increases the
list of species known from the Central eastern Pacic to
56. Of these 56 species, 5 were originally described from
other regions of the world and later reported for the Central
eastern Pacic: Synallactes ishikawai Mitsukuri, 1912
(now a junior synonym of S. sagamiensis Augustin, 1908),
from Japan; Leptosynapta albicans (Selenka, 1867), from
the North Atlantic; Rynkatorpa duodactyla (H.L. Clark,
1907) (as Protankyra duodactyla), from the NW Pacic;
Ceraplectana trachyderma H.L. Clark, 1907, from the
NW Pacic; and Molpadia musculus Risso, 1826, from the
Maluf (1991) proposed a checklist of echinoderms
reported from the Galapagos Islands, including 38
Holothuroidea, 16 with records below 350 m depth.
Lambert (1996) presented a recapitulation table with
distribution, habitat and morphological data for all 8
species of Psolidium known from the eastern Pacic. Of
these 8 species, only 2 (P. panamense Ludwig, 1894, and
P. gracile Ludwig, 1894) occur deeper than 350 m (at
2 323 m, both at the same and unique known locality,
in the Gulf of Panama), and both were cited by Maluf
(1988). Solís-Marín et al. (1997) presented a compilation
of species of echinoderms known from the Bay of La Paz.
Mostly based on literature records, the list includes 27
Holothuroidea with 3 species occurring below 500 m.
Further north, a large series of exploratory surveys were
performed off the coast of California and Oregon, USA.
Echinoderms, particularly Holothuroidea, were often
identied to species level. A review of the information
available was presented by Nybakken et al. (1998), who
also reported on 13 species of Holothuroidea collected
or observed by either a beam trawl or a camera sled
off central California, many of which were previously
reported for Mexico.
Additional information related to deep-water
Holothuroidea from off the Galapagos was provided by
Pawson and Ahearn (2001). More recently, Maluf and
Brusca (2005) have reported 57 species of Holothuroidea
for the Gulf of California, most from shallow water, and
only 9 deep-water (>350 m depth) species are included in
their list. Solís-Marín et al. (2005) published a compilation
of echinoderm species from the Gulf of California. This
list, based exclusively on records veried in the Dra.
María Elena Caso Muñoz (Universidad Autónoma de
México, México City) and in the Smithsonian Institution
(Washington D.C.) echinoderms collections, includes
45 species of Holothuroidea from all kinds of habitats.
Subsequently, Honey-Escandón et al. (2008) presented a
list of species for the Mexican Pacic (i.e., excluding the
Gulf), based on the same collections, with 46 species of
Holothuroidea, and Solís-Marín et al. (2009) summarized
our present knowledge of the Gulf of California
Holothuroidea in a monograph that includes 55 species.
Many contributions to our knowledge of Mexican
echinoderms were authored by M. E. Caso. Because
she worked exclusively on intertidal and shallow-water
holothurians (see among others Caso 1964, 1966, 1968),
her contributions will rarely be cited in this study.
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Revista Mexicana de Biodiversidad 82: 413-443, 2011
Materials and methods
Holothuroidea were collected between 377 and 2 200
m depth on the continental slope along the Pacic coast of
Mexico using an Agassiz dredge (ca 2.7 m width) and a
benthic sledge (2.35 m width, 0.95 m high) equipped with a
collecting net of ca 5.5 cm (2 1/4”) stretched mesh aperture
and an internal net with a ca 2.0 cm (3/4”) stretched mesh
aperture. A total of 13 cruises were organized in the Gulf
of California and along SW Mexico, south of Banderas
Bay, from 1989 to 2008. Specimens of Holothuroidea
were collected during 9 cruises: TALUD III, September
1991; TALUD VI, March 2001; TALUD VII, June 2001;
TALUD VIII, April 2005; TALUD IX, November 2005;
TALUD X, February 2007; TALUD XI, June 2007; TALUD
XII, March-April 2008; and TALUD XIII, January 2009.
Positional coordinates for each station were plotted using
a GPS navigation system. Depth was measured with an
EdoWestern, analog recorder (TALUD III-VIII) or a digital
recorder (TALUD IX-XIII). Epibenthic water temperature
was measured with a Seabird CTD, and dissolved oxygen
content was measured by the Winkler method (all cruises)
and with a probe attached to the CTD (TALUD VIII-XIII).
Holothuroidea were xed on board with a 4% formaldehyde
sea water solution, washed with tap water and preserved in
70% ethanol. In the systematic section below, a restricted
synonymy including the prime synonym, references to
new combination or reviews, and references for the Pacic
coast of Mexico are included for each species, together with
comments and additional data related to their distribution
and ecology. The material examined herein is deposited in
the Regional Collection of Invertebrates (EMU), ICML,
UNAM, Mazatlán, Sinaloa, Mexico, in the collection of
Holothuroidea at the Royal Belgian Institute of Natural
Sciences (IG/HOL/RBINS), Brussels, Belgium, and in the
Colección Nacional de Equinodermos “Dra. María Elena
Caso” (ICML-UNAM), in México D.F., Mexico. Material
examined also include 2 lots received on loan from the
Scripps Institution of Oeanography, University of California
San Diego (SIO, UCSD), La Jolla, USA, and a series of
ossicle preparations from the Zoology Museum of Hamburg
(ZMH), Hamburg, Germany. Other abbreviations used are:
L, total length; St., sampling station: id.; identicator; coll.,
collector. Based primarily on Maluf (1988) contribution,
a list of all species with at least 1 record within Mexican
waters of the Pacic Ocean was established (Table 1). In
most cases the records used by Maluf were checked in the
original literature. Literature records posterior to 1988 or
based on the material collected during the TALUD cruises
were incorporated in this list. The sources of these records
are indicated in table 1.
Generality and abiotic factors. During the TALUD
cruises, a total of 135 localities were sampled with benthic
sledges and Holothuroidea were captured in 25 of these.
No specimens of Holothuroidea were caught during
the TALUD I, II, IV and V cruises. A total of 13 species
represented by 74 specimens were collected. Eleven were
identied to species level, 2 of these recognized as new
species. The collection includes representatives of 5 of
the 6 orders of Holothuroidea, i.e., 3 Dendrochirotida, 2
Dactylochirotida, 2 Aspidochirotida, 4 Elasipodida and 2
Molpadiida. Apodida were not represented. In 23 stations
only 1 species was collected. A maximum of 3 species
were found in a single station and 2 in another. All species
considered, the material obtained in this survey was caught
in a depth range of 377 to 2 200 m, and in epibenthic
temperature and dissolved oxygen ranges of 2.0 to 10.52
°C and <0.05 to 2.14 ml O2/l, respectively (Table 1).
Class Holothuroidea de Blainville, 1834
Order Dendrochirotida Grube, 1840
Family Cucumariidae Ludwig, 1894
Genus Abyssocucumis Heding, 1942
Abyssocucumis albatrossi (Cherbonnier, 1941)
Figs. 1 A-F, 2
Cucumaria albatrossi Cherbonnier, 1941: 93-103, g. 2,
g. 3 a-h, i, k-m.
Abyssocucumis albatrossi; Cherbonnier, 1947: 462;
Panning, 1949: 454; Madsen, 1955: 165, 167; Carney and
Carey, 1976: 69; Maluf, 1988: 92, 155; Nybakken et al.,
1998: 1778.
Cucumaria abyssorum; Ludwig, 1894; 122, pl. XIII, gs.
1-5; Ludwig and Heding, 1935: 179, textg. 42.
Staurocucumis abyssorum; Hansen, 1988: 302, Fig. 1;
Solís-Marín et al., 2009: 84, pl. 17A-E.
Non: Cucumaria abyssorum Théel, 1886: 66.
Material examined. One specimen (L = 66 mm), TALUD
XIII, St. 37 (EMU-8630).
Body cylindrical, 66 mm long, 25 mm across, slightly
tapering at both extremities. Mouth and anus terminal.
Tentacles fully retracted. Colour in alcohol grayish to
black. Body wall gritty to the touch, parchment like, 0.7
mm thick. Tube feet only in the ambulacral zone aligned
in 2 rows. Ossicles of the body wall as cross-shaped
669.indd 3 14/06/2011 03:42:43 p.m.
416 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
Cruise Station Species Position Date Depth Temp. Oxygen
(m) °C ml/l
TALUD III St. 14B Molpadia musculus 24°39’12”N, 108°37’54”W 8-Sep-1991 1 188-1 208 ND 0.4
TALUD VI St. 25 Molpadia intermedia 24°51’41”N, 108°57’53”W 16-Mar-2001 830-850 ca 5.0 0.22
TALUD VI St. 26 Ypsilocucumis californiae 24°56’17”N, 109°6’39”W 16-Mar-2001 1 190-1 270 3.7 1.4
TALUD VIII St. 3 Pannychia moseleyi 24º32’36”N, 109º30’30”W 16-Apr-2005 1 100 3.0 0.39
TALUD VIII St. 11 Ypsilocucumis californiae
Synallactes alexandri
Pannychia moseleyi
24°54’24”N, 110°25’30”W 17-Apr-2005 920 50.2
TALUD VIII St. 16 Ypsilocucumis californiae
Synallactes virgulasolida
25°24’48”N, 110°34’48”W 18-Apr-2005 1 030 50.2
TALUD VIII St. 20 Dendrochirotida sp. und. 25°56’54”N, 110°45’0”W 19-Apr-2005 1 150 4 0.30
TALUD VIII St. 22 Elasipodida sp. und. 26°03’42”N, 110°21’18”W 19-Apr-2005 2 200 2.0 1.27
TALUD IX St. 10 Pannychia moseleyi 24°56’24”N, 110°16’42”W 12-Nov-2005 969-1 225 3.6 0.11
TALUD IX St. 16 Molpadia musculus 25°23’48”N,110°36’42”W 13-Nov-2005 997-1 021 4.6 0.15
TALUD IX St. 17 Psolidium gracile 25°20’48”N, 110°46’30”W 13-Nov-2005 826-836 5.75 <0.05
TALUD X St. 3 Ypsilocucumis californiae
Psolidium gracile
28°16’40”N, 112°35’10”W 9-Feb-2007 377-379 10.52 1.05
TALUD X St. 9 Mitsukuriella unusordo 27°52’51”N, 112°15’53”W 10-Feb-2007 1 205-1 215 3.77 0.32
TALUD XI St.1 Psolidium gracile 16°52’N, 100°20’W 7-Jun-2007 740-790 5.9 <0.05
TALUD XII St. 5 Ypsilothuria bitentaculata 16°58’17”N, 100°55’12”W 28-Mar-2008 1 925-1 977 2.6 1.43
TALUD XII St. 8 Elasipodida sp. und. 17°04’09”N, 101°39’16”W 29-Mar-2008 1 880-1 940 2.35 1.99
TALUD XII St. 9 Laetmogone scotoeides 17°10’15”N, 101°37’23W 28-Mar-2008 1 392-1 420 3.28 1.01
TALUD XII St. 10 Laetmogone scotoeides 17°11’18”N, 101°28’30W 29-Mar-2008 1 180-1 299 3.73 0.68
TALUD XII St. 15 Benthodytes cf. sanguinolenta 17°25’33”N, 102°07’20”W 30-Mar-2008 2 015-2 080 2.16 2.14
TALUD XII St. 25 Pannychia moseleyi 18°26’45”N, 104°16’10”W 1-Apr-2008 1 858-1 879 2.45 1.38
TALUD XII St. 27 Psolus aff. squamatus 18°40’28”N, 104°35’51”W 2-Apr-2008 1 040-1 095 4.26 0.26
TALUD XII St. 28 Psolus aff. squamatus 18°50’19”N, 104°34’14W 2-Apr-2008 1 101-1 106 4.25 0.36
TALUD XII St. 29 Pannychia moseleyi 19°19’37”N, 105°26’20”W 2-Apr-2008 1 609-1 643 2.82 1.38
TALUD XIII St. B Elasipodida sp. und. 26°17’04”N, 110°27’53”W 13-Jan-2009 1 295-1 330 3.62 0.87
TALUD XIII St. 37 Abyssocucumis albatrossi 25°59’30”N, 110°19’21”W 15-Jan-2009 2 056-2 195 2.56 1.68
Table 1. Sampling stations of the TALUD cruises where specimens of Holothuroidea were collected and list of species
per station. Position, depth, and epibenthic water temperature and dissolved oxygen concentration are indicated for each
station. Precision of data may vary according to the method used during the survey
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Revista Mexicana de Biodiversidad 82: 413-443, 2011
Species CC GC SWM OI
Psolidium gracile Ludwig, 1894 PS PS
Psolus cf. squamatus (O.F. Müller, 1776) PA PS MA PS
Abyssocucumis abyssorum (Théel, 1886) (1) PA MA HE MA MB SO1 PA MA HE
Abyssocucumis albatrossi (Cherbonnier, 1941) (2) PS SO3
Mitsukuriella unusordo sp. nov. PS
Ypsilocucumis californiae sp. nov. PS
Ypsilothuria bitentaculata (Ludwig, 1893) (3) PA MA MA MB MA PS
Paelopatides confundens Théel, 1886 MA MB
Pseudostichopus mollis Théel, 1886 PA MA MB PA MA
Bathyplotes sp. MA PA
Synallactes sagamiensis Augustin, 1908 (4) PA MA
Synallactes alexandri Ludwig, 1894 PS
Synallactes virgulasolida Massin and Hendrickx, 2010 PS
Oneirophanta mutabilis mutabilis Théel, 1979 PA PA MA
Laetmogone scotoeides (H.L. Clark, 1913) MA PS
Laetmogone wyvillethomsoni Théel, 1879 MA
Pannychia moseleyi Théel, 1882 MA MA MB SO1 SO2 SO3 PS MA PS
Psychronaetes hanseni Pawson, 1983 MA (Clarion)
Benthodytes sanguinolenta Théel, 1882 (5) PA MA HE PS
Psychropotes longicauda Théel, 1882 (6) PA MA PA MA
Peniagone intermedia Ludwig, 1894 MA(?)
Peniagone papillata Hansen, 1975 MA
Peniagone sp. PA
Scotoplanes clarki Hansen, 1975 SO3
Scotoplanes globosa (Théel, 1879) MA
Leptosynapta albicans (Selenka, 1867) MA (Socorro)
Molpadia granulata (Ludwig, 1894) (7) PA MA MB
Table 2. Species (total 31) of deep-water (> 350 m depth) Holothuroidea occuring off the coast of Mexico (northernmost
limit set at 32o28’16”N; southernmost limit set at 14o32’27’N), including the California Current area (CC), the Gulf of
California (GC), the area of souwestern Mexico, south of Banderas Bay (SWM), and the offshore islands (OI). Data
used in the table were taken from the following sources: PA, Parker (1964); MA, Maluf (1988); SO1, Solis-Marín et
al. (1997); SO2, Solis-Marín et al. (2005); SO3, Solis-Marín et al. (2009); MB, Maluf and Brusca (2005); HE Honey-
Escandón et al. (2008); PS, present study. Boldface: species collected during the TALUD cruises. (?) Dubious record. The
unidentiable specimens collected during the TALUD cruises are not included in this list (see text for further comments).
For convenience species are listed following Maluf’s (1988) sequence
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418 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
ossicles, 270-330 µm long, very spiny, with or without
perforation at the extremities of the arms (Fig. 1A). Most
of these cross-shaped ossicles have 4 arms, a few have 3 or
5 arms. In the tube feet same kind of ossicles (Fig. 1B, C),
somewhat larger (up to 450 µm), the most common with 2
long and 2 short arms (Fig.1B). Extremities of these arms
multi-perforated (Fig. 1C). In the tentacles 2 kinds of rods:
very thin, with strong spines on 1 side (Fig.1D), or with a
central apophysis, 180-250 µm long, and large straight to
V-shaped rods, smooth or spiny, up to 600 µm long, with a
central apophysis (Fig. 1E-F),
The present specimen ts particularly well with the
detailed description of Abyssocucumis albatrossi given by
Cherbonnier (1941).The spiny arms of the cross-shaped
ossicles are characteristic of the species whereas smooth
arms are characteristic of Abyssocucumis abyssorum
Théel, 1886. Ludwig (1894: pl. XIII, gs. 1-5) illustrated
an A. albatrossi, but named his material abyssorum.
