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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 Pacic 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 Pacíco 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.
*Correspondent: michel@ola.icmyl.unam.mx
Abstract. Research cruises aboard the R/V “El Puma” were organized to collect deep-water benthic and pelagic
specimens off the Pacic coast of Mexico. Seventy four specimens of Holothuroidea were collected off the Pacic 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 identied 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 Pacic Mexico Oxygen Minimum Zone. An updated checklist of species of
Holothuroidea recorded below 350 m depth along the Pacic 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 Pacic, Mexico, new species.
Resumen. Una serie de campañas oceanográcas fue organizada a bordo del B/O “El Puma”, frente a las costas del
Pacíco 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 identicadas 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 Pacíco 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 Pacíco mexicano. Se anexa una lista actualizada de las especies de Holothuroidea
registradas en profundidades mayores a 350 m frente a la costa del Pacíco 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, Pacíco este,
México.
Introduction
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;
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414 Massin and Hendrickx.- New Mexican Pacic 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 Pacic
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 Pacic, 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
Pacic (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 Pacic 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 Pacic to
56. Of these 56 species, 5 were originally described from
other regions of the world and later reported for the Central
eastern Pacic: 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 Pacic;
Ceraplectana trachyderma H.L. Clark, 1907, from the
NW Pacic; and Molpadia musculus Risso, 1826, from the
Mediterranean.
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 Pacic. 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
identied 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 veried 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 Pacic (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 Pacic 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 Pacic
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.; identicator; coll.,
collector. Based primarily on Maluf (1988) contribution,
a list of all species with at least 1 record within Mexican
waters of the Pacic 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.
Results
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
identied 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).
Descriptions
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, textg. 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
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416 Massin and Hendrickx.- New Mexican Pacic 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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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
unidentiable 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 Pacic 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),
Remarks
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 dened 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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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 Pacic,
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,
textg. 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 dened
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.
Remarks
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 Pacic are not conspecic.
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-
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420 Massin and Hendrickx.- New Mexican Pacic 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 Pacic 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
difcult 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 identied 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 identication 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 Pacic 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 Pacic, 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.
Remarks
According to Maluf (1988) there are 6 species of
Psolidium along the Central eastern Pacic 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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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 Pacic 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).
Remarks
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 identication.
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
radius.
Remarks
The presence of 15 digitate tentacles and of a calcareous
ring without posterior processes are characteristic of the
family Vaneyellidae as dened 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. inexa
(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. inexa 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. inexa.
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
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422 Massin and Hendrickx.- New Mexican Pacic 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,
difcult 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).
Remarks
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 (Pacic
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,
textgs. 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).
Remarks
Ypsilothuria bitentaculata is considered a
cosmopolitan species, collected in the East and West
Pacic 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,
1894).
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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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
Pacic (Maluf and Brusca, 2005). Widely distributed
along the Pacic 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.
Remarks
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 Pacic, 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 Pacic, and features a
very wide Indo-Pacic 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)
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424 Massin and Hendrickx.- New Mexican Pacic 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 ramied rods (g 10E).
Remarks
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 identied 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 Pacic 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 Pacic coast of Mexico. It
conrms the restricted distribution of L. scotoeides which
seems to be endemic of Pacic 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.
Remarks
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
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425
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-Pacic, 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).
Unidentied 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).
Remarks
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 difcult 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.
10-19.
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.
Remarks
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
Pacic, all from water deeper than 350 m, between SW
Mexico and northern Peru. The only previous record of
the genus for Pacic Mexico is for Synallactes ishikawai
f. ind. (= S. sagamiensis Augustin, 1908), reported from
the Gulf of Tehuantepec, SW Mexico, by Parker (1964).
Whether this identication (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
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426 Massin and Hendrickx.- New Mexican Pacic 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).
Remarks
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
nudum).
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.
Remarks
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 Pacic 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 Pacic (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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427
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.
Remarks
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 Pacic,
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 Pacic 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.
Discussion
It should be emphasized that in most surveys either
survey methods, sampling effort, or depth range are distinct,
thus rendering formal comparison difcult. 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
identied at species level and 2 at genus level) have at least
1 record from off the Pacic 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
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428 Massin and Hendrickx.- New Mexican Pacic 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 Pacic (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 difculty 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 Pacic 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 veried (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-Pacic 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 conrms the
presence of a second species of the genus in Mexican
waters and is the rst conrmed 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 conrms the abundance and high occurrence of this
species along western Mexico. Synallactes alexandri is
recorded for the rst time in Mexican waters.
Ecology
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
Pacic 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 conrm its afnity
669.indd 16 14/06/2011 03:42:45 p.m.
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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 Pacic. Bluhm (1994) reported on holothurians
collected in manganese module sites off northern Peru (ca
90°W) and in the central Pacic, but none of his species
(most identied 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 identied 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 Pacic 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 Pacic deep-water Holothuroidea
al., 1985) with Pacic 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 Pacic Mexico OMZ, which
is not present in the 2 other areas. Rockall Trough and the
Mexican Pacic 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 Pacic. Yet, every single study
has fully demonstrated that even limited sampling effort
on the bathyal seaoor 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 Pacic.
Acknowledgments
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.
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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.
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436 Massin and Hendrickx.- New Mexican Pacic 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
tentacles.
Figure 2. Distribution of exam-
ined species of Dendrochirotida
off the Pacic 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 Pacic 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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Figure 7. Distribution of
examined species of Dactylo-
chirotida off the Pacic 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
tentacles.
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440 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
Figure 9. Distribution of
examined species of Elasi-
poda off the Pacic coast
of Mexico, including previ-
ous records (open symbols)
and localities where mate-
rial was collected (solid
symbols).
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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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 Pacic
coast of Mexico, including
previous records (open
symbols) and localities where
material was collected (solid
symbols).
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442 Massin and Hendrickx.- New Mexican Pacic deep-water Holothuroidea
Figure 13. Molpadia
intermedia (Ludwig,
1894). Tables of the tail.
Figure 14. Distribution
of examined species
of Molpadiida off the
Pacic 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 Pacic 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 Bibliográcas” 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.
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