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Cah. Biol. Mar.

(2010) 51 : 459-466

Biodiversity of deep-sea organisms associated with


sunken-wood or other organic remains sampled
in the tropical Indo-Pacific
Sarah SAMADI1, Laure CORBARI1, Julien LORION1, Stphane HOURDEZ2,3, Takuma HAGA4, Jolle DUPONT5,
Marie-Catherine BOISSELIER1 and Bertrand RICHER DE FORGES1
(1) Systmatique, Adaptation et Evolution, UMR 7138 UPMC-IRD-MNHN-CNRS (UR IRD 148), Musum National
d'Histoire Naturelle, Dpartement Systmatique et Evolution, CP 26, 57 Rue Cuvier, F-75231 Paris Cedex 05, France
E-mail: sarah@mnhn.fr ; tel: 33 1 40 79 37 59; fax: 33 140 79 38 44
(2) UPMC Univ Paris 06, UMR 7144, Gntique de ladaptation au milieu extrme, Station Biologique de Roscoff,
Place G. Teissier, 29680 Roscoff, France
(3) CNRS, UMR 7144, Gntique de ladaptation au milieu extrme, Station Biologique de Roscoff,
Place G. Teissier, 29680 Roscoff, France
(4) Department of Biological Science, Graduate School of Science, The University of Tokyo, 113-0033,
7-3-1 Hongo, Bunkyo-ku, Tokyo
(5) Origine, Structure et Evolution de la Biodiversit", UMR 7205 MNHN-CNRS, Musum National d'Histoire Naturelle,
Dpartement Systmatique et Evolution, 57 Rue Cuvier, F-75231 Paris Cedex 05, France.

Abstract: Since 80s, fauna associated to sunken woods has been collected during exploration cruises of the Tropical DeepSea Benthos cruises program (TDSB). In 2004 we started a specific program of cruises devoted to the exploration of
sunken-wood habitats. Seven cruises provided sunken-wood organisms in depth ranging from 100-1 500 metres, at three
locations corresponding to large basins of sunken-wood accumulation in the south west Pacific: (i) Philippines, (ii) Salomon
islands and (iii) Vanuatu. Extra-samples from other TDSB cruises are also available. Indeed, far from these basins, pieces
of woods are unevenly catch by trawling and dredging and these erratic substrata, found for example on seamounts, are
usually colonized by a typical sunken wood fauna. To enhance the collection of species associated to organic substrata that
are difficult to catch at the deep sea floor, we also immersed down traps containing different kinds of baits: pieces of wood,
blue whale bones, green turtle shell, and stag horn. These experiments deployed between 12-30 months off New Caledionia
and Vanuatu. Samples yielded an abundant and original fauna, notably Bathymodiolinae and Galatheids. Specific diversity
of collected zoological groups with taxa specific to this habitat was explored using molecular tools. Here this paper, we
report the preliminary finding about the biodiversity of major zoological groups but also fungi found during this program
of cruises.
Keywords: Sunken-wood l Exploration cruises l Biodiversity l Deep-sea

Reu le 16 fvrier 2010 ; accept aprs rvision le 19 aot 2010.


Received 16 February 2010; accepted in revised form 19 August 2010.

460

SUNKEN-wOOD FAUNA

Introduction
The presence of animals on plant remains at the deep-sea
floor was first documented by the Challenger expedition
(1872-1876). Indeed, during this pioneering circumnavigation, organisms associated with plant remains were
sampled at 9 stations off the Philippines and New Guinea,
at depth ranging from 1500 to 3900 m (Murray, 1895). The
Galathea expedition (1950-1952) also recorded plant
remains at 56 stations at depth ranging from 400 to 1050 m
(Bruun, 1959). The organisms associated with these
remains were studied mostly from a taxonomic point of
view. For example Knudsen (1961) examined the woodboring bivalves of the Xylophagaidae and identified 30
species, among which 17 were new to science. Incidentally,
he also mentioned the presence of others organisms
associated with plant debris, sunken in the deep-sea. Until
recently, just as in Knudsen papers, apart from taxonomic
considerations, such organisms were mainly looked at as
zoological and/or ecological curiosities and thus only
anecdotally studied.
The interest for these organisms was renewed by Distel
et al. (2000) that revealed the evolutionary affinities of the
giant hydrothermal vent mussels with those found on pieces
of sunken wood or whale falls. This close phylogenetic
relationship was then confirmed by several studies (see
references in Lorion et al., 2010). Moreover, it was shown
that mussels sampled on whale bones or on pieces of
sunken-wood harbor chemosynthetic bacteria in their gills,
just as their hydrothermal-vent or cold-seep relatives
(reviewed in Duperron et al., 2009). Based on these phylogenetic results, Distel et al. (2000) formulated the
wooden-step hypothesis according to which, mussels
found at hydrothermal vents and cold seeps evolved
recently from ancestors associated with organic falls.
Moreover, paleontological data confirmed that many
organisms found at deep-sea vents, including the mussels,
belong to lineages much younger than previously thought,
challenging previous ideas about the evolutionary history
of the fauna found at vents and seeps (Little & Vrijenhoek,
2003). However, until recently, mostly, only the spectacular
fauna associated with whale falls have been studied (see for
example the review of Fujiwara et al., 2007)
In this context, a series of expeditions launched in the
early 1980s by the Institut de la Recherche pour le
Dveloppement (formerly ORSTOM) and the Museum
National dHistoire Naturelle (MNHN), following the
traces of the famous historical expeditions, explored the
deep-sea benthos of the tropical Indo-Pacific at depths
ranging from 100 to 1500 m. This series of cruises, allowed
the establishment of an impressive collection of tropical
deep-sea animals that is studied thanks to an informal
network of taxonomists. The cruises, the network of