Hansen (1988), who considered albatrossi a synonym of
abyssorum, moved abyssorum to the genus Staurocucumis.
Hansen considered that the cross-shaped ossicles are
characteristic of the juveniles and disappear in adults to be
replaced by perforated plates. However, the 60 mm long
specimen here examined is an adult with a well developed
gonad. Ludwig (1894) observed also only cross-shaped
ossicles in large specimens (up to 90 mm long) and no
perforated plates. Another argument to sustain the opinion
of Hansen (1988) is the fact that among the dendrochirotes
(particularly in the genus Staurocucumis) the ossicles
become more spiny with increasing body size (Massin,
1994). Indeed, in the paper of Cherbonnier (1941) the A.
abyssorum (smooth ossicle arms) are small specimens, <30
mm long, whereas the A. albatrossi (spiny ossicle arms) are
large specimens, 60 mm long. Abyssocucumis abyssorum
could thus be the juveniles of A. albatrossi. However, this
does not t with the presence of a well developed gonad
in both species, whatever the body size (Théel, 1886;
Ludwig, 1894; Cherbonnier, 1941). Moreover, small and
large specimens of A. abyssorum present smooth ossicle
arms (Théel, 1886; Ludwig, 1894; Heding, 1942). Spiny
or smooth ossicle arms seem to not be related to body
size. Consequently, we consider that A. albatrossi and A.
abyssorum are well separated species, both belonging to
the genus Abyssocucumis as dened by Heding (1942).
The Staurocucumis abyssorum illustrated by Solís-
Marín et al. (2009: 84, pl. 17, Fig. B) is, according to us, a
specimen of Abyssocucumis albatrossi because of the very
spiny arms of the cross-shaped ossicles.
Distribution. Records in Mexico. Known from the type
locality (“Albatross” St. 3414, 10o14’N, 96o28’W) located
about 500 km off the coast of Chiapas, SW Mexico; 4 085
m (2 232 fm) (Fig. 2). According to Cherbonnier (1941),
the 2 specimens he examined had been collected at the
“Albatross” St. 3414, located off SW Mexico (above). He
also stated, however, that A. albatrossi is found in the Gulf
Molpadia intermedia (Ludwig, 1894) MA HE MA MB SO1 SO3 PS
Molpadia musculus Risso, 1826 MA HE PA MA SO1 SO3 MB PS PA MA
Heldingia californica (Ludwig, 1894) (8) MA SO1 SO3
Paracaudina chilensis (J. Müller, 1850) MA SO2 MA
(1) Also cited in the genus Staurocucumis.
(2) See text for further details.
(3) Previously in the genus Sphaerothuria.
(4) Cited as S. ishikawai f. ind.
(5) Not cited by Maluf and Brusca (2005).
(6) Cited by Parker (1964) as P. dubiosa Ludwig, 1894, and P. raripes Ludwig, 1894, both junior synonyms of P. longicauda.
(7) Cited by Parker (1964) as M. granulosa.
(8) Also cited in the genus Caudina.
Species CC GC SWM OI
Table 2. Continues
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Revista Mexicana de Biodiversidad 82: 413-443, 2011
of Panama (1o05’N, 29o40’N) and in the Gulf of California
(27o34’N, 110o53’40”W), at 1 466-3 615 m depth, but it
seems he did not actually examine material from these
2 localities. According to Maluf (1988), A. albatrossi is
known only from off Peru (ca 6oS) and from a (doubtful)
locality off California (ca 32oN), in depths of 1 585-5 690
m. In addition to the original description by Cherbonnier
(1941), Maluf (1988) only cited the reference of Madsen
(1955) as sources for the geographic and bathymetric
ranges of A. albatrossi, and the data provided by her are
exactly the same as those provided by Madsen. Although
much more detailed than the information provided by
Madsen, the compilation presented by Maluf (1988) does
not include the Chiapas (type locality) record, which is
rather surprising. We were not able, however, to trace the
records for California, the Gulf of California and Peru cited
by Cherbonnier (1941) and Maluf (1988). Abyssocucumis
albatrossi has been reported from Oregon, NE Pacic,
in a depth range of ca 2 700-4 000 m, and appears to
dominate the holothurian community below 3 000m
(Carney and Carey, 1976; not cited by Maluf). It was
also recently reported by Nybakken et al. (1998) during
a survey off central California, but this Californian record
could obviously not be used as a source of information by
earlier authors. The geographic range of A. albatrossi is
provisionally set as off Oregon, USA, to off the coast of
Chiapas, Mexico, including the central Gulf of California,
Mexico (Fig. 2).
Family Psolidae R. Perrier, 1902
Genus Psolus Oken, 1815
Psolus aff. squamatus (O.F. Müller, 1776)
Fig. 2
Holothuria squamata O.F. Müller, 1776: 232.
Cuvieria squamata; Koren, 1845: 211, pls. 2, 3.
Psolus squamatus; Lütken, 1857:14, 69, 81, 104; Ludwig,
1900: 158 (list of citations); Vaney, 1906a: 27, pl. 2, gs.
16a-c, 17a-c; Mitsukuri, 1912: 225, pl. 7, gs. 61, 62,
textg. 42 ( list of citations); Ohshima, 1915: 280; Ekman,
1923: 1-56 (passim), gs. 12-14, 20-24, 26-27, 29-30,
36-37; Bergen, 1980: 275; Imaoka, 1980: 361, gs. 1-9;
Maluf, 1988: 88, 152 (in part, probably not the record of P.
squamatus segregatus Perrier, 1905); Lambert, 1997: 51,
gs. 21, 22; Maluf and Brusca, 2005: 343.
Psolus pauper Ludwig, 1894: 139.
Psolus valvatus Östergren, 1904: 659.
Material examined. Two specimens (L = 68 mm,
EMU-8609; L = 64 mm, IG 31487/HOL 1511 RBINS/
HOL/738992), TALUD XII, St. 27. One specimen (L = 45
mm), TALUD XII, St. 28 (EMU-8610).
Additional material examined. Three specimens (L
= 10, 16, 27 mm), SW of Punta Banda (31°38’18”N,
116°51’24”W), Baja California, 12/February/1960, 183-
458 m (coll. R. Parker: id. E. Deichmann) (SIO, UCSD,
SOB1-31-6333). Ossicle preparations from a specimen of
Psolus squamatus segregatus, Patagonia (ZMH, E 4170).
Body elongate to nearly rounded. Dorsal surface
covered by large overlapping scales; ventral surface thick
with a double row of tube feet along the margin. Midventral
radius without tube feet or with a few at the front and at
the rear. Mouth and anus dorsal, each at the top of a cone
covered by irregular, elongated plates. No clearly dened
oral or anal valves. Dorsal scales multilayered, 0.6 to 8.0
mm across, covered or not with rounded or ovoid granules.
When present, granules are 150-250 µm across, whatever
the size of the specimen. In the ventral sole small perforated
plates with 2-9 large holes in the smallest specimen and
3-5 in the large specimens. Largest plates with a few
knobs. Size of the plates from 150 to 250 µm across in
small specimens to 130-165 µm across in large specimens.
Ossicles of tentacles smooth, perforated plates, rounded,
triangular or elongated, slightly curved, 100-500 µm long.
In the tube feet elongated perforated plates, 160-320 µm
long; end plate in 1 piece, 240-450 µm across.
Vaney (1906a), Ekman (1923), and Deichmann
(1941) considered P. squamatus segregatus as a valid
species. The characters separating P. squamatus from
P. squamatus segregatus are: ossicles of the ventral
sole (large perforated plates, 150-300 µm across,
with large holes vs. small massive perforated plates,
75-110 µm across, with small holes, respectively) and
the size of the dorsal granules (150-250 µm across vs.
330-470 µm across, respectively) (Ekman, 1923: Fig.
33). The specimens examined t particularly well with
P. squamatus as far as the body size and the general
aspect of dorsal and ventral ossicles are concerned. H.L.
Clark (1913), studying material from California, came
to the same conclusion. However, H.L. Clark (1913),
Mortensen (1927), Madsen (in Maluf, 1988), and Madsen
and Hansen (1994) considered that the P. squamatus
from Norway and the East Pacic are not conspecic.
Contrary to the specimens from the Gulf of California,
those from Japan show an increase in the number of
holes of the ventral perforated plates with increasing
body size (Imaoka, 1980). The small specimen from
Japan (L = 41 mm) is very close to the small specimen
(L = 45 mm) from SW Mexico examined herein (EMU-
669.indd 7 14/06/2011 03:42:44 p.m.
420 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
8610), particularly in the ossicles of the tentacles and
the tube feet. The species is also reported to be present
in the Pacic waters from Japan (Imaoka, 1980) to
Patagonia (Pawson, 1969), through Canada (Lambert,
1997). For the specimens from Patagonia, R. Perrier
(1905) erected the variety segregatus. Ekman (1923)
and Deichmann (1941) consider that this variety extends
from Patagonia to the Bering Sea. According to Ekman
(1923) and Imaoka (1980), P. squamatus is a highly
variable species. Moreover, most of the citations in the
literature are restricted to a name without description
and/or illustrations. In these circumstances, it is very
difcult to establish the limit of its distribution. Parker
(1964: 165) reported a capture of Psolus squamatus var.
segregatus from the west coast of Mexico (ca 183-458 m
depth, SW of Ensenada, off Banda Point; 31° 38’20”N,
116° 51’24”W), in the California Current area. The
material was presumably identied by E. Deichmann.
The same lot was later deposited at Scripps Institution
of Oceanography and included by Luke (1982: 56) in
his catalogue of echinoderms as Psolus squamatus.
Three of the 117 specimens contained in this lot were
examined during this study. Unfortunately, this material
corresponded to juveniles that had been previously kept
dry and ossicles could not be properly examined, thus
making a positive identication impossible.
We are convinced that P. squamatus and P. squamatus
segregatus are distinct species. Considering the very
wide distribution range of P. squamatus s.l., they could
even represent more than 2 species. Psolus squamatus
segregatus seems to be restricted to the Pacic coast of
South America. We believe that this taxonomical problem
will not be solved until material preserved for DNA analysis
from Japan, Canada, Central and South America, and the
North Atlantic is available. Such a study is obviously
beyond the scope of the present paper.
Distribution. Records in Mexico. Type locality of Psolus
pauper, “Albatross” St. 3424 (21°15’N, 106° 23’W), off
Tres Marías Islands, SW Gulf of California; 1 237 m (676
fm), 3.3oC (Ludwig, 1894). H.L. Clark (1923: 161) reported
on a single specimen (L = 55 mm) of Psolus squamatus,
collected off California, “Albatross” St. 5695 (33°33’N,
120°17’W), in 977 m depth (534 fm). Probably restricted
to the East Pacic, north of Central America to the Bering
Sea. Present records extend the Mexican distribution of
this species to SW Mexico (18o40’28”N) (Fig. 2).
Psolidium Ludwig, 1887
Psolidium gracile Ludwig, 1894
Figs. 2, 3A-G, 4A-F, pl. 1A-D
Psolidium gracile Ludwig, 1893: 184 (nomen nudum).
Psolidium gracile Ludwig, 1894: 132, pl. XIII, gs. 17-19;
Maluf, 1988: 88, 152; Lambert, 1996: table; Nybakken et
al., 1998: 1760.
Material examined. Six specimens (L = 13-17 mm, EMU-
8611; L = 16 and 17 mm, IG 31487/HOL 1512 RBINS/
HOL/738988), TALUD IX, St. 17. Seventeen specimens
(L = 12-18 mm), TALUD XI, St. 1 (EMU-8612). Three
specimens (L = 15-17 mm), TALUD X, St. 3 (EMU-8624).
Small species 13-18 mm long, 6-11 mm wide and 5
mm height (pl. 1 A-C). Ventral sole more or less 70% of
body length, packed with ossicles visible to naked eyes.
One row of 30-35 tube feet on each side of the ventral sole
(pl. 1D). Along the mid-ventral radius 1 row of a 10 of tube
feet located mainly at the rear and at the front, more widely
spaced out at mid-body. Dorsal tube feet very small, easily
overlooked. Dorsally, very large, rounded scales up to 1
200 µm across (pl. 1C-D); 16-20 scales between mouth
and anus. No valves closing mouth and anus. Dorsal scales
180-1 200 µm across, small ones 180-350 µm across and
made of 1 layer (Fig. 3B), large ones half or ¾ of their
surface with 1 layer and the remaining surface with 2 layers.
Calcareous ring simple, without posterior processes; radial
and interradial plates of the same width but radial plates
nearly twice the height of the interradial (Fig. 3A). Ventral
sole packed with rounded, perforated plates (3-15 holes),
125-290 µm across (Fig. 3C), surface of plates smooth,
edge slightly knobbed or with blunt spines (Fig. 3C). In
large specimens, ventral plates larger, more elongate, up
to 450 µm long, perforated by 4-19 holes (Fig. 3D). In the
ventrolateral tube feet, curved rods 120-250 µm long (Fig.
3F), perforated plates (250-360 µm long) (Fig. 3G), and a
small end-plate (170-220 µm across) (Fig. 3E). Tentacles
with spiny curved rods, 200-400 µm long, perforated at
the extremities by 1-9 small holes (Fig. 4A); also a few
perforated plates, 330-350 µm long with large holes (Fig.
4B-C). In the gonads, spiny rods, 150-300 µm long (Fig.
4D-F); spines often ending in 2-3 spinules.
According to Maluf (1988) there are 6 species of
Psolidium along the Central eastern Pacic coast. Two
deep-water (P. panamense Ludwig, 1894, and P. gracile)
and 4 shallow-water species (Psolidium dorsipes Ludwig,
1886; P. ekmani Deichmann, 1941; P. eubullatum
Deichmann, 1941; and P. planum Deichmann, 1941).
The specimens here observed t particularly well with
Psolidum gracile. Only the number of tube feet in the mid-
ventral radius and the size of the dorsal scales are different
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Revista Mexicana de Biodiversidad 82: 413-443, 2011
from the holotype. Unfortunately, the material reported
by Nybakken et al. (1998) from off central California
was not illustrated or described, and the species is known
only from the description of the holotype. The differences
observed in the TALUD specimens could be ascribed to
species variability.
Distribution. No previous records in Mexico. Known
from the type locality, “Albatross” St. 3392 (7o05’30” N,
79o40’W), Cabo Mala, Gulf of Panama, 2 323 m (1 270
fm) (Lambert, 1996), and off central California, between
2 300 and 3 075 m depth (Nybakken et al., 1998). The
material collected during the TALUD survey was obtained
in much shallower water (377-920 m) and represents the
rst record for the Pacic coast of Mexico (off north of
Acapulco and off southern Baja California) (Fig. 2).
Dendrochirotida sp. und.
Material examined. One specimen (not measured),
TALUD VIII, St. 20 (EMU-8625).
The specimen is damaged and completely eviscerated,
calcareous ring included. The presence of 5 longitudinal
muscle bands and of retractor muscles of the pharynx
indicate that it belongs with the Dendrochirotida.
Unfortunately, the absence of ossicles in the tissues does
not allow for further identication.
Order Dactylochirotida Pawson and Fell, 1965
Family Vaneyellidae Pawson and Fell, 1965
Genus Mitsukuriella Heding and Panning, 1954
Mitsukuriella unusordo sp. nov.
Figs 5A-F, 6, 7, pl. 1E-G
Material examined. Holotype (L = 13.4 mm), TALUD X,
St. 9 (EMU-8629).
Small specimen 13 mm long, cylindrical, tapering at
both ends (pl. 1E-F), 4 mm across at the rear, 2 mm across
at the front and 4 mm across at the end. Mouth and anus
terminal. Mouth surrounded by 15 digitiform tentacles (pl.
1G), 5 large and 10 very small, a pair of small tentacles
between 2 large tentacles. Anus surrounded by 5 anal
papillae. Tube feet on 1 row in each radius (pl. 1E), more
crowded and longer at mid-body (pl. 1F). Dorsally, 15
tube feet on radii, and 25-30 per radius ventrally. Color
in alcohol yellow-grayish. Body wall gritty to the touch,
very thin, packed with large perforated plates visible to the
naked eye. Calcareous ring without posterior processes;
gonad present as a few undivided tubules.
In the body wall large, elongated, perforated plates,
375-850 µm long (Fig.5A-B), smaller plates smooth (Fig.
5A), larger plates partly knobbed (Fig. 5B).
Ossicles of the tube feet small perforated plates with
2-4 holes, 90-125 µm long (Fig. 5C-D). A few larger
plates, 160 µm long, with up to 8 holes and partly knobbed
(Fig.5D). At the apex of the tube feet a small perforated
end-plate (Fig. 5E) surrounded by V-shaped ossicles 100-
190 µm long, with a central apophysis (Fig. 5F). Ossicles
of the tentacles spiny curved rods, 130-200 µm long (Fig.
6), with 1 hole at each extremity, spines large, blunt.
Etymology. From the Latin unus”, “one”, and “ordo”,
row”, in reference to to the single row of tube feet in each
The presence of 15 digitate tentacles and of a calcareous
ring without posterior processes are characteristic of the
family Vaneyellidae as dened by Pawson and Fell (1965)
and of the genus Mitsukuriella. According to Heding and
Panning (1954) 2 species are known in this genus: M.
squamulosa (Mitsukuri, 1912) from Japan, and M. inexa
(Koehler and Vaney, 1908) from India. Mitsukuriella
unusordo sp. nov. is easily separated from M. squamulosa
by the presence of knobs on the large perforated plates,
and from M. inexa by the presence of a single row of
tube feet in each radius and by the shape of the large plates
(elongated vs. rounded, respectively).
Distribution. Known only from the type locality (Fig. 7),
Mitsukuriella unusordo sp. nov. has been collected deeper
(1 205-1 215 m) than the 2 other species of the genus: 250
m depth for M. squamulosa and 170 m for M. inexa.
Family Ypsilothuriidae Heding, 1942
Genus Ypsilocucumis Panning, 1949
Ypsilocucumis californiae sp. nov.
Figs 7, 8A-F, pl. 1H
Material examined. Eight specimens from the Gulf of
California. Holotype, TALUD VI, St. 26 (L = 15 mm)
(EMU-8613). Paratypes. Two specimens (L = 12 and 15
mm), TALUD VIII, St. 11 (EMU-8614), 3 specimens (L =
13, 14 and 21 mm), TALUD VIII, St. 16 (ICML-UNAM
5.177.0), and 2 specimens (L = 13 and 17 mm), TALUD X,
St. 3 (IG 31487/HOL 1514 RBINS/HOL/738995).
Specimens spherical, with 2 dorsal tubes (1 oral and
1 anal) contracted and well separated from each other
(pl. 1H). Length of specimens (tubes not included) from
669.indd 9 14/06/2011 03:42:44 p.m.
422 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
13 to 21 mm. Distance between mouth and anus from 7
to 12 mm. Body wall bristle, thin, translucent and gritty
to the touch, packed with large scales (visible to naked
eye) with spires protruding outside. Tube feet very small,
difcult to observe, apparently only present along the
radius. Calcareous ring brittle, very thin, with radial and
interradial plates of the same size and without posterior
process (Fig. 8A); anterior process of radial plates with a
deep groove for the insertion of the retractor muscles of
the introvert.
All specimens contracted, with the introvert inside.
After dissection 2 very long (at least 2 mm long when
contracted) digitiform tentacles observed. Other tentacles
minute, not counted. Retractor muscles of introvert very
thin, attached to the body wall at mid-body length. Gonad
present, reduced to a few, short, divided tubules, some
containing very large oocytes.
Body wall with large scales, developing from small,
single layered perforated plates, 200-700 µm across,
without pillar (Fig. 8B), to 2-layered perforated scales
(edge of the scale very often single layered) with an
eccentric spire (Fig. 8C). Scales are 700 to 1 000 µm
across, their spire, made of the fusion of several spiny
pillars, 350-400 µm high (Fig. 8D). In the long digitiform
tentacles curved rods, 150-370 µm long, with lateral blunt
spines and enlarged perforated extremities (Fig. 8E), most
rods very thin (± 10 µm across), a few thicker (20-25 µm
across) (Fig. 8 F).
The specimens at hand t well with the diagnosis of
the family Ypsilothuriidae. The presence of multilayered
scales, the eccentric position of the spire of the scales,
and the retractile oral and anal cones clearly indicate that
the new species belongs to the genus Ypsilocucumis (see
Panning, 1949) which included 3 species: Ypsilocucumis
asperrima Théel, 1886, Y. turricata (Vaney, 1906),
and Y. scotiae (Vaney, 1906). The latter 2 species,
erected by Vaney (1906b), have since been moved by
O’Loughlin (2002) and O’Loughlin et al. (2009) to the
genera Paracucumis Mortensen, 1925 and Crucella
Gutt, 1990, respectively. Ypsilocucumis californiae sp.
nov. differs from Y. asperrima in the size of the scales
(700-1 000 µm vs. >2 000 µm, respectively), in the
number of layers of the scales (maximum 2 vs. several,
respectively) and in its zoogeographic distribution (Pacic
coast of North America vs. Caribbean Sea, respectively).
According to Théel (1886) and Deichmann (1930, 1954)
small specimens of Y. asperrima (length of body, oral
and anal cones not included, <15 mm) have many single-
layered scales. Ypsilocucumis californiae sp. nov. presents
the same characteristics. The rods of the tentacles are
identical to those of Ypsilothuria bitentaculata attenuata
E. Perrier, 1886 (see Massin 1996: Fig. 2A).
Etymology. The species is a noun in the genitive, referring
to the geographic area (Gulf of “California”) of collection.
Distribution. Central and southern Gulf of California,
Mexico (Fig. 7).
Ypsilothuria bitentaculata (Ludwig, 1893)
Fig. 7
Sphaeroturia bitentaculata Ludwig, 1893: 184; 1894: 141,
pl. XII, gs. 16-17, pl. XIV, gs. 5-14; H.L. Clark, 1913:
229; Ohshima, 1915: 266; Ludwig and Heding, 1935: 76,
textgs. 55-57 (list of citations and synonymy); Parker,
1964: 165; Hansen, 1975: 216; Luke, 1982: 56.
Ypsilothuria bitentaculata; R. Perrier, 1902: 517; Koehler
and Vaney, 1905: 87; Panning, 1949: 455; Madsen, 1955:
167; Caso, 1961: 371; Thandar, 1984: 226, Fig. 39a-k
(list of citations and synonymy); Maluf, 1988: 95, 156;
Nybakken et al. 1998: 1759, 1778; Maluf, 1991: 358;
Lane et al., 2000: 491; Maluf and Brusca, 2005: 342; Tilot,
2006: 59; Sastry, 2007: 254.
Material examined. One specimen (L = 12 mm), TALUD
XII, St. 5 (EMU-8615).
Ypsilothuria bitentaculata is considered a
cosmopolitan species, collected in the East and West
Pacic Ocean, Indian Ocean, North Atlantic Ocean and
Mediterranean Sea. Two varieties have been recognized
by Heding (1942): Y. b. attenuata E. Perrier, 1886 and
Y. b. virginiensis Heding, 1942. The latter is only known
from the North Atlantic Ocean, whereas the former is
cosmopolitan. Due to the complexity in separating the
varieties or subspecies of Y. bitentaculata and the lack
of reliable information related to their distribution, we
have considered the Mexican material as belonging
to Sphaerothuria bitentaculata sensu Ludwig (1893,
Distribution. Records in Mexico. “Albatross” St. 3424
(21o15’N, 106o23’W), 1 237 m (676 fm) (Ludwig,
1894). “Albatross” St. 5675, SW of San Cristobal Bay
(27°07’08”N 114°33’10”W), Baja California, 520 m
(284 fm) (Parker, 1963). Off Salina Cruz (14o28’30”N,
93o09’30”W), 3 539-3 610 m (Parker, 1964; Luke, 1982).
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Revista Mexicana de Biodiversidad 82: 413-443, 2011
Off Tres Marias Islands (probably “Albatross” St. 3424)
(Maluf and Brusca, 2005). Syntypes were collected at
different stations of the “Albatross”, roughly between
00o54’N and 21o15’N (Ludwig, 1894). From central
California (Nybakken et al. 1998), where it is the most
abundant species, San Cristobal Bay, Baja California,
Mexico, to San Francisco Cape, Equator; Indo-West
Pacic (Maluf and Brusca, 2005). Widely distributed
along the Pacic coast of Mexico (Fig. 7).
Order Elasipodida Théel, 1882
Family Laetmogonidae Ekman, 1926
Genus Pannychia Théel,1882
Pannychia moseleyi Théel, 1882
Fig. 9
Pannychia moseleyi Théel, 1882: 3, 88, pl. XVII, gs. 1-2,
pl. XXXII, gs. 1-13; Hansen, 1975: 72, g. 26 (synonyms
and citations); Maluf, 1988: 101, 161 (synonyms and
citations); Solís-Marín et al., 1997: 256; Nybakken et al.,
1998: 1778; Pawson and Ahearn, 2001: 42; Solís-Marín et
al., 2005: 132; Tilot, 2006: 42, 43, g. 75, 60; Anonymous,
2004: 4, 1 pl.; Pawson, 2009: 398; Solís-Marín et al., 2009:
144, pl. 47. gs. A-H.
Material examined. Two specimens (L = 180 mm,
EMU-8632; L = 182 mm, IG 31487/HOL 1509 RBINS/
HOL/738972), TALUD VIII, St. 3. One specimen (L = 11
mm), TALUD VIII, St. 11 (EMU-8617). One specimen (L =
85 mm), TALUD IX, St. 10 (EMU-8616). Four specimens
(L = 41-89 mm; EMU-8631; L = 79 mm, IG 31487/HOL
1516 RBINS/HOL/739010) and 3 specimens (L = 40 and
80 mm, EMU-8619; L = 61 mm, IG 31487/HOL 1510
RBINS/HOL/738978), TALUD XII, St. 25. One specimen
(L = 104 mm), TALUD XII, St. 29 (EMU-8618).
The 12 specimens examined are 40-182 mm long and
6-34 mm across. The body wall contains a few characteristic
wheels, 120-235 µm across, with 11-15 spokes.
As noticed by Hansen (1975), the number and size
of wheels are highly variable and not related to the size
of the specimens. The observed wheels are similar to
the ones described and gured by Hansen (1975) from a
Challenger’s specimen from the Gulf of Panama.
Distribution. Records in Mexico. “Albatross” St. 3431
(23o59’N, 108o40’W), 1 748 m (955 fm); St. 3432
(24o22’30”N, 109o03’20”W), 2 600 m (1 421 fm); St. 3436
(27o34’N, 110o53’40”W), 1 656 m (905 fm); 2.9-3.9oC
(Ludwig, 1894). “Albatross” St. 5676, off San Juanico
(25o31’15”N, 113o29’30”), 1 165 m (647 fm); St. 5685,
SW of Ballenas Bay (25o42’45”N, 113o38’30”), 1 180m
(645 fm); 3.8oC (39oF) (H.L. Clark, 1913). “Albatross” St.
3418 (16o33’N, 99o52’30”W), 1 208 m (660 fm); St. 3425
(21o19’N, 106o24’W) 1 245 m (680 fm); St. 3435 (26o48’N,
110o45’20”W) 1 572 m (859 fm); St. 3436 (27o34’N,
110o53’40”W), 1 656 m (905 fm); 2.9-3.2oC (Ludwig, 1894;
as Laetmophasma fecundum) (Fig. 9). The record of Luke
(1982: 58), at 32°24’42”N-117°27’45”W, is at the southern
limit of the Mexican-USA border (1 204-1 226 m, San
Diego Trough). Solís-Marín et al. (1997) cited this species
off Isla Espíritu Santo (ca 24°30’N, 110°17’W), Baja
California, Mexico, and Solís-Marín et al. (2005) include
2 records for the Gulf of California, both corresponding to
material collected by the “Albatross” (Solís-Marín, pers.
comm.). Two additional records are provided by Solís-
Marín et al. (2009) for the Gulf of California (25o43’50”N,
109o53’59”W) and the California Current area (28o03’N,
1115o10’W) (Fig. 9). Syntypes are from “Challenger” Sts.
164 and 169, 34o8’S, 152oE, and 37o34’S, 179o22’E, E of
New Zealand, in depths of 1 281-1 739 m (700-950 fm);
2.2-4.2oC (Théel, 1882). Widely distributed throughout
the East Pacic, Pannychia moseleyi has been reported
from off Central California (Nybakken et al. 1998) and
San Diego (Luke, 1982) to ca 6oS, off northern Peru, with
many intermediate localities (Maluf, 1988: 101). It also
occurs off Hawaii and in the SW Pacic, and features a
very wide Indo-Pacic distribution (Hansen, 1975; Maluf,
1988; Thandar, 2008). It ranges from 212 to 2 599 m depth,
and the TALUD material was collected within that depth
range (920-1 879 m). Information based on deep-water
trawls and photographs taken in the Gulf of California
and off the coast of California (Solís-Marín et al., 1997;
Anonymous, 2004; Nybakken, 2010) and off California-
Oregon-Washington (Keller et al., 2007) indicates that P.
moseleyi is an abundant species in these areas. According
to Tilot (2006: g. 75) this species is present (photographic
evidence) in the Clipperton-Clarion fractures zone.
Genus Laetmogone Théel, 1879
Laetmogone scotoeides (H.L. Clark, 1913)
Figs 9, 10, A-E
Laetmenoceus scotoeides H.L. Clark, 1913:231.
Laetmogone scotoeides Hansen, 1975: 61, Fig. 23; Maluf,
1988: 100, 160; Thandar, 1998: 87.
Material examined. One specimen (L = 90), TALUD XII,
St 9 (EMU-820). One specimen (L = 150 mm), TALUD
XII, St. 10 (EMU-8634).
Additional material examined. One specimen (L= 86 mm)
669.indd 11 14/06/2011 03:42:45 p.m.
424 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
of Laetmogone wyvillethomsoni, Cruise MV69-VI-9-W,
Patton Escarpment, off Baja California, Mexico (31o11’N,
119o36’W), 18/December/1969, 3600-3676 m depth
(SIO-E 8) (coll. C. Hubbs, S. Luke).
The 90 mm long specimen is in poor condition and
does not allow for an anatomical description. The 150 mm
long specimen is dark violet. On each side of the ventral
sole, a single row of approximately 20 very large tube feet.
Mouth ventral, surrounded by 15 tentacles; anus terminal.
In the body wall numerous wheels. Small wheels 55-130
µm in diameter with 13 spokes and 4 central rays. Large
wheels 120-280 µm in diameter with 8-12 spokes and 5
central rays (g 10 A). No clear-cut distinction between
large and small wheels. Tube feet with spiny rods (190-420
µm long) (Fig. 10D), wheels, and an end-plate. Wheels
similar to those of the body wall, small wheels 30-100
µm in diameter (Fig. 10B), large wheels 120-290 µm in
diameter (Fig. 10C). The end-plate is composed of several
spiny, irregularly ramied rods (g 10E).
According to Hansen (1975), the number of tentacles
(15), the size of the large wheels (up to 300 µm) with 5
central rays are typical of Laetmogone scotoeides. The size
of the large wheels is particularly striking: 50% to 100%
larger than in any other Laetmogone species. (Hansen,
1975; Thandar, 1998). The specimen in poor condition
(L = 150 mm) was rst identied as a L. wyvillethomsoni.
However, examination of the wheel diameter of a
specimen of L. wyvillethomsoni collected off northern Baja
California and obtained on loan (see additional material
examined) shows that this poorly preserved specimen also
belongs to L. scotoeides.
Distribution. Record in Mexico. Type locality, “Albatross”
St. 5685, SW of Ballenas Bay (25o42’45”N, 113o38’30”W),
west coast of Baja California, 1 180 m (645 fm) (H.L. Clark,
1913). Now from off Petatlán, Guerrero (17°11’18”N,
101°28’30W), SW Mexico (Fig. 9). According to Maluf
(1988) L. scotoeides had been reported only from the type
locality. The record for the Bay of La Paz (east coast of
the southern Baja California Peninsula), Mexico, in Solís-
Marín et al. (1997: 255) is an error (Solís-Marín, pers.
comm.). It is not cited by Solís-Marín et al. (2005) and
Honey-Escandón et al. (2008) for Pacic Mexico and we
therefore conclude that no additional record is available for
this species. Present records off SW Mexico occur in a depth
range of 1 180-1 420 m, very similar to the type locality
(1 173 m) and are new for the Pacic coast of Mexico. It
conrms the restricted distribution of L. scotoeides which
seems to be endemic of Pacic Mexico. This is in contrast
with the other Laetmogone species which generally have a
wide distribution (Hansen, 1975).
Family Psychropotidae Théel, 1882
Genus Benthodytes Théel, 1882
Benthodytes cf. sanguinolenta Théel, 1882
Fig. 9
Benthodytes sanguinolenta Théel, 1882: 3-4, 104, pl. XXIII,
pl. XL, gs. 4-5, pl. XLII, g. 6; Ludwig, 1894: 53, pl. I, gs.
1-8; H.L. Clark, 1913: 233; 1920: 142; 1923: 162; Hansen,
1975: 94, pls. III-VI, pl. IX, gs. 6-7, pl. XII, gs. 4-5, g.
116 (list of citations); Parker, 1964: 165; Luke, 1982: 58;
Maluf, 1988: 101, 161; Nybakken et al. 1998: 1778; Maluf,
1991: 360; Honey-Escandón et al., 2008: 58.
Material examined. One specimen (L = 120 mm), Talud
XII, St 15 (EMU-8627).
Specimen attened, with mouth ventral and anus
dorsal. Skin slightly damaged. Unpaired dorsal appendage
absent. Circum-oral papillae present. Mid-ventral tube feet
present, well developed. Color in alcohol grey-white, with
patches of purple. Tentacles purple. No ossicle in body
wall and tube feet.
According to Hansen (1975) the general aspect of
the specimen examined herein ts well with the genus
Benthodytes. Because of the absence of ossicles and the
size and position of the ventral tube feet (see Ludwig, 1894:
pl. I, g. 1), the specimen examined is most probably B.
sanguinolenta but some doubt remains, and it was labeled
B. cf. sanguinolenta.
Distribution. Records in Mexico. “Albatross” St. 3415
(14o46’N, 98o40’W), 3 383 m (1 879 fm); 2.2oC (Ludwig,
1894). Off San Tomas Point, SW of Magdalena Bay, and
off Rosario Bay; 1 969-3 221 m (1 076-1 760 fm); 2.83-
3.30oC (37.1-38.1oF) (H.L. Clark, 1913). “Albatross”
station 4732 (16o32’30”N, 119o59’W); 3 682 m (2 012 fm);
1.5oC (34.8oF) (H.L. Clark, 1920). Undetermined stations
of the “Albatross”, off Baja California, in depths exceeding
1 000 fm (H.L. Clark, 1923). W of Punta Banda (31o18’N,
117o36’W), Patton Escarpment (30o52’N, 116o53’W), and
basin off Magdalena Bay (23o59’30”N, 113o11’54”W),
Baja California, Mexico; 1 975-3 518 m depth (Parker,
1964; Luke, 1982). Off western Baja California
(29o40’12”N, 117o06’36”W); 2 708-2 763 m (Parker,
1964). Environmental data of the material cited by Parker
(1964): 2.0-2.5oC and 1.3-2.8 ml O2/l. Honey-Escandón et
669.indd 12 14/06/2011 03:42:45 p.m.
Revista Mexicana de Biodiversidad 82: 413-443, 2011
al. (2008) reported this species for the California Current
area, in 2 localities off Baja California (W of San José
Point, 31o23’45”N, 118o31’30”W; SW of San Carlos Point,
29o29’N, 116o18’W) (Solís-Marín, pers. comm.) (Fig. 9).
Syntypes are from off Chile (34o7S, 73o56’W, and 38o7’S,
94o4’W), in depths of 2 745-4 072 m (1 500-2 225 fm);
1.3-1.4oC (Théel, 1882). Throughout almost the entire
Indo-Pacic, from 768 to 7 250 m depth, mostly in depths
>2 000 m (Hansen, 1975). Central California and the
Channel Islands, California, USA, to Chile (Maluf, 1988;
Nybakken et al., 1998).
Unidentied Elasipodida
Material examined. One specimen (L = 95 mm), TALUD
VIII, St. 22 (EMU-8626). One specimen (L = 115 mm),
TALUD XII, St. 8. (EMU-8623). One specimen (L = 152
mm), TALUD XIII, St. B (EMU-8628).
The 3 specimens examined look like Elasipodida
because of their general shape and color pattern (dark
violet). The body wall of each specimen, however, is
damaged making it very difcult to ascertain the number
and position of the tube feet and the presence/absence of a
brim. Moreover, ossicles are completely lacking.
Order Aspidochirotida Grube, 1840
Family Synallactidae Ludwig, 1894
Genus Synallactes Ludwig, 1893
Synallactes alexandri Ludwig, 1893
Figs. 11A-H, 12
Synallactes alexandri Ludwig, 1893: 178; Hansen, 1975:
215; Maluf, 1988: 99, 160; Maluf, 1991: 360; Solís-Marín
2003: 249, gs. 194-200.
Scotodeima alexandri; Ludwig, 1894: 21, pl. IX, gs.
Bathyplotes hancocki; Domantay, 1953:136 (nomen
nudum); 1961: 333 (nomen nudum).
Bathyplotes macullochae; Domantay, 1953: 136 (nomen
nudum); 1961: 333 (nomen nudum).
Bathyplotes hancocki Domantay, 1961: 334.
Bathyplotes maccullochae Domantay, 1961: 335.
Material examined. Six specimens (L = 46-97 mm,
EMU-8633; L = 81 mm, IG 31487/HOL 1515 RBINS/
HOL/739007), TALUD VIII, St. 11.
Six specimens examined, 46-97 mm long and 4-12 mm
across. Disposition of mouth, anus, ventral tube feet and
dorsal papillae as described by Ludwig (1894), with the
rows of dorsal papillae less visible than in the type material.
18-20 small peltate tentacles. No ossicles in the ventral body
wall; a few, gathered in heaps, in the dorsal body wall. In
the dorsal body wall and dorsal papillae only cross-shaped
ossicles with 3-4 arms and 1 central pillar (Fig. 11A-B);
each arm forked or perforated at the extremity. Cross-shaped
ossicles 80-100 µm across in the body wall (Fig. 11A), and
100-115 µm across in the dorsal papillae (Fig. 11B). In the
tube feet very numerous, spiny, slightly curved rods, 300-
700 µm long (Fig. 11C), a 1 piece end-plate, 550-650 µm
across, and cross-shaped ossicles similar to those in the
body wall (Fig. 11D), some transformed in table with 4-5
large perforations (Fig. 11E). Central pillar of cross-shaped
ossicles ending in a few spines, its height never exceeding
cross diameter. Tentacles with numerous spiny rods, straight
to strongly curved, 40-700 µm long (Fig. 11F-H), longer
rods (Fig. 11H) located at the base of the tentacles.
The specimens examined are much smaller than in the
types series (46-97 mm vs. 145-175 mm) but the ossicles
are very similar. According to Maluf (1988) 4 species
of Synallactes have been reported in the Central eastern
Pacic, all from water deeper than 350 m, between SW
Mexico and northern Peru. The only previous record of
the genus for Pacic Mexico is for Synallactes ishikawai
f. ind. (= S. sagamiensis Augustin, 1908), reported from
the Gulf of Tehuantepec, SW Mexico, by Parker (1964).
Whether this identication (presumably by E. Deichmann)
was correct or not is impossible to determine. Haney
(2004) reports S. alexandri from off California, but a
close examination of the illustrations included in this
contribution indicates that her material most probably
belongs to a new species recently described: Synallactes
virgulosolida Massin and Hendrickx, 2010. Nybakken et
al. (1998) reported S. aenigma from central California,
with no illustrations, but this species is clearly distinct
from S. alexandri.
Solís-Marín (2003) re-examined material and
descriptions of Synallactes worldwide and retained 22 valid
species. He synonymized both Bathyplotes hancocki and B.
maccullochae with Synallactes alexandri, thus extending
the distribution of the latter to southern California. The
record of B. hancocki for the Gulf of California, cited
by Domantay (1961), however, is in error. All specimens
examined by this author (9 specimens from 5 stations)
and presently in the echinoderms collection at the Los
669.indd 13 14/06/2011 03:42:45 p.m.
426 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
Angeles County Museum of Natural History (except the
holotype from off San Clemente Island, California, which
is apparently missing), were collected off Santa Catalina
Islands, California (G. Hendler, pers. comm.). The material
examined herein, however, extends the distribution of S.
alexandri to the Gulf of California.
Distribution. The species is new to Mexico. The syntypes
are from “Albatross” Sts. 3354, off Mariato Point, Panama
(07o45’N, 80o50’W), and 3406, off the Galapagos Islands
(00°16’S, 90°21’30”W) (Ludwig, 1894). Off Santa
Barbara, California, in depths of 278-550 m (152-300 fm)
(Domantay, 1961, as B. hancocki and B. maccullochae;
Solís-Marín, 2003). In depths of 585-1 018 m (Hansen,
1975; Maluf, 1988; Maluf, 1991). Present record
extends the distribution range of S. alexandri within the
southwestern Gulf of California (Fig. 12) and lls the
distribution gap between Panama and California.
Synallactes virgulasolida Massin and Hendrickx, 2010
Synallactes virgulasolida Massin and Hendrickx, 2010:
600, gs. 1-3.
Material examined. Three specimens (L = 50-85 mm),
TALUD VIII, St. 16 (EMU-8608; IG.31487-HOL 1506;
ICML-UNAM 5.171.0).
This species has been recently described in a separate
paper (Massin and Hendrickx, 2010).
Distribution. Known only from the SW Gulf of California
(Fig. 12), Mexico, at 1 030 m depth.
Order Molpadiida Haeckel, 1896
Family Molpadiidae J. Müller, 1850
Genus Molpadia Risso, 1826
Molpadia intermedia (Ludwig, 1894)
Figs 13, 14
Trochostoma intermedium Ludwig, 1893: 185 (nomen
Trochostoma intermedium Ludwig, 1894: 161, pl. XVI,
gs. 7-21.
Molpadia intermedia; H.L. Clark, 1907: 162, pl. 12, gs.
5-15; 1913: 228; Deichmann, 1937: 174; Caso, 1961: 375;
Maluf, 1988: 105, 163; Nybakken et al. 1998: 1778; Maluf
and Brusca, 2005: 343; Solís-Marín et al., 2005: 132; Tilot,
2006: 62; Honey-Escandón et al., 2008: 58; Solís-Marín et
al., 2009: 148, pl. 49.
Haplodactyla intermedia; Heding, 1931: 280.
Material examined. One specimen (L = 92 mm), TALUD
VI, St. 25 (EMU-8621).
Body very similar to M. musculus, with a short tail
(± 10% of body length). No ossicles in body wall but
numerous phosphatic deposits. Ossicles of the tail tables
only, no fusiform rods. Most of the tables are very irregular
and broken. The disc is 130-210 µm across, perforated by
3-5 large holes (Fig. 13). Spire made of 2-3 pillars, 100-
120 µm high, fused at the top.
Molpadia intermedia is said to have a long tail (20-
25% of total body length; Ludwig, 1894; H.L. Clark, 1907;
Solís-Marín et al., 2009). The tail of the observed specimen
is very short (10% of body length), making it close to M.
musculus. The absence of fusiform rods and the presence
of irregular tables in the tail, however, are characteristic of
M. intermedia (see Ludwig, 1894: pl. XVI, gs. 16-19).
Molpadia intermedia is the most common species of the
genus along the Central eastern Pacic coast (H.L. Clark,
1907; Deichmann, 1937) and it is therefore surprising that
only 1 specimen was collected during the TALUD survey.
Distribution. Records in Mexico. “Albatross” Sts. 2838
(28o12’N, 115o09’W), 79 m depth, and 3431 (23o59’N,
108o40’W), 1 791 m depth (H.L. Clark, 1907); “Albatross”
Sts. 5676 (off San Juanico; 25o31’15”N, 113o29’30”W),
5683 (off Cabo San Lucas; 22o46’45”N, 109o50’15”W),
5688 (off Cedros Island; 27°38’5”N 115°17’40”W), 5689
and 5690 (off Ballenas and Rosario Bay; 29°23’00N
116°14’W and 29°29’N 116°18’W), Baja California,
in depths of 961-2 015 m (525-1 101 fm); 3.30-4.38oC
(38.1-39.9oF) (H.L. Clark, 1913). East of Cedros Island
(St 126-DA, Templeton Crocker), Baja California
(Deichmann, 1937). Solís-Marín et al. (2005) reported 2
records for the Gulf of California and Honey-Escandón et
al. (2008) 2 records for the California Current area. These
records are probably the same as those cited by Solís-
Marín et al. (2009; 5 records in total), and were included in
this compilation (28o12’00” N, 115o09’09” W; 24o15’18”
N, 108o24’06” W; 24o53’12” N, 108o59’24” W; 24o56’24”
N, 109o05’36” W; 24o51’41” N, 108o57’52” W) (Fig. 14).
From Alaska to Cabo Mala, Panama; West Pacic (Maluf
and Brusca, 2005); in depths of 55-2 014 m (Maluf, 1988).
Molpadia musculus Risso, 1826
Fig. 14
Molpadia musculus Risso, 1826: 293; H.L. Clark, 1907:
35, 158, 165, pl. XI; 1913: 228; 1923: 161; Caso 1961:
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Revista Mexicana de Biodiversidad 82: 413-443, 2011
375; Pawson, 1977: 100, gs. 3a-d, map 1 (synonymy and
list of citations); Luke, 1982: 59; Maluf, 1988: 105, 163;
Nybakken et al., 1998: 1778; Maluf and Brusca, 2005:
343; Solís et al., 2005: 132; Honey-Escandón et al., 2008:
58; Solís-Marín et al., 2009: 150, pl. 50.
Molpadia musculus forma violacea; Parker 1964: 165.
Molpadia musculus forma spinosa; Parker, 1964: 165.
Material examined. Four specimens (L = 17 and 47 mm,
EMU-4203; L = 52 and 58 mm, IG 31487/HOL 1513
RBINS/HOL/738993), TALUD III, St. 14B. One specimen
(L = 98 mm), TALUD IX, St. 16 (EMU-8622).
No ossicles in the body wall, but only numerous
phosphatic deposits. In the tail fusiform rods only. The
smaller the specimen, the more numerous the ossicles.
Rods in the tail show the same diversity as the one
reported and illustrated by Pawson (1977: g. 2a-f) for
M. musculus from off southern Chile and South Shetland
Island. Molpadia musculus is considered a cosmopolitan
and highly variable species, described repeatedly under
different names or varieties (see Maluf, 1988). A detailed
review of species of Molpadia occurring in the southern
oceans is available in Pawson (1977), including the
redescription of M. musculus. Records for Molpadia
musculus forma musculus and for Molpadia musculus
forma spinosa by Parker (1964: 165) are for southern
California and northern Guatemala, respectively, in both
cases close to the border limit with Mexico.
Distribution. Records in Mexico. “Albatross” Sts. 3418,
3429 (16o33’N, 99o52’30”W; 22o30’30”N, 107o01’W),
1 188-1 654 m (H.L. Clark, 1907). SW of Magdalena Bay
(“Albatross” St. 5684; 23o23’30”N, 112o30’W) and off
Santo Tomas Point (“Albatross” St. 5692, 31o23’45”N,
118o31’30”W), 1 969-3 170 m (1 076-1 760 fm); 2.83
(37.1oF) (H.L.Clark, 1913, 1923). “Albatross” St. 3434
(25o29’30”N, 109o48’W), 2 906 m (1 588 fm) (Ludwig,
1894; as Trochostoma violaceum). “Albatross” St. 3436
(27o34’N, 110o53’40”W), 1 656 m (905 fm); 2.9° C
(Ludwig, 1894; as Ankyroderma spinosum). Off Salina
Cruz (15o38’N, 95o18’30”W), Gulf of Tehuantepec, and
NW of San Juanico Island (22o11’12”N, 107o46’06”W),
Gulf of California, in depths of 1 006-3 001 m; 2.0-8.0oC
and 0.1-2.0 ml O2/l (Parker, 1964). Solís-Marín et al.
(2005) reported 1 record for the Gulf of California, off
Mazatlán (22o30’30”N, 107o01’W), and Honey-Escandón
et al. (2008) 1 record over the Lusitania Bank (23o23’30”N,
112o00’30”W), off Baja California Sur, in the California
Current area (Solís-Marín pers. comm.). Solís-Marín et
al. (2009) reported an additional lot in the holdings of
the Smithsonian Institution, from off Mazatlán (23o12’N,
106o25’W) (Fig. 14). The type locality is the Gulf of Nice,
Mediterranean Sea. Widely distributed in the East Pacic,
it is known from Monterey, California, to southern Chile.
A cosmopolitan species also recorded worldwide except
north of the Arctic Circle (Pawson, 1977; Maluf, 1988;
Borrero-Peréz et al., 2005).
The depth range provided by Maluf (1988) for this
species (4 to 5 205 m depth) is astonishing. A search in the
literature indicates that the sample presumably collected at
only 4 m depth most certainly corresponds to a record by
Parker (1964) at a locality situated in the Bay of La Paz (St. 4,
depth 4-7 m, water temperature 22oC), sampled in 1959. The
extremely high water temperature and the subtidal character
of the sampling site indicate that this record is uncertain.
Perhaps this sample had been confused with the preceding
sample (St. 3, visited 2 days earlier, same area, 2 710 m depth)
and erroneously labeled. Depth records for Pacic Mexico are
from 830 to 3 170 m. According to Pawson (1977), however,
M. musculus has been collected from 35 to 5 205 m, still a
remarkably wide bathymetric range.
It should be emphasized that in most surveys either
survey methods, sampling effort, or depth range are distinct,
thus rendering formal comparison difcult. Depth range,
for example, is very important for the distribution of the
Elasipodida, particularly for the families Psychropotidae
and Elpidiidae (Hansen, 1975; Gebruk, 1990). Most of the
species belonging to those families live deeper than 2 000
m. Consequently, Elasipodida collected at a depth range
of 350-2 200 m (present study) or 400-2 900 m (Gage et
al., 1985) represent 31% and 27 % of the species diversity,
respectively. If the sampled depth is 2 000-4 000 (Sibuet,
1977) or 350-4 000 (Ludwig, 1894) the Elasipodida species
diversity can reach up to 37% and 45%, respectively.
As a result of the compilation of the data available
in the literature, and including the 13 species collected
during the TALUD cruises, 31 deep-water species (29
identied at species level and 2 at genus level) have at least
1 record from off the Pacic coast of Mexico: 13 in the
California Current area (CC, south of the USA border), 20
in the Gulf of California (GC), and 14 (15 if the doubtful
record of Peniagone intermedia is considered) along
southwestern Mexico (SWM, south of Banderas Bay to
the Guatemala border). Two species have been collected
off Mexican oceanic islands (see Table 2). It should be
noted, however, that the Gulf of California southern limit
669.indd 15 14/06/2011 03:42:45 p.m.
428 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
used in table 2 is a line extending from San Lucas Cape
(ca 22o53’N, 109o58’W) to Corrientes Cape (20o24’44”N,
105o43’38”W), on the southern edge of Banderas Bay
(see Hendrickx et al., 2005). Consequently, the record
of Molpadia granulata off Mazatlán (ca 23o13’N,
106o27’W), a locality sometimes considered as being
outside the Gulf limits, allows us to include it in the Gulf of
California species list. Parker (1964) included 17 species
living deeper than 350 m in this data base: Peniagone
sp. could correspond to any of the 3 species reported
for deep water in the Central eastern Pacic (see Maluf,
1988). Psychropotes dubiosa and P. raripes are now
considered junior synonyms of P. longicauda, and Parker’
s information is therefore considered as valid record for
the later species in the California Current and along SW
Mexico. The record for S. sagamiensis should be taken
with care due to the difculty to properly identify species
in this genus. In this survey we collected 3 species cited
by Parker (i.e., B. sanguinolenta, Y. bitentaculata, and M.
musculus). Seven of the 23 “Mexican” species reported
by Maluf (1988) were collected during this survey (see
Table 2). Peniagone leander was described after Maluf
(1988) had completed her search but has not been found
off Mexico. The contribution by Solís-Marín et al. (1997)
refers to 3 records of deep-water species off SW Baja
California (Laetmogone scotoeides, Pannychia moseleyi
and Paracaudina chilensis) (see Table 2). The record of
L. scotoeides, however, is in error (F. Solís-Marín, pers.
comm.). Within the Gulf of California, Maluf and Brusca
(2005) have reported 57 species of Holothuroidea, most
from shallow water. Eight deep-water (>350 m depth)
species are included in their list, all previously included in
Maluf‘s 1988 list of species (see Table 2).
Compilations by Solís-Marín et al. (2005) and Honey-
Escandón et al. (2008) include, when combined, 55 species
of Holothuroidea for Pacic Mexico. Unfortunately, depth
ranges were not indicated in these lists, but review of
bathymetric records available in literature indicates that 6
of these species are from water deeper than 350 m (see
Table 2), all also previously reported by Maluf (1988)
for the Gulf of California. In their synopsis of the Gulf
of California Holothuroidea, Solís-Marín et al. (2009)
include 11 species with bathymetric range reaching
depths greater than 350 m. We believe, however, that the
following 5 records are either doubtful [i.e., Cucumaria
crax Deichmann, 1941; Pseudocnus californicus
(Semper, 1868); Holothuria leucospilota (Brands, 1835);
Parastichopus californicus (Stimpson, 1857)] or need to
be veried (i.e., Chiridota aponocrita H.L. Clark, 1920,
which depth information was taken from another species)
(see Appendix I). Holothuria leucospilota, for example, a
very common species throughout the Indo-Pacic Ocean,
is known as an intertidal species with a maximal depth
record of 10 m (Samyn and Massin, 2003). The 695 m
cited by Solís-Marín et al. (2009) seems unlikely. We have
considered as valid only the record for Abyssocucumis
albatrossi (cited as Stereocucumis abyssorum; however,
presence of spiny arms on the ossicles indicates the material
belongs to A. albatrossi), Pannychia moseleyi, Molpadia
intermedia, and M. musculus, all 4 species collected during
this survey, and for Scotoplanes clarki and Heldingia
californica (see Table 2), not collected during the TALUD
cruises. Finally, a new deep water species of Synallactes
was recently described from the Gulf of California (Massin
and Hendrickx, 2010) and has also been included in the list
of Mexican species (Table 2).
The capture of 1 large specimen of Abyssocucumis
albatrossi in the central Gulf of California conrms the
presence of a second species of the genus in Mexican
waters and is the rst conrmed record for the Gulf. The
rediscovery of Psolidium gracile, reported only once
from off California since its original description, based on
material from off Panama, allows us to report it from off
SW Mexico and within the Gulf of California. Laetmogone
scotoeides was previously known only from the type
locality, SE of Ballenas Bay, in the California Current
area, and is now cited for SW Mexico. The large series
of specimens of Pannychia moseleyi collected during this
survey conrms the abundance and high occurrence of this
species along western Mexico. Synallactes alexandri is
recorded for the rst time in Mexican waters.
For most species studied herein, depth and epibenthic
water temperature are similar to the data available in the
literature (e.g., Ludwig, 1894; H.L. Clark, 1913, 1920;
Parker, 1964). Information on epibenthic dissolved oxygen
concentration at sampling sites is rarely available for deep-
water species. In this study, focused on the fauna living
at the edge and below the minimum oxygen zone off the
Pacic coast of Mexico, we were able to measure close-to-
bottom oxygen concentration associated with the capture of
Holothuroidea (Table 1). These data indicate adaptation to
mild hypoxic conditions for Laetmogone scotoeides (0.68-
1.01 ml O2/l) and to more severe hypoxic condition for
Psolus squamatus (0.26-0.36 ml O2/l), Molpadia musculus
(0.15-0.40 ml O2/l), and both species of Synallactes (0.20
ml O2/l). It is to be noted, however, that Molpadia musculus
has been reported by Parker (1964) in dissolved oxygen
concentration of 1.80-2.00 ml O2/l in localities just north
and south of Mexico. Mitsukuriella unusordo sp. nov. was
also collected in low oxygen concentration (0.32 ml O2/l)
but additional information is needed to conrm its afnity
669.indd 16 14/06/2011 03:42:45 p.m.
Revista Mexicana de Biodiversidad 82: 413-443, 2011
to a hypoxic environment. Some species were collected
in a wide range of dissolved oxygen values (Psolidium
gracile, <0.05-1.05 ml O2/l; Yspilocucumis californiae
sp. nov., 0.20-1.40 ml O2/l; Pannychia moseleyi, 0.11-
1.38 ml O2/l) corresponding to a wide bathymetric range,
thus indicating the possibility for these species to extend
their vertical distribution from the edge of the anoxic zone
into deeper water. Moereover, the number of P. gracile
collected in 2 stations (i.e., 6 specimens in St. 17, TALUD
IX, and 17 in St. 1, TALUD XI) indicates that this species
can be abundant in this habitat. Benthodytes sanguinolenta,
a species occurring mostly below 2 000 m depth, was
collected at a similar depth, in relatively well oxygenated
waters (i.e., 1.61 ml O2/l) during this survey, and its known
range of tolerance to oxygen concentration is 1.30-2.80 ml
O2/l (Parker, 1964). The unique locality where Ypsilothuria
bitentaculata and Abyssocucumis albatrossi were found
features similar environmental conditions (1 925-1 977 m,
1.43 mlO2/l for the former; 2 056-2 195 m, 1.68 ml O2/l
for the latter). Off western Mexico, oxygen concentrations
increase to values superior to 1.50 ml O2/l below 2 000
m, and this might indicate that B. sanguinolenta, Y.
bitentaculata, and Abyssocucumis albatrossi are not able to
tolerate the low oxygen concentrations found in shallower
water (i.e., between 700 and 1 300 m where oxygen values
range from almost zero to ca 1.00 ml O2/l) (Table 1) (see
Hendrickx, 2001; Hendrickx and Serrano, 2010). Off
Oregon, these species were collected below 2 000 m, also
much deeper that the oxygen minimum zone occurring in
that area (Carney and Carey, 1976). The possibility for
several species of deep-water holothurians to survive in
severe or extreme hypoxic conditions represents a decisive
advantage with regards to competition for food or potential
predators. On the other hand, as noted by Alton (1972) and
Carney and Carey (1976) off Oregon , the presence of a
minimum oxygen zone might serve as a physiological
barrier to both upward and downward extension of range
for many species, a similar pattern to the one described
for shrimp (Dendrobranchiata and Caridea) along the west
coast of Mexico (Hendrickx and Serrano, 2010). Still, it
is interesting to note that the 2 stations where more than
1 species of Holothuroidea were found during this survey
experienced severe hypoxia (see Table 1).
Distribution and biodiversity
Comparison with neighbouring areas. There are few
recent data available to compare the material collected
during the TALUD survey with the rest of the Central
eastern Pacic. Bluhm (1994) reported on holothurians
collected in manganese module sites off northern Peru (ca
90°W) and in the central Pacic, but none of his species
(most identied to genus) matches with the TALUD
material. Bluhm and Gebruk (1999) reported 20 holothurian
species observed by means of a remote control camera in
the Peru Basin, roughly between 3 800 and 4 200 m depth.
Many species were identied to genus level, and only 1
species, Benthodytes sanguinolenta, coincides with our
study. Pawson and Ahearn (2001) studied a small series
of holothurians collected off the Galapagos Islands and
reported the presence of 7 species, 3 probably undescribed
and only 1 (Pannychia moseleyi) common with our study.
On the other hand, there has been a large series of
surveys of the megafauna on the continental slope and
abyssal plains off the west coast of the USA, using both
conventional trawls and camera sled. A comparative analysis
of the composition of the Holothuroidea fauna collected in
this area is provided by Nybakken et al. (1998). According
to this review, the highest species richness was reported
by Carney and Carey (1976) off Oregon, who reported 28
species below the continental shelf zone (>300 m to the 4
000 m depth range). Six of these species were found during
the TALUD project, including 2 species (i.e., A. albatrossi
and B. sanguinolenta) reported by these authors only in
water much deeper (>2 700 m) than in our study. Carney
and Carey (1982) collected 22 species of Holothuroidea
between 2 162 and 3 961 m on Cascadia Basin and
Tufts Abyssal Plain, off Oregon, including 5 species (A.
albatrossi, B. sanguinolenta, P. moseleyi, M. musculus, and
Y. bitentaculata) found during the TALUD project in much
shallower waters. Of the 13 species collected by Nybakken
et al. (1998) off central California, 6 were found during the
TALUD cruises. Quite remarkably, these 6 species are the
same as those common between the “1976” Oregon and the
TALUD surveys. A more recent survey along the coast of
California, Oregon and Washington, detected the presence
of 8 deep-water species ranging to 721-1 285 m depth,
including Molpadia intermedia, Pannychia moseleyi, and
Psolus squamatus, all 3 considered among the moderately-
frequent species (Keller et al., 2007). According to Lambert
(2007), there are 45 species of sea cucumbers reported
for British Columbia, 34 occurring below 200 m. His list
includes Pannychia moseleyi, Ypsilothuria bitentaculata,
Psolus squamatus, and Molpadia intermedia, but the
former 2 are only found above 200 m off British Columbia.
Including the material collected during the TALUD survey,
the deep-water (>350 m) Holothuroidea fauna occurring
of the Pacic coast of Mexico comprises of 31 species, a
number quite similar to those numbers reported for Oregon
(28 in >300 m) and British Columbia (34 in >200 m), but
apparently higher than those reported for California.
Comparison with remote areas. If we compare
deep-water species richness from the Philippines (Cherbonnier
and Féral, 1981) and the Rockall Trough (U.K.) (Gage et
669.indd 17 14/06/2011 03:42:46 p.m.
430 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
al., 1985) with Pacic Mexico (all 3 with similar sampling
depth: 379-1 125, 400-2 900, and 350-2 200, respectively),
the Philippines and the U.K. areas appear much richer. The
number of species per number of samples is 1.20 for the
Philippines, 0.97 for U.K. and only 0.09 for Mexico. This
difference, of an order of magnitude of 10, could be linked
to the limiting effect of the Pacic Mexico OMZ, which
is not present in the 2 other areas. Rockall Trough and the
Mexican Pacic differ also drastically by their holothurian
species composition. Of the 34 and 31 species collected
respectively only 2 are shared, i.e. Psychropotes longicauda
and Ypsilothuria bitentaculata.
Even considering the strong limitations of this
study (i.e., reduced number of days at sea and samples,
sampling depth, extension of the study area), the TALUD
exploratory survey initiated in 1989, with major collecting
efforts in 2000-2001 and 2005-2008, represents one of the
most important event for the knowledge of deep-water
Holothuroidea of western Mexico since the “Albatross”
collected material between Guatemala and the southern
Gulf of California, in 1891-92. There have been very few
recent studies of deep-water Holothuroidea (and other
Echinodermata) in the East Pacic. Yet, every single study
has fully demonstrated that even limited sampling effort
on the bathyal seaoor can bring very interesting results,
particularly considering the impact of the Minimum
Oxygen Zone on the composition and distribution of the
deep-water benthic communities in the East Pacic.
The authors thank all scientists, students and crew
members who took an active part into the TALUD cruises
aboard the R/V “El Puma”. One of us (MEH) is grateful
to the Royal Belgian Institute of Natural Sciences for
its hospitality during his sabbatical leave, to Thierry
Backeljau and Claude Massin for their invitation, and to
the DGAPA, PASPA, UNAM, Mexico, for supporting his
sabbatical stay. We thank Harim Cha, SCRIPPS Institution
of Oceanography, La Jolla, for the loan of specimens, José
Salgado Barragán for his technical support during the
study of the material, A. Van Haelen for the photographs
of plate 1. Gordon Hendler provided data related to Allan
Hancock expeditions material, and Mercedes Cordero
made the nal edition of the manuscript. Thanks to our
colleagues and friends Manuel, Sammy, José-Antonio and
David U., and to R. García-Tenorio and J. Ontiveros for
helping with availability of specimens. This project was
partly supported by CONACyT, Mexico (project 31805-
N) and DGAPA (project IN-217306-3), UNAM, Mexico.
Literature cited
Alton, M. S. 1972. Bathymetric distribution of echinoderms off
the Northern Oregon Coast. In The Columbia River Estuary
and Adjacent Ocean Waters, A. T. Pruter and D. L. Alverson
(eds.). University of Washington Press, Seattle p. 475-536.
Anonymous. 2004. SCAMIT Newsletter. 23:1-8.
Belyaev, G. M. 1971. Deep water holothurians of the genus
Elpidia. In Fauna of the Kurile-Kamchatka Trench and its
environment, Vol. 92 (in Russian), V. Grigor’yevich Bogorov
(ed.). Trudy Instituta Okeanologii, P.P. Shirshova. p. 326-367.
Bergen, M. 1980. Holothuroidea. In A taxonomic listing
of common marine invertebrate species from Southern
California, D. Straugham and R. W. Klink (eds.). Technical
Reports of the Allan Hancock Foundation 3:274-277.
Bluhm, H. 1994. Monitoring megabenthic communities in
abyssal manganese nodule sites of the East Pacic Ocean
in association with commercial deep-sea mining. Aquatic
Conservation: Marine and Freshwater Ecosystems 4:187-
Bluhm, H. and A. Gebruk. 1999. Holothuroidea (Echinodermata)
of the Peru Basin – ecological and taxonomic remarks based
on underwater images. Marine Ecology 20:167-195.
Borrero-Peréz, G. H, M. Benavides-Serrato, O. D. Solano
and G. R. Navas. 2005. Holothuroideos (Echinodermata:
Holothuroidea) recolectados en el talud continental superior
del Caribe colombiano. Boletín del Instituto Oceanográco,
Universidad de Oriente, Venezuela 42:65-85.
Carney, R. S. and A. G. Carey, Jr. 1976. Distribution pattern
of holothurians on the Northeastern Pacic (Oregon,
U.S.A.) continental shelf slope, and abyssal plain. Thalassia
Jugoslavica 12:67-74.
Carney, R. S. and A. G. Carey, Jr. 1982. Distribution and diversity
of holothuroids (Echinodermata) on Cascadia Basin and
Tufts Abyssal Plain. Deep-Sea Research 29:597-607.
Caso, M. E. 1961. Estado actual de los conocimientos acerca
de los equinodermos de México. Tesis doctorado, Facultad
de Ciencias, Universidad Nacional Autónoma de México,
México, D.F. 388 p.
Caso, M. E. 1964. Contribución al conocimiento de los
Holoturoideos de México. Descripción de un nuevo subgénero
del género Holothuria. Holothuria (Paraholothuria) y de
una nueva especie Holothuria riojae. Anales del Instituto
de Biología, Universidad Nacional Autónoma de México
Caso, M. E. 1966. Estudios sobre Equinodermos de México.
Contribución al conocimiento de los Holoturoideos de
Zihuatanejo y de la Isla de Ixtapa (primera parte). Anales del
Instituto de Biología, Universidad Nacional Autónoma de
México, Serie Ciencias del Mar y Limnología 36:253-291.
Caso, M. E. 1968. Contribución al estudio de los Holoturoideos
de México. La familia Psolidae. Descripción de una nueva
669.indd 18 14/06/2011 03:42:46 p.m.
Revista Mexicana de Biodiversidad 82: 413-443, 2011
especie del género Psolus, Psolus conchae sp. nov. Anales
del Instituto de Biología, Universidad Nacional Autónoma de
México, Serie Ciencias del Mar y Limnología 39:1-20.
Cherbonnier, G. 1941. Étude anatomique et biogéographique
sur deux Cucumaria abyssaux: C. abyssorum Théel et C.
albatrossi n. sp. Bulletin du Muséum national d’Histoire
naturelle (Paris) ser. 2, 13:93-103.
Cherbonnier, G. 1947. Note sur une holothurie abyssale:
Abyssocucumis ingol (Deichmann, 1927). Bulletin du
Muséum national d’Histoire naturelle (Paris) ser. 2, 19:459-
Cherbonnier, G. and J-P. Féral, 1981. Echinodermes: Holothuries.
In Résultats des Campagnes MUSORSTOM. I. Philippines
(18-28 mars 1976). Tome 1. Collection Mémoires ORSTOM
Clark, H. L. 1907. The Apodous holothurians. A monograph of
the Synaptidae and Molpadiidae. Smithsonian Contributions
to Knowledge 35:1-206.
Clark, H. L. 1913. Echinoderms from Lower California, with
descriptions of new species. Bulletin of the American
Museum of Natural History 32:185-236.
Clark, H. L. 1920. Holothuroidea. Report XXXII on the Scientic
Results of the Expedition of the “Albatross” to the Tropical
Pacic, 1899-1900 and 1904-1905. Memoirs of the Museum
of Comparative Zoology at Harvard College 39:121-154.
Clark, H. L. 1923. Echinoderms from Lower California, with
descriptions of new species: Supplementary report. Bulletin
of the American Museum of Natural History 48:147-163.
Deichmann, E. 1930. The holothurians of the western part of
the Atlantic Ocean. Bulletin of the Museum of Comparative
Zoology Harvard 71:43-226.
Deichmann, E. 1937. Holothurians from the Gulf of California,
the west coast of Lower California and Clarion Island.
Zoologica, N.Y. 22:161-176.
Deichmann, E. 1938a. New holothurians from the western coast
of North America, and some remarks on the genus Caudina.
Proceedings of the New England Zoological Club 17:23-25.
Deichmann, E. 1938b. New records of Paracaudina chilensis (J.
Müller) from the west coast of Central America and Mexico.
Proceedings of the New England Zoological Club 16:103-
Deichmann, E. 1938c. Holothurians from the western coasts
of Lower California and Central America, and from the
Galapagos Islands; Eastern Pacic Expeditions of the New
York Zoological Society. Zoologica, N.Y. 23:361-387.
Deichmann, E. 1941. The Holothuroidea collected by the “Velero”
III during the years 1932 to 1938. Part 1. Dendrochirota.
Allan Hancock Pacic Expeditions 8:61-153.
Deichmann, E. 1954. The holothurians of the Gulf of Mexico.
Bulletin of the United States Fishery Commission 55:381-41.
Domantay, J. S. 1953. A brief summary of the Pacic and Atlantic
Holothuroidea of the Allan Hancock Foundation Collections.
Philippine Journal of Science 82:133-140.
Domantay, J. S. 1961. New forms of Holothuroidea from the
vicinity of Southern California and Mexico in the collection
of the Allan Hancock Foundation, University of Southern
California. Philippine Journal of Science 90:333-346.
Ekman, S. 1923. Über Psolus squamatus und verwandte Arten.
Zugleich ein Beitrag zur Bipolaritätsfrage. Arkiv för
Zoölogie 15:1-59.
Gage, J.D., D.S.M. Billett, M. Jensen and P.A. Tyler. 1985.
Echinoderms of Rockall Trough and adjacent areas.2.
Echinoidea and Holothurioidea. Bulletin of the British
Museum (Natural History), Zoology 48:173-213.
Gebruk, A.V. 1990. Deep-Sea holothurians of the family
Elpidiidae. Akademia Nauka, Moscow, URSS. 160 p. [In
Grassle, J. F. 1989. Species diversity in deep-sea communities.
Trends in Ecology and Evolution 4:12-15.
Haney, L. 2004. Holothuroidea. SCAMIT Newsletter 23:10-30.
Hansen, B. 1975. Systematics and Biology of the Deep-Sea
Holothurians, Part 1: Elasipoda. Galathea Reports 13:1-262.
Hansen, B. 1988. The genus Staurocucumis Ekman and its
possible afnity with Echinocucumis Sars (Holothuroidea,
Dendrochirota). In Echinoderm Biology, R.D. Burke, P.V.
Madlenov, P. Lambert and R.I. Parsley (eds.). Balkema,
Rotterdam p. 301-308.
Heding, S.G. 1931. On the classication of the Molpadids.
Videnskabelige Meddelelser fra Dansk naturhistorisk
Forening i Kǿbenhavn 92:275-284.
Heding, S.G. 1942. Holothurioidea. II. Danish Ingolf-Expedition
Heding, S.G. and A. Panning. 1954. Phyllophoridae. Ein
Bearbeitung der polytentaculaten dendrochiroten
Holothurien des zoologischen Museums in Kopenhagen.
Spolia Zoologica Musei Hauniensis XIII:1-209.
Hendrickx, M. E. 2001. Occurrence of a continental slope
decapod crustacean community along the edge of the
minimum oxygen zone in the southeastern gulf of California,
Mexico. Belgian Journal of Zoology 131:95-109.
Hendrickx, M. E. and D. Serrano. 2010. Impacto de la zona de
mínimo de oxígeno sobre los corredores pesqueros en el
Pacíco mexicano. Interciencia. 35:12-18.
Hendrickx, M. E., R. C. Brusca and L. T. Findley. 2005. A
Distributional Checklist of the Macrofauna of the Gulf
of California, Mexico. Part I. Invertebrates. [Listado y
Distribución de la Macrofauna del Golfo de California,
México, Parte I. Invertebrados]. M. E. Hendrickx, R. C.
Brusca and L. T. Findley (eds.). Arizona-Sonora Desert
Museum. Tucson, AZ, USA. 429 p.
Hessler, R. R. and H. L. Sanders. 1967. Faunal diversity in the
deep sea. Deep-Sea Research 14:65-78.
Honey-Escandón, M., F. A. Solís-Marín and A. Laguarda-
Figueras. 2008. Equinodermos (Echinodermata) del Pacíco
669.indd 19 14/06/2011 03:42:46 p.m.
432 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
Mexicano. Revista de Biología Tropical 56 (Suppl. 3):57-73.
Imaoka, T. 1980. Observations on Psolus squamatus (Koren) from
the Okhotsk Sea (Dendrochirota: Psolidae). Publications of
the Seto Marine Biological Laboratory 25:361-372.
Keller, A. A., V. H. Simon, B. H. Homess, J. R. Wallace, V. J.
Tuttle, E. L. Fruh, K. L. Bosley, D. M. Kamikawa and J. C.
Buchaman. 2007. The 2003 U.S. West Coast bottom trawl
survey of groundsh resources off Washington, Oregon,
and California: Estimates of distribution, abundance, and
length composition. U.S. Department of Commerce, NOAA
Technical Memo. NMFS-NWFSC-86. 130 p.
Koehler, R. and C. Vaney. 1905. II. Les Holothuries Littorales.
In An account of the littoral Holothurioidea collected by
the R.I.M.S.S. “Investigator”. Echinoderma of the Indian
Museum, Part IV, Holothurioidea. Indian Museum, Calcuta
p. 1-123.
Koren, J. 1845. Beskrivelse over Thyone fusus og Cuviera
squamata. Nyt Magazin for Naturvidenskaberne 4:203-225.
Lambert, P. 1986. Northeast Pacic holothurians of the
genus Parastichopus with a description of a new species
Parastichopus leukothele (Echinodermata). Canadian
Journal of Zoology. 64:2266-2272.
Lambert, Ph. 1996. Psolidium bidiscum, a new shallow water,
psolid sea cucumber (Echinodermata: Holothuroidea) from
the northeast Pacic, previously misidentied as Psolidium
bullatum Ohshima. Canadian Journal of Zoology 74:20-31.
Lambert, Ph. 1997. Sea cucumbers of British Columbia,
Southeast Alaska and Puget Sound. University of British
Columbia Press, Vancouver. 166 p.
Lane, D. J. W., L. M. Marsh, D. VandenSpiegel and F. W. E.
Rowe. 2000. Echinoderm fauna of the South China Sea: an
inventory and analysis of distribution patterns. The Rafes
Bulletin of Zoology Supplement 8:459-493.
Levin, L. A. and J. D. Gage. 1998. Relationships between oxygen,
organic matter and the diversity of bathyal macrofauna.
Deep-Sea Research II 45:129-163.
Levin, L. A., R. J. Etter, M. A. Rex, A. J. Gooday, C. R. Smith,
J. Pineda, C. T. Stuart, R. R. Hessler and D. Pawson. 2001.
Environmental inuences on regional deep-sea species
diversity. Annual Review Ecology System 32:51-93.
Ludwig, H. 1893. Vorläuger Beritch über die auf den Tiefsee-
Fahrten des “Albatross” (Fruhling 1891) im östlichen Stillen
Ocean erbeuteten Holothurien. Zoologischer Anzeiger
Ludwig, H. 1894. Reports of an Exploration 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, Lieutenant
Commander Z.L. Tanner, U.S.N., Commanding, XII: The
Holothurioidea. Memoirs of the Museum of Comparative
Zoology at Harvard College 17:1-183.
Ludwig, H. 1900. Arktische und subarktische Holothurien.
Chapter V. In Fauna Arctica, F. Römer and F. Schaudin
(eds.). Gustav Fisher, Jena 1900 1:134-178.
Ludwig, H. and S. G. Heding. 1935. Die Holothurien der
Deutschen Tiefsee-Expedition. 1. Fusslose und dendrochirote
Formen. Wissenschaftliche Ergebnisse der Deutschen
Tiefsee-Expedition auf dem Dampfer Valdivia 1898–1899
Luke, S. R. 1982. Catalog of the Benthic Invertebrate Collections,
Echinodermata. Scripps Institution of Oceanography
Reference Series, No. 82-5. University of California. 71 p.
Lütken, Chr. 1857. Oversigt over Grölands Echinodermata.
Videnskabelige Meddelelser fra Dansk naturhistorisk
Forening i Kǿbenhavn 1857:1-55.
Madsen, F. J. 1955. Holothuroidea. Reports of the Swedish Deep-
Sea Expedition, Part II. Zoology 12:151-173.
Madsen, F. J. and B. Hansen. 1994. Echinodermata Holothurioidea.
Marine invertebrates of Scandinavia, 9. Scandinavian
University Press. Oslo, Norway. 143 p.
Maluf, L. I. 1988. Composition and distribution of the Central
eastern Pacic Echinoderms. Technical Report, Natural
History Museum of Los Angeles County 2:1-242.
Maluf, L. Y. 1991. Echinoderm Fauna of the Galapagos Islands.
Chapter 16. In Galapagos Marine Invertebrates: Taxonomy,
Biogeography and Evolution in Darwin’s Islands, M. J.
James (ed.). Plenum Press, New York p. 345-367.
Maluf, L. I. and R. C. Brusca. 2005. Echinodermata. Chapter
18. In A Distributional Checklist of the Macrofauna of the
Gulf of California, Mexico. Part I. Invertebrates. [Listado
y Distribución de la Macrofauna del Golfo de California,
México, Parte I. Invertebrados], M. E. Hendrickx, R. C.
Brusca and L. T. Findley (eds.). Arizona-Sonora Desert
Museum, Tucson, Az. USA p. 327-343.
Massin, C. 1994. Ossicle variation in Antarctic dendrochirote
holothurians (Echinodermata). Bulletin de l’Institut Royal
des Sciences Naturelles de Belgique, Biologie 64:129-146.
Massin, C. 1996. Holothuries (Echinodermata) récoltées sur le
talus continental méditerranéen lors de la Campagne DEPRO
96. Mésogée 55:43-48.
Massin, C. and M. E. Hendrickx. 2010. A new species of
deep-water Holothuroidea (Echinodermata) of the genus
Synallactes from off western Mexico. Scientia Marína
McClain, C. R. 2004. Connecting species richness, abundance
and body size in deep-sea gastropods. Global Ecology
Biogeography 13:327-334.
McClain, C. R. and M. A. Rex. 2001. The relationship between
dissolved oxygen concentration and maximun size in deep-
sea turrid gastropods: an application of quantile regression.
Marine Biology 139:681-685.
Méndez, N. 2006. Deep-water polychaetes (Annelida) from the
southeastern Gulf of California, Mexico. Revista de Biología
Tropical 54:773-785.
669.indd 20 14/06/2011 03:42:46 p.m.
Revista Mexicana de Biodiversidad 82: 413-443, 2011
Mitsukuri, K. 1912. Studies on Actinopodous Holothurioidea.
Journal of the College of Science, Tokyo Imperial University
Mortensen, T. 1925. On a small Collection of Echinoderms from
the Antarctic Sea. Arkiv för Zoologi 17A:1-12.
Mortensen, T. 1927. Handbook of the echinoderms of the British
Isles. Humphrey Milford, Oxford University Press, London.
471 p.
Müller, J. 1850. Anatomische Studien über die Echinodermen.
Archiv für Anatomie und Physiologie. 1850:115-155; 225-
Müller, O. F. 1776. Zoologiae Danicae Prodromus, Havniae
1776. Copenhagen 274 p.
Nybakken, J. 2010. Deeper bottom habitats. III. Deep Sea
Sedimentary Megafaunal Communities (>2000 m
(last access: 01.II.2010).
Nybakken, J., S. Craig, L. Smith-Beasley, G. Moreno, A.
Summers and L. Weetman. 1998. Distribution density and
relative abundance of benthic invertebrate megafauna from 3
sites at the base of the continental slope off central California
as determined by camera sled and beam trawl. Deep-Sea
Research II 45:1753-1780.
Ohshima, H. 1915. Report on the holothurians collected by
the U.S. sheries steamer “Albatross” in the N.W. Pacic
during the summer of 1906. Proceedings of the United States
National Museum 48:213-291.
Olabarria, C. and M. H. Thurston. 2003. Latitudinal and
bathymetric trends in body size of the deep-sea gastropod
Troschelia barniciensis (King). Marine Biology 143:723-730.
Olabarria, C. and M. H. Thurston. 2004. Patterns of morphological
variation of the deep-sea gastropod Troschelia berniciensis
(King, 1846) (Buccinidae) from the northeastern Atlantic
Ocean. Journal of Molluscan Studies 70:59-66.
O’Loughlin, P. M. 2002. Report on selected species of BANZARE
and ANARE Holothuroidea, with reviews of Meseres Ludwig
and Heterocucumis Panning (Echinodermata). Memoirs of
Museum Victoria 59:297-325.
O’Loughlin P. M., M. E.Manchon-Cabeza and F. Moya Ruiz.
2009. Antarctic holothuroids from the Bellingshausen
Sea, with descriptions of new species (Echinodermata:
Holothroidea). Zootaxa 2016:1-6.
Östergren, H., 1904. Sitzung den 8 Mai 1903. Zoologische
Anzeiger 27:659.
Panning, A. 1949. Versuch einer Neuordnung der Familie
Cucumariidae (Holothuroidea; Dendrochirota). Zoologische
Jahrbücher. Abteilung für Systematik 78:404-470.
Parker, R. H. 1964. Zoogeography and ecology of some macro–
invertebrates, particularly mollusk, in the Gulf of California
and the continental slope off Mexico. Videnskabelige
Meddelelser fra Dansk naturhistorisk Forening i Kǿbenhavn
Pawson, D. L. 1969. Holothuroidea from Chile. Report No. 46
of the Lund University Chile Expedition 1948-1949. Sarsia
Pawson, D. L. 1977. Molpadiid sea cucumbers (Echinodermata:
Holothuroidea) of the southern Atlantic, Pacic and Indian
Oceans. Antarctic Research Series 26:97-123.
Pawson, D. L. 1983. Psychronaetes hanseni, a new genus
and species of elasipodan sea cucumber (Echinodermata:
Holothuroidea) from the eastern central Pacic. Proceedings
of the Biological Society of Washington 96:154-159.
Pawson, D. 2009. Holothuroidea (Echinodermata). In New
Zealand Inventory of Biodiversity. Volume One: Kingdom
Animalia, D. Gordon. (ed.). Sydney. 584 p.
Pawson, D. L. and E. J. Foell, 1986. Peniagone leander
new species, an abyssal benthopelagic sea cucumber
(Echinodermata, Holothuroidea), from the eastern central
Pacic Ocean. Bulletin of Marine Science 38:293-299.
Pawson, D. L. and C. Ahearn, 2001. Bathyal echinoderms of the
Galapagos Islands. In Echinoderms 2000, M. Barker (ed.). A.
A. Balkema, Rotterdam. p.41-46.
Pawson, D. L. and H. B. Fell. 1965. A revised classication of the
Dendrochirote holothurians. Breviora 214:1-7.
Perrier, R., 1902. Holothuries. In Expéditions Scientiques du
Travailleur et du Talisman pendant les années 1880, 1881,
1882, 1883. Ouvrage publié sous les auspices du Ministère
de l’instruction Publique sous la direction de A. Milne-
Edwards de 1888 à 1890 et continuée par E. Perrier. Masson
et Cie, Editeurs, Paris. p. 273-554, 11 plates.
Perrier, R. 1905. Holothuries antarctiques du Muséum d’Histoire
Naturelle de París. Annales des Sciences Naturelles Zoologie
Ser. 9:1-146.
Rex, M. A. and R. J. Etter. 1998. Bathymetric patterns of body
size: implications for deep-sea biodiversity. Deep-Sea
Research II 45:103-127.
Risso, A. 1826. Histoire naturelle des principales productions de
l’Europe méridionale et particulièrement de celles des environs
de Nice et des Alpes Maritimes. Levrault, Paris. 4, 439 p.
Rosenberg, R., W. E. Arntz, E. Chumán de Flores, L. A. Flores, G.
Carvajal. I. Finger and J. Tarazona. 1983. Benthos biomass
and oxygen deciency in the upwelling system off Peru.
Journal of Marine Research 41:263-279.
Rogers, A. D. 2000. The role of oceanic oxygen minimum zones
in generating biodiversity in the deep sea. Deep-Sea Research
II 47:119-148.
Samyn, Y, and Cl. Massin, 2003. The holothurian subgenus
Mertensiothuria (Aspidochirotida: Holothuriidae) revisited.
Journal of Natural History 37:2487-2519.
Sanders, H. L. and R. R. Hessler. 1969. Ecology of the deep-sea
benthos. Science 163:1419-1424.
Sastry, D. R. K. 2007. Echinodermata of India: An annotated list.
Records of the Zoological Survey of India, Occasional Paper
669.indd 21 14/06/2011 03:42:46 p.m.
434 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
Sibuet, M. 1977. Répartition et diversité des Echinodermes
(Holothurides-Astérides) en zone profonde dans le Golfe de
Gascogne. Deep-Sea Research 24:549-563.
Smith, C. R., L. A. Levin and S. Mullineaux. 1998. Deep-
sea biodiversity: a tribute to Robert R. Hessler. Deep-Sea
Research II 45:1-11.
Solís-Marín, F. A. 2003. Systematics and phylogeny of the
holothurian family Synallactidae. Ph.D. thesis, University of
Southampton, UK. 356 p.
Solís-Marín, F. A., H. Reyes-Bonilla, M. D. Herrero-Pérezrul,
O. Arizpe-Covarrubias and A. Laguarda-Figueras. 1997.
Sistemática y distribución de los equinodermos de la bahía
de La Paz. Ciencias Marinas 23:249-263.
Solís-Marín, F. A., A. Laguarda-Figueras, A. Durán-González,
C. Gust Ahearn and J. Torres Vega. 2005. Echinoderms
(Echinodermata) from the Gulf of California, Mexico.
Revista de Biología Tropical 53 (suppl. 3):123-137.
Solís-Marín, F. A., J. A. Arriaga-Ochoa, A. Laguarda-Figueras,
S. C. Frontana-Uribe and A. Durán-González. 2009.
Holoturoideos (Echinodermata: Holothuroidea) del Golfo
de California. Comisión Nacional par el Conocimiento y
Uso de la Biodiversidad e Instituto de Ciencias del Mar y
Limnología, UNAM, México, D.F. 177 p.
Thandar, AS., 1984. The holothurian fauna of Southern Africa.
Ph.D thesis, University of Durban-Westville, Durban. 566 p.
Thandar, A. S. 1998. A new genus and 3 new species of deep-
sea holothuroids from the west coast of South Africa
(Echinodermata). Journal of Zoology, London 244:79-88.
Thandar, A. 2008. Additions to the holothuroid fauna of the
southern African temperate faunistic provinces, with
descriptions of new species. Zootaxa 1697:1-57.
Théel, H. 1879. Preliminary report on the Holothuridae of the
exploring voyage of H.M.S. “Challenger” I. Kunglica
Svenska, Vetenskapsasakademiens Handlingar 5:1-20.
Théel, H. 1882. Report on the Holothurioidea dredged by H.M.S.
“Challenger” during the years 1873-1876, Part I: Reports of
the scientic results of the voyage of H.M.S. “Challenger”
1873-1876, 4:1-176.
Théel, H. 1886. Report on the Holothurioidea dredged by H.M.S.
“Challenger” during the years 1873-1876, Part II. Report on
the scientic results of the voyage of H.M.S. “Challenger”
during the years 1873-76. Zoology 14:1-290.
Tilot, V. 2006. Biodiversité et distribution de la mégafaune. Vol.
2. Atlas photographique annoté des échinodermes de la zone
de fractures de Clarion et de Clipperton. Paris, UNESCO/
IOC. IOC Technical Series 69:1-62.
Vaney, C. 1906a. Expédition Antarctique Française (1903-1905)
commandée par le Dr. Jean Charcot. Sciences naturelles:
Documents scientiques. Echinodermes. Holothuries.
Expédition Antarctique Française, Masson, Paris. 30 p.
Vaney, C. 1906b. Deux nouvelles holothuries du genre Thyone
provenant des Orcades du Sud. Bulletin du Muséum national
d’Histoire naturelle, Paris 12:400-402.
Verrill, A. E. 1867. Comparison of the tropical echinoderm fauna
of the east and west coasts of America. Transactions of the
Connecticut Academy of Arts and Sciences 1:339-351.
Verrill, A. E. 1868. Notice of a collection of echinoderms from
La Paz, Lower California, with descriptions of a new genus.
Transactions of the Connecticut Academy of Arts and
Sciences 1:371-376.
Verrill, A. E. 1870. Descriptions of echinoderms and corals from
the Gulf of California. American Journal of Science 2:93-100.
Verrill, A. E. 1871a. Additional observations on echinoderms,
chiey from the Pacic coast of America. Transactions of the
Connecticut Academy of Arts and Sciences 1:568-593.
Verrill, A. E. 1871b. On the echinoderm fauna of the Gulf
of California and Cape San Lucas. Transactions of the
Connecticut Academy of Arts and Sciences 1:593-596.
Zamorano, P., M. E. Hendrickx and A. Toledano Granados. 2006.
Distribution and ecology of deep-water mollusks from the
continental slope, southeastern Gulf of California, Mexico.
Marine Biology 150:883-892.
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Revista Mexicana de Biodiversidad 82: 413-443, 2011
Plate 1. A-D. Psolidium gracile Ludwig, 1894. A: ventral view. B: lateral view. C: lateroventral view. D: detail of lateroventral side
(arrows indicate a ventrolateral row of tube feet). E-G. Mitsukuriella unusordo sp. nov.(holotype). E: dorsal view. F: ventral view. G:
tentacle crow (arrows indicate long tentacles). H: Ypsilocucumis californiae sp. nov. (holotype) lateral view.
669.indd 23 14/06/2011 03:42:49 p.m.
436 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
Figure 1. Abyssocucumis albatrossi
(Cherbonnier, 1941). A: cross-shaped
ossssicles from body wall. B: cross-
shaped ossicles from tube feet. C: details
of the arm’s end of B. D: small, curved
rods from tentacles. E, F: large rods of the
Figure 2. Distribution of exam-
ined species of Dendrochirotida
off the Pacic coast of Mexico,
including previous records
(open symbols) and localities
where material was collected
(solid symbols).
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Figure 3. Psolidium gracile Ludwig, 1894. A:
calcareous ring (r: radial piece; ir: interradial
piece). B: small dorsal scales. C: perforated
plates from ventral sole (L= 16.7 mm). D:
perforated plates from ventral sole (L= 18.5
mm). E: end plate of ventral tube feet. F: rods
of the ventral tube feet. G: perforated plate of
the tube feet.
Figure 4. Psolidium gracile Ludwig,
1894. A: spiny curved rods of the
tentacles. B: elongated perforated
plates from tentacles. C: details of B
extremity. D, E, F: rods of the gonad.
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438 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
Figure 5. Mitsukuriella unusordo sp. nov.
(holotype) A: smooth body wall scales. B:
knobbed body wall scales. C-D: perforated
plates of tube feet. E: end-plate of the tube feet.
F: V-shaped rods from tube feet.
Figure 6. Mitsukuriella unusordo sp.
nov. (holotype) Rods of the tentacles.
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Revista Mexicana de Biodiversidad 82: 413-443, 2011
Figure 7. Distribution of
examined species of Dactylo-
chirotida off the Pacic coast
of Mexico, including previous
records (open symbols) and
localities where material was
collected (solid symbols).
Figure 8. Ypsilocucumis
californiae nov sp. (holotype)
A: calcareous ring (r: radial
piece; ir: interradial piece); B:
small body wall scales; C: large
multilayered body wall scales;
D: excentric spire of the large
body wall scale; E-F: rods of the
669.indd 27 14/06/2011 03:43:00 p.m.
440 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
Figure 9. Distribution of
examined species of Elasi-
poda off the Pacic coast
of Mexico, including previ-
ous records (open symbols)
and localities where mate-
rial was collected (solid
Figure 10. Laetmogone
scotoeides (H.L. Clark,
1913). A: large wheel
of body wall. B: small
wheels of the tube feet.
C: large wheel of the tube
feet. D: rods of the body
wall and tube feet. E:
irregular, branching rods
of the tube feet end-plate.
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Revista Mexicana de Biodiversidad 82: 413-443, 2011
Figure 11. Synallactes alexandri
Ludwig, 1893. A: cross-shaped
ossicles from body wall. B:
cross-shaped ossicles from dorsal
papillae. C: rods of the tube feet.
D: cross-shaped ossicle from tube
feet. E: pseudo table of tube feet.
F, G, H: rods of the tentacles.
Figure 12. Distribution
of examined species of
Aspidochirotida off the Pacic
coast of Mexico, including
previous records (open
symbols) and localities where
material was collected (solid
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442 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
Figure 13. Molpadia
intermedia (Ludwig,
1894). Tables of the tail.
Figure 14. Distribution
of examined species
of Molpadiida off the
Pacic coast of Mex-
ico, including previous
records (open symbols)
and localities were
material was collected
(solid symbols).
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Appendix 1. Remarks concerning doubtful or erroneous depth records in Solís-Marín et al. (2009).
Cucumaria crax Deichmann, 1941, is reported by Solis-Marin et al. (2009) in depths of 8-549 m. According to Maluf (1988), this
species occurs between 15 and 55 m depth. We could not locate a reference to the 549 m record. When the list of material cited by
Solis-Marin et al. (2009; 3 records) is examined, the only option for this record seems to be the one of “Bahía Sta. María”, 24°56.3’N,
108°44.6’W (the 2 others are cited by Maluf, 1988). However, there is no citation of a cruise sampling at that depth in this area and,
according to ecological data gathered in that area, bottom at ca 549 m depth is almost anoxic (Hendrickx and Serrano, 2010). In
addition, plotting the latitude-longitude data on a map indicates that the “Sta. María” station actually corresponds to a depth of less
than 200 m.
Pseudocnus californicus (Semper, 1868), in 0-717 m (Solis-Marin et al. 2009). An intertidal to 190m species according to Maluf
(1988). We were not able to locate a record at this depth. All records by Solis-Marin et al. (2009) are from intertidal or very shallow
water; only 1 is from the same sample as previous species (i.e., C. crax; Bahía Sta. María), but the authors provide a distinct maximum
depth range (717 m vs. 549 m for C. crax).
Holothuria leucospilota (Brands, 1835), in 0-695 m (Solis-Marin et al. 2009). An intertidal species according to Maluf (1988).
According to Samyn and Massin (2003), deeper record is from 8 m. We found no trace of a 695 m record in the Gulf of California or
in the eastern Pacic for this species.
Parastichopus californicus (Stimpson, 1857), in 5-5 640 m (Solis-Marin et al. 2009). From the intertidal to 180 m (Maluf, 1988) and to
216 m (Lambert, 1986). There seem to be no such depth in the Gulf of California. The 2 lots cited by Solis-Marin et al. (2009) are from
San Benedito and Angel de la Guarda Islands, certainly from much shallow water. Also, the reference to Théel (1886) in the synonymy
refers to Théel´s “Challenger” contribution but it should be the “Blake” contribution. See “Referencias Bibliográcas” in Solis-Marin
et al. (2009).
Chiridota aponocrita H.L. Clark, 1920, in 9-4 755 m (Solis-Marin et al. 2009). In the intertidal (Maluf 1988). Actually the data reported by
Solis-Marin et al. (2009) correspond to 1 of the 2 Albatross stations where another species, Protankyra abyssicola” (see H.L. Clark, 1920),
was collected (top of page 125 in H.L. Clark, 1920). The depth range reported for Ch. aponocrita is therefore due to a confusion.
669.indd 31 14/06/2011 03:43:08 p.m.
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... La contribución de Ludwig (1894) es considerada entre las más importantes para la región, pues en ésta describió 21 especies distribuidas por debajo de los 350 m de profundidad. Además de estos trabajos considerados como "clásicos", una amplia serie de especies nuevas encontradas en aguas profundas de la región han sido descritas por diversos autores (véase síntesis en Massin & Hendrickx, 2011) lo que incrementó de manera significativa el número de especies conocidas en el Pacífico este. ...
... El espécimen que aparece en la fotografía es parecido a una especie del género Abyssocucumis (= Staurocucumis), del cual hay dos especies señaladas en el Pacífico mexicano: A. albatrossi (Cherbonnier, 1941) y A. abyssorum (Théel, 1886) . La primera fue recolectada durante la campaña TALUD XIII en la cuenca del Carmen, a 2056-2195, y es conocida desde Oregon, USA, hasta Chiapas y en el Golfo de California (Massin & Hendrickx, 2011). La segunda tiene una distribución mucho más amplia, desde los 32ºN hasta Chile en el Pacífico este, incluyendo registros en el norte del Golfo de California así como en los océanos Atlántico e Indico (Maluff, 1988). ...
... Müller, 1776) y Psolidium gracile (Ludwig, 1894). El estatuto de P. squamatus fue discutido en detalles por Massin & Hendrickx (2011) quienes limitaron su posible distribución a la porción del Pacífico este que se extiende desde Centro-América hasta el mar de Bering. Al igual que las demás especies de Psolidae, P. squamatus requiere de un sustrato duro (e.g., rocas, conchas de moluscos) para poder asentarse lo cual reduce hasta cierto punto su distribución potencial en aguas profundas. ...
... Two major oceanographic cruises of the TALUD project were carried out off the west coast of the Baja California Peninsula, in NW Mexico (TALUD XV and TALUD XVI-B). These expeditions returned many specimens of sea cucumber (Holothuroidea) (see Luna-Cruz & Hendrickx 2018), including additional material of the recently described Ypsilocucumis californiae Massin & Hendrickx, 2011. The genus Ypsilocucumis Panning, 1949, is currently represented worldwide by two species: Y. asperrima (Théel, 1886), from the Atlantic Ocean, and Y. californiae, which was originally described for the Gulf of California, western Mexico, in the eastern Pacific (Massin & Hendrickx 2011). ...
... These expeditions returned many specimens of sea cucumber (Holothuroidea) (see Luna-Cruz & Hendrickx 2018), including additional material of the recently described Ypsilocucumis californiae Massin & Hendrickx, 2011. The genus Ypsilocucumis Panning, 1949, is currently represented worldwide by two species: Y. asperrima (Théel, 1886), from the Atlantic Ocean, and Y. californiae, which was originally described for the Gulf of California, western Mexico, in the eastern Pacific (Massin & Hendrickx 2011). Previously, the genus included two additional species, Y. turricata (Vaney, 1906) and Y. scotiae (Vaney, 1906), which have been transferred to Paracucumis Mortensen, 1925and Crucella Gutt, 1990respectively (O'Loughlin 2002O'Loughlin et al. 2009). ...
... Density of organisms (orgs/ha) was estimated using the swept area method (width of the sledge * distance of the haul). The identity of the specimens of Y. californiae was determined using traditional morphological taxonomy, using as main character the shape of the ossicles (see Massin & Hendrickx 2011). Specimens were compared to the holotype which is housed in the UNAM collection in Mazatlán. ...
Four specimens of the sea cucumber Ypsilocucumis californiae Massin & Hendrickx, 2011 were obtained during sampling operations off western Mexico. These specimens permit identification of this species as a member of the deep-water holothuroid community off the west coast of the Baja California Peninsula. Previous records correspond to four locations (including the type locality) in the Gulf of California, where eight specimens were collected. SEM ossicles images are provided for the first time and new ecological data associated with the presence of this species are available: temperature, 5.34‒8.38 °C; dissolved oxygen, 0.15‒0.28 ml O2/l and salinity, 34.42‒34.51 ups. The specimens were present in a wide variety of sediments with an organic carbon content of 3.18‒5.20 mg C/g (5.47‒8.95 % organic matter). Density values indicated low abundance of this species in the area (2.63‒3.94 orgs/ha). Records presented here were in a depth range from 540 to 776 m, which corresponds to the lower limit of the Oxygen Minimum Zone of the eastern Pacific. Additional records are provided for the West Atlantic Ypsilocucumis asperrima (Théel, 1886) and a key to the species of Ypsilocucumis is provided.
... More recently, Hendrickx (2010, 2011) studied a collection of 71 specimens collected below 300 m depth during the TALUD cruises (1991)(1992)(1993)(1994)(1995)(1996)(1997)(1998)(1999)(2000)(2001)(2002)(2003)(2004)(2005)(2006)(2007)(2008)(2009) in the same area and off the SW coast of Mexico, and reported 13 species, including 3 species new to science. Based on their studies and a compilation of previous literature, Massin and Hendrickx (2011) reported 31 species known from off the Pacific coast of Mexico, of which 20 have been recorded in the Gulf of California. ...
... Maluf (1988) included 627 entries in a review, each corresponding to a species, subspecies or unidentified genus that were known at that species. In the particular case of P. squamatus, Massin and Hendrickx (2011) briefly discussed the taxonomic complexity of the species of the "squamatus" complex, noting that further studies using molecular phylogeny are needed to elucidate the relationships among these and their geographic distribution. ...
... Diversity information of deep-sea echinoderm fauna through recent research is scarce (Pawson, 1982;Stöhr and Segonzac, 2005;Mecho et al., 2014;Moles et al., 2015;Calero et al., 2017;Mironov et al., 2018;Setyastuti and Wirawati, 2018;Stöhr and O'Hara, 2021). Strong progress has been made, especially for Colombia, Chile, Brazil, Argentina and Mexico (González et al., 2002;Borrero-Pérez et al., 2003;Benavides-Serrato and Borrero-Pérez, 2010;Campos et al., 2010;Manso, 2010;Massin and Hendrickx, 2011;Borrero-Pérez et al., 2012;Hendrickx et al., 2014;Solís-Marín et al., 2014;Martínez et al., 2014;Martínez et al., 2015;Martínez, 2016;Conejeros-Vargas et al., 2017;Martinez and Penchaszadeh, 2017;Martínez et al., 2017, Rivadeneira et al., 2017Luna-Cruz and Hendrickx, 2018;Borrero-Pérez et al., 2019;Flores et al., 2019;Pertossi et al., 2019;Martínez et al., 2019;Borrero-Pérez et al., 2020;Martinez et al., 2020;Rivadeneira et al., 2020;Luna-Cruz and Hendrickx, 2020;Catalán et al., 2020;Luna-Cruz and Hendrickx, 2021;Flores et al., 2021). ...
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Echinoderms are a highly diverse group and one of the most conspicuous in the deep sea, playing ecological key roles. We present a review about the history of expeditions and studies on deep-sea echinoderms in Costa Rica, including an updated list of species. We used literature and information gathered from the databases of the California Academy of Sciences, the Benthic Invertebrate Collection of the Scripps Institution of Oceanography, the National Museum of Natural History, the Museum of Comparative Zoology and the Museo de Zoología from the Universidad de Costa Rica. A total of 124 taxa (75 confirmed species) have been collected from the Costa Rican deep sea, 112 found in the Pacific Ocean, 13 in the Caribbean Sea, and one species shared between the two basins. We report 22 new records for the Eastern Tropical Pacific, 46 for Central American waters, and 58 for Costa Rica. The most specious group was Ophiuroidea with 37 taxa, followed by Holothuroidea (34 taxa), Asteroidea (23 taxa), Echinoidea (17 taxa), and Crinoidea (11 taxa). The highest number of species (64) was found between 800 m and 1200 m depth. Only two species were found deeper than 3200 m. Further efforts on identification will be required for a better comprehension of the diversity of deep-sea echinoderms. Limited research has been done regarding the biology and ecology of deep-sea echinoderms in Costa Rica, so additional approaches will be necessary to understand their ecological functions.
... The crinoid Hyocrinus foelli (Solís-Marín et al., 2013a;Solís-Marín et al., 2013b;Solís-Marín, Laguarda-Figueras, & Honey-Escandón, 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 Aporocidaris 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. ...
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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 elaborated 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 species richness.
... The affinity of the Gulf of California fauna is markedly tropical (Hendrickx 1992, Brusca & Hendrickx 2010, with a relatively high percentage of endemic species: 15.8% overall and about 26% in the case of the Gastropoda . Although many attempts have been made in order to divide the Gulf of California into faunistic units, distribution patterns of benthic invertebrates is far from being settled due to the frequent discovery of new species and reports on new distributions (e.g., Massin & Hendrickx 2011, Hendrickx & Ayón-Parente 2014, Tovar-Hernández et al. 2016, Gómez & Díaz 2017, Salgado-Barragán et al. 2017, Berschauer et al. 2018. ...
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Los invertebrados conforman el grupo más abundante y diverso entre los animales marinos, no obstante, son menos apreciados y estudiados en comparación con los vertebrados. En el Pacífico sur de México, formado por los estados de Guerrero, Oaxaca y Chiapas, está comprobada la enorme diversidad biológica terrestre que contiene; no es así el caso sobre su biodiversidad marina, en especial sobre los invertebrados marinos, cuya información parece ser escasa o muy dispersa. Con la intención de subsanar esta falta de información y cambiar la percepción sobre la biodiversidad de los invertebrados marinos del Pacífico sur de México, se preparó esta obra colectiva, la cual está compuesta de 10 capítulos realizados por especialistas. El inventario completo incluye casi 2,600 especies de invertebrados marinos registrados en la región, pertenecientes a 16 filos. La información de cada uno de los registros, su presencia en cada estado del Pacífico sur de México, así como su distribución batimétrica, entre otros datos de interés, puede ser consultada en cada uno de los capítulos respectivos.
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This contribution provides an overview of the scientific career of the late Dr Claude Massin (1948-2021), listing his scientific activities (academic career, participation to and organization of expeditions and scientific conferences, publications) as well as the taxa he described as new to science and the eponyms that were dedicated to him. The scientific career of Claude Massin is briefly sketched against the background of the personal family-life.
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It is proposed to name the Atlantic holothuria Psolidium complanatum Cherbonnier, 1969, whose denomination is already occupied by Psolidium complanatum (Semper, 1867) from the Philippines, as Psolidium bathygalego nom. nov. 843 specimens of Psolidium bathygalego nom. nov., collected between 417 and 1191mdeep in the NW and W of Galicia and in the Galicia Bank, were studied. A detailed description of the external and internal anatomy of Psolidium bathygalego nom. nov. is made by studying the ossicles and the skeletal structure by means of scanning electron microscopy (SEM) as well as the introvert, calcareous ring, retractor muscles, watervascular system, digestive system, respiratory trees, and reproductive system by means of micro-computed tomography (micro-CT). The habitat, feeding system, and geographical distribution of Psolidium bathygalego nom. nov. are also described.
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Between 1998 and 1999 the expedition INVEMAR-MACROFAUNA 1 investigated the upper continental slope of the Caribbean off Colombia at depths ranging from 200 to 500 m. The collection of echinoids comprised 714 individuals belonging to 7 orders, 10 families, 14 genera and 15 species. Stylocidaris lineata, Trigonocidaris albida, Echinocyamus grandiporus, Palaeobrissus hilgardi and Archaeopneustes hystrix are recorded for the first time in the Colombian Caribbean. Descriptions and identification keys are provided.