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Two projects coordinated by the Research & Development Center of the Brazilian energy company PETROBRAS have collected benthic physical and image samples at Campos Basin, Brazilian continental margin, between 21-22°S. The Synallactidae (Echinodermata: Holothuroidea) constitute a very representative group amongst the holothurians. Taxonomic details of the sampled material are condensed here. Pseudostichopus depressus Hérouard, 1902 had been reported by Deichmann (1930), and this species is now referred as Molpadiodemas depressus (Hérouard, 1902). Also, here is the first register of Mesothuria verrilli (Théel, 1886) in Brazilian waters.
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Echinoderms: Durham – Harris et al. (eds)
© 2010 Taylor & Francis Group, London, ISBN 978-0-415-40819-6
Synallactidae (Echinodermata: Holothuroidea) from Campos Basin,
Southwest Atlantic
R.B. Moura & L.S. Campos
Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, RJ, Brazil
M.P. Curbelo-Fernandez
Departamento de Biologia Marinha, Instituto de Biologia,Universidade Federal do Rio de Janeiro, RJ, Brazil
Consultant for Centro de Pesquisas e Desenvolvimento Leopoldo Américo Miguez de Mello – CENPES/
PETROBRAS, RJ, Brazil
G.H. Cavalcanti
Centro de Pesquisas e Desenvolvimento Leopoldo Américo Miguez de
Mello – CENPES/PETROBRAS, RJ, Brazil
ABSTRACT: Two projects coordinated by the Research & Development Center of the Brazilian energy com-
pany PETROBRAS have collected benthic physical and image samples at Campos Basin, Brazilian continental
margin, between 21–22S. The Synallactidae (Echinodermata: Holothuroidea) constitute a very representative
group amongst the holothurians. Taxonomic details of the sampled material are condensed here. Pseudosticho-
pus depressus Hérouard, 1902 had been reported by Deichmann (1930), and this species is now referred as
Molpadiodemas depressus (Hérouard, 1902). Also, here is the first register of Mesothuria verrilli (Théel, 1886)
in Brazilian waters.
1 INTRODUCTION
In general, about 40 holothurian species from 11 fam-
ilies have been recorded in Brazilian waters to date
as in Tommasi (1999). Tiago & Ditadi (2001) recog-
nized this holothurian fauna as dominated by species
with a wide geographical distribution that reach at
least the Caribbean. The three families of the Order
Aspidochirotida (Holothuriidae, Stichopodidae and
Synallactidae) have been represented by 8 species in
Brazil (Tommasi 1999).
Synallactids are found almost exclusively in the
deep-sea (Solis-Marín 2003). They possess shield-
shaped tentacles, lack tentacle ampullae, have their
gonads in one or two tufts, and the ossicles, when
present, include basically tables and rods (Solis-Marín
2003, Solis-Marín et al. 2004, O’Loughlin & Ahearn
2005).
According to Tommasi (1999), Pseudostichopus
depressus Hérouard, 1902 was the only synallac-
tid Holothurian recorded for Brazilian waters. He
included it into the Brazilian list because of the ‘Alba-
tross Results’ report on holothurians from theWestern
part of the Atlantic (Deichmann 1930).
More recently, energy industry programmes have
contributed to a significant increase in the knowl-
edge on the Southwest Atlantic deep-sea fauna. Both
Campos Basin Deep-Sea Environmental Project’ and
Campos Basin Deep-Sea Coral Assessment Project’
are main Brazilian initiatives coordinated by the
Research and Development Center of the Brazilian
energy company PETROBRAS (CENPES). Through
these programmes, echinoderms have been sampled
and observed in situ, and have been found as one of
the most abundant megabenthic groups. The Synallac-
tidae has been the most conspicuous group amongst
the holothurians, and this paper aims to provide
information on their taxonomy, depth and geographic
distribution, and general biology wherever possible.
2 MATERIAL & METHODS
Synallactid specimens have been sampled from the
Brazilian Continental Margin at Campos Basin,
Southwest Atlantic with the use of an Otter Trawl
Semi-Balloon (OTSB) from 1100 to 1700m, and a
Remoted Operated Vehicle (ROV SCV 3000 from
Acergy Offshore) from 700 to 1100m (Fig. 1).
In February and August, 2003, 6 replicate trawls
were accomplished at each of 1100, 1300 and 1600m
depth, covering a total area of 1.2km2. Specimens
were fixed on board in 4% formaline and preserved
in 70% ethanol.
During November 2004, a preliminary survey was
carried out where parallel transects of 50 m each were
viewed with the ROV along 8 selected areas. This
survey provided images of benthic communities asso-
ciated with deep-sea coral banks and soft sediments
nearby.
245
Figure 1. Study sites, at Campos Basin, SWAtlantic.
External and internal anatomy based on original
descriptions, keys and further reviews by Solis-Marín
(2003) and O’Loughlin & Ahearn (2005) were used
for taxonomic identification.
Ossicles were isolated especially from the spec-
imens body wall and tentacles using concentrated
sodium hypochlorite solution. They were washed off
with distilled water, mounted on slides, analyzed, and
photographed using a Carl Zeiss®Axioscop2 light
microscope equipped with a video camera. The speci-
mens studied here have been deposited at the collection
of Echinodermata, Department of Zoology at the Insti-
tute of Biology, Federal University of Rio de Janeiro
(UFRJ), Brazil.
3 TAXONOMY
Order Aspidochirotida Grube, 1840
Family Synallactidae Ludwig, 1894
Genus Mesothuria Ludwig, 1894
Mesothuria verrilli (Théel, 1886a)
Holothuria verrilli Théel, 1886a; (passim).
Mesothuria verrilli (Théel, 1886a); Deichmann,
1930: 93–94, pl. 6 figs. 1–8; Deichmann, 1940:
192–193; Deichmann, 1954: 385.
Non Holothuria verrilli Théel, 1886a; Marenzeller,
1893: 7–9, pl. 1 fig. 2, pl. 2 fig. 2.
Non Mesothuria verrilli (Théel, 1886a); Östergren,
1986: 345; Perrier, 1902: 307–312, pl. 16 figs. 22–31;
Hérouard, 1923: 10–13; Mortensen, 1927: 381–382;
Grieg, 1932: 4.
Non Allantis intestinalis var. verrilli (Théel,
1886b); Hérouard, 1902: 18–21, pl. 1 fig. 306.
Mesothuria gargantua Deichmann, 1930: 96-96
(passim).
Material examined. 17 specimens: Campos Basin,
Oceanprof I – IBZ/UFRJ-EH0176 (n =2), IBZ/UFRJ-
EH0177 (n =4); Oceanprof II – IBZ/UFRJ-EH178
(n =5), IBZ/UFRJ-EH179 (n =4), IBZ/UFRJ-EH180
(n =2).
Distribution. Reliable records in the Caribbean, off
Ambergris Cay (Belize), from 700 to 1800 m, but may
be found deeper (Solis-Marín 2003).
Figure 2. Side view of Mesothuria verrilli (Théel, 1886).
Scale bar =50 mm.
Figure 3. Ossicles from a 205mm specimen of Mesothuria
verrilli: (A) Side view of body wall quadriradiate tables;
(B) Upper view of body wall quadriradiate tables; (C) Spiny
rods from the tentacles. Scale bars: (A), (B) =100 µm,
(C) =200 µm.
Diagnosis. (after Théel 1886a, Deichmann 1930,
and Solis-Marín 2003). Specimens studied are
72–225 mm long. Body cylindrical as a whole and
elongated. Body surface whitish to dirty yellow in
color. Skin thick and wrinkled in preserved specimens
(Fig. 2). Tentacles about 20 in number, pale brown-
ish in color, small and retracted in most specimens.
Mouth and anus located almost terminally on each
end. Pedicels small and hard to observe in larger spec-
imens, but easily found on the ventral surface near
the anus. Body wall ossicles consist of well developed
quadriradiate tables usually with circular to irregular
margin disk (95–150 µm in diameter) and moderate to
low spire (38–76µm) (Fig. 3). Disk has a large cen-
tral hole surrounded mostly by 8–11 holes of different
sizes. Range of number of the marginal holes tend to
increase from 8–11 (8 and 10 more common in 72 mm
specimen) to 8–13 (8,9,10 and 11 more common in
175 mm specimen) and decrease back to 8–11 (9 and
10 more common in 225 mm specimen). Another 2–14
smaller accessory holes can be present, especially in
larger specimens. Spire composed by 4 pillars united
by 1, or sometimes 2, transverse beams. Tips of the
pillars a little diverging, smooth or bearing 2 or 3
spinelets. Tentacles supported by straight to slightly
bent rough spiny rods of 175–346 µm long.
Four sedentary individuals of this species were
observed on muddy substrate during the ROV survey
246
Figure 4. Individual of Mesothuria verrilli at Campos
Basin, Southwest Atlantic, 1015m depth: (A) Overall body
shape; (B) Detail of the body showing attached pteropod
shells and pebble like structures.
Figure 5. Dorsal and ventral view of Molpadiodemas
depressus (Hérouard, 1902). Scale bar =30 mm.
at depths greater than 1000 m. All of them had their
bodies partially encrusted with pteropod shells, sharp
sponge spicules, and black unidentified pebble-like
structures (Fig. 4).
Remarks.A comprehensive review on the taxonomy
and phylogeny of the Synallactidae was performed by
Solis-Marín (2003). The genus Mesothuria comprises
of 27 valid species and had never been reported in
the Brazilian continental margin before. This record
increases this species distribution from the Caribbean
to Campos Basin, Southwest Atlantic. Sampling and
conservation methods may strongly affect the qual-
ity of synallactids external morphology, which can
influence the identification. Even specimens care-
fully collected by submersibles have arrived on board
in very poor conditions (Pawson, 1982). Combined
taxonomy and imaging of live specimens may help
understand not only the main external morphological
features, but also their biology and behavior.
Genus Molpadiodemas Heding, 1935
Molpadiodemas cf. depressus (Hérouard, 1902)
O’Loughlin & Ahearn, 2005
Pseudostichopus depressus Hérouard, 1902: 15–16,
pl. 2 figs 15–18.
Platystichopus depressus Heding, 1940: 353–358.
Material examined. 7 specimens: Campos Basin,
Oceanprof I – IBZ/UFRJ-EH0181 (n =2), IBZ/UFRJ-
EH182 (n =2), IBZ/UFRJ-EH0183 (n =1); Ocean-
prof II – IBZ/UFRJ-EH184(n =1), IBZ/UFRJ-EH185
(n =1).
Figure 6. Molpadiodemas depressus (40 mm long speci-
men): (A) Anteroventral view, mouth and brim. (B) Detail
of foraminiferan shells attached to its body wall.
Figure 7. Braidedcur vedrods from the tentacles of a 36 mm
specimen. Scale bar =100µm.
Distribution. North and South Atlantic Ocean, West
European Basin, Sargasso Sea, off Bahamas and West
Indies from 1353 to 5690 m.
Diagnosis. (after Hérouard, 1902; Deichmann,
1930 and O’Loughlin & Ahearn, 2005). Specimens
are 38–60 mm in length when preserved. Body dorso-
ventrally flat, not tapered, surrounded by a thick round
broad brim (Fig. 5). Body wall opaque and gelatinous.
Pale faint body surface, dorsally smooth and ventrally
wrinkled, often encrusted with foraminiferan shells
(Fig. 6). Mouth anterior and ventral. Tentacles shield-
shaped in form and partially outwards in most of the
specimens. Anal opening situated in a posterior ver-
tical furrow. Small and delicate tube feet arranged all
over the body, but more conspicuous on the dorsal sur-
face, near the furrow and brim. Very distinctive double
flat midventral longitudinal muscles bands, attached
to internal body wall. Body wall and gonad ossicles
completely lacking. Ossicles in tentacles consist of
different patterns of large narrow braided curved rods
from 183–320 µm long (Fig. 7). No distinctive dor-
sal madreporite was observed, such as illustrated by
Hérouard (1902) for Pseudostichopus depressus.
Remarks. Both genera Molpadiodemas Heding,
1935 and Pseudostichopus Théel, 1882 were rediag-
nosed by O’Loughlin & Ahearn (2005). As a conse-
quence, Pseudostichopus depressus Hérouard, 1902
has been recently reassigned to Molpadiodemas. This
genus is cosmopolitan and it is represented by 16 valid
species. Molpadiodemas is mainly characterized by
the absence of tube feet or papillae along the paired
radii, flat longitudinal muscular bands attached to the
inner body wall, branched gonad tubules and rods in
the tentacles. Further review on the taxonomic charac-
ters used in the identification of this species needs to
be addressed and included in the list of holothurians
reported for Brazilian waters.
247
4 CONCLUDING REMARKS
A revision on the Brazilian holothurian key is neces-
sary, but new deep-sea sampling activities are likely to
occur.These should provide a better understanding of
bathymetric and spatial distribution of the holothuri-
ans. As the very large Brazilian Marine ecosystem is
hydrologically and topographically complex with con-
trasting dominant ecosystems, with the presence of
major river outflows, canyons, deep-water reefs, with
different types of substrates and the continental mar-
gin subject to different water masses effects (Lavrado,
2006), there is still much to be found and described
in the Southwest Atlantic. Further comparisons of the
fauna from this region with that found in other areas of
the Atlantic are also necessary to recognize biogeog-
raphy patterns of these organisms, and especially in
deeper waters, to test the degree of cosmopolitanism
or endemism closest to seamount areas for instance
and adjacent abyssal plain.
Also, combined sampling strategies should pro-
vide a better understanding on feeding, reproduction,
growth, development, and general life history pro-
cesses, biology and behavior including interactions
with other species. For the first time echinoderms have
been recorded in situ at the Brazilian continental mar-
gin (Campos et al. in prep/ or this volume if accepted).
Certainly, this has helped in the identification of the
deep-sea holothurians described herein, as those origi-
nated from OTSB, other trawling or dredging methods
often damage the organisms, and do not allow their
real picture in the natural environment.
ACKNOWLEDGEMENTS
We would like to thank CENPES/PETROBRAS for
the opportunity of sampling and studying this unique
material. To Dr. Anaize Henriques (Universidade Fed-
eral do Rio de Janeiro) for using the facilities of the
Laboratory of Image Capturing – PROIN/Capes to
make the images of the ossicles. We are also grateful
to Dr. Francisco Solis-Marín (Universidad Nacional
Autónoma de México), Mrs. Cynthia Ahearn (National
Museum of Natural History, Smithsonian Institu-
tion), Mr. P. Mark. O’Loughlin (Museum of Victoria)
and Dr. Ahmed Thandar (University of KwaZulu-
Natal) for providing appropriate literature and
support.
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... For instance, the last new holothuroid species was described by Freire & Grohmann (1989) (Leptosynapta brasiliensis) and few contributions have been made since then (e.g. Tiago & Ditadi, 2001;Moura et al., 2010). Furthermore, most of the genera need revision, especially with the use of new techniques (e.g. ...
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Pseudostichopus villosus var. violaceus Théel is raised to species status; Pseudostichopus alatus Imaoka, P. trachus Sluiter and P. nudus Ohshima are junior synonyms of Pseudostichopus mollis Théel; Pseudostichopus globigerinae Hérouard and P. dilatorbis Imaoka are junior synonyms of Meseres (= Molpadiodemas) involutus Sluiter; Pseudostichopus propinquus Fisher and P. lapidus Hérouard are junior synonyms of Meseres (= Pseudostichopus) peripatus Sluiter; and Pseudostichopus arenosus Ohshima and P. molpadioides Ohshima are junior synonyms of Meseres (= Pseudostichopus) hyalegerus Sluiter. New combinations are: Molpadiodemas atlanticus (Perrier), M. depressus (Hérouard), M. involutus (Sluiter), M. pustulosus (Sluiter), M. villosus (Théel), Pseudostichopus elegans (Koehler and Vaney), P. hyalegerus (Sluiter), P. papillatus (Djakonov), P. peripatus (Sluiter) and P. spiculiferus (O'Loughlin). Other species discussed are: Molpadiodemas violaceus (Théel), Pseudostichopus aemulatus Solís-Marín and Billett, P. echinatus Thandar, P. mollis Théel, P. occultatus Marenzeller, and P. profundi Djakonov. A key is provided for the species of Molpadiodemas and Pseudostichopus. Tables are provided of genera and subgenera with current systematic status, and species with original and current combinations. Editorial alert re misguiding editorial mistake in O’Loughlin & Ahearn 2005 concerning pygal-furrowed species of Synallactidae and genus Meseres Ludwig by Mark O’Loughlin (July 2013). O’Loughlin 2002 maintained the referral of many pygal-furrowed synallactid species to genus Meseres Ludwig, 1894. See: 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(2): 297-325. This paper was in part superseded by a comprehensive review of pygal-furrowed synallactid species by O’Loughlin & Ahearn 2005. Most of the species previously assigned to Meseres Ludwig, 1894 were then assigned to Molpadiodemas Heding, 1935 or Pseudostichopus Théel, 1886. Meseres Ludwig, 1894 was listed and discussed incertae sedis. See: O’Loughlin, P.M. and Ahearn, C. 2005. A review of pygal-furrowed Synallactidae (Echinodermata: Holothuroidea), with new species from the Antarctic, Atlantic and Pacific oceans. Memoirs of Museum Victoria 62(2): 147–179. The publication of O’Loughlin & Ahearn, 2005 was accompanied by two editorial “disasters”, and one of them remains at best confusing and at worst misleading. The late Cynthia Ahearn and I were not given proofs before publication and the first evidence of the disasters was in the hard copy volume after publication. First error: All of the published figures were little better than rorschsch blotches as the printer seemed to have confused printing colour and black & white images. Neither the editor nor the publisher noticed this, and Cynthia and I had not been given an opportunity to see the proofs. After a long struggle with the editor he agreed to correct the figures and produce a hard copy offprint. The figures in the on-line version (see above) are correct and as originally submitted. Second error: In the figure captions that were submitted for publication we used “M.” for “Molpadiodemas” to abbreviate the many usages of Molpadiodemas. In the context of the paper the “M.” could not have referred to anything but “Molpadiodemas”. Understandably the editor judged that the first “M.” in each caption should be the full generic name. But he then unilaterally and most unfortunately inserted “Meseres” when the genus should very obviously have been “Molpadiodemas”. Cynthia and I were never consulted. Nothing could convince the editor to correct his mistake. So in the printed and on-line versions of the paper the figure captions 4, 5, 6, 7, 8 and 12 have Meseres when it should be Molpadiodemas.