taxonomists and the publications are gathered under the


umbrella Tropical Deep-Sea Benthos. A review of this
program, focusing on the results obtained for mollusks, is
available (Bouchet et al., 2008). Apart from the spectacular
discoveries of species belonging to lineages known only
from the fossil record (e.g. Hemicrinidae, Bourseau et al.,
1987), the most amazing result of this loose program is the
amount of species newly described (cf. data for mollusks in
Bouchet et al., 2008). As in the historical expeditions,
these cruises repeatedly pulled up from the deep-sea floor
plant remains, principally around the islands, that were
usually heavily colonized by a remarkable fauna.

Materials and Methods


To more accurately sample these organisms, a series of
cruises (gathered under the banner Biodiversit des
Organismes Associs aux bois couls BOA) devoted to
the exploration of sunken wood piles started in 2004. Seven
cruises provided specimens associated with plant remains
in the depth range explored by the Tropical Deep Sea
Benthos program (100-1500 m) from three important
basins in the southwest Pacific where sunken wood is
known to accumulate (Fig. 1): (i) Philippines, (ii) Salomon
islands and (iii) Vanuatu. Additional samples from other
cruises, taking place far from these basins, also provided
specimens associated with organic remains (Fig. 1). Indeed,
for example on seamounts, small pieces of wood and seeds
(e.g. coconuts) are unevenly pulled up by the trawls or the
dredges and these erratic substrates, are usually also
colonized by the organisms typically found on pieces of
wood at accumulation areas.
This sampling is however biased toward substrates that
are easy to localize at the deep-sea floor (such as large
pieces of wood off the mouth of the rivers of forested
islands). Therefore, to more accurately catch the diversity
of the organisms associated with all kinds of organic
substrates that may sink to the deep-sea floor, traps covered
by netting with mesh of 3 mm were deployed (Fig. 2). The
baits put in the traps were: seeds and pieces of wood of
various tree species, sugar canes, whale bones, green turtle
shell, stag horn, feathers, cephalopod beaks and cuttlebones. A first line of traps was deployed in 2003 for 20
months at 1100 m depth, off the barrier of New Caledonia.
Then, in 2005, three lines were deployed, two, respectively
at 500 and 400 m depth, off Santo Island in Vanuatu for one
year and one at 900 m depth off New Caledonia for 30
months. These traps yielded an abundant and remarkable
macrofauna, notably Bathymodiolinae settled on turtle
bones (see Lorion et al., 2010 for more details) but also
galatheids that were not thought to be tightly associated
with organic falls.
These collections are under study by the network of

S. SAMADI, L. CORBARI, J. LORION, S. HOURDEZ, T. HAGA, J. DUPONT, M.-C. BOISSELIER, B. RICHER DE FORGES

461

Figure 1. Location of Tropical deep-sea benthos cruises (TSBS) with emphasis on BOA cruises and others cruises providing wood
samples. Boxes mention respectively: cruise name (year, number of stations).
Figure 1. Localisation des campagnes Benthos profond tropical (TSBS) et dtail des campagnes BOA et de celles ayant
chantillonn des bois couls. Les encarts mentionnent le nom de la campagne (lanne, le nombre de stations).

taxonomists of the Tropical Deep Sea Benthos program but


other studies, on other biological aspects, are also starting
(e.g. study of the diet and the microflora in the gut content
of a galatheid crab, Hoyoux et al., 2009 or of a sea-urchin,
Becker et al., 2009; botanical identification of the plant
remains, Pailleret et al., 2007; prokaryotic diversity,
Palacios et al., 2009). Although the taxonomic results are
mostly not yet available, we give here an overview of the
diversity of some of the major taxa collected by the BOA
cruises on organic remains at the deep-sea floor. Images for
some of these taxa are provided in Figure 2.

Results and Discussion


Mollusks
Table1 provides a preliminary list of mollusks collected on
pieces of wood of various sizes or on seeds, during the
BOA1 cruise in Vanuatu. Specimens were sorted in the
field to families and then separated into morphospecies.
Many of the identified families are also known from

hydrothermal vents (e.g. Lepetidae or Mytilidae). Others


are known only from wooden substrates (e.g.
Xylophagaidae, Teredinidae or Pectinodonta spp.) or
organic substrates (e.g. Cocculinidae). Most of this material
is still under study for a more accurate identification but the
first available results suggest higher species diversity
within families. For example, among leptochitons, 17
species attributed to 3 genera were identified by an
anatomical study, of which 4 need a new name (Sigwart,
2008). Similarly, using an integrative approach to
taxonomy, Lorion et al. (2010) showed that mytilids
collected during this cruise may be divided into 10 species.
Crustaceans
Isopods, amphipods and decapods but also barnacles were
found among plants debris in the trawls or the dredges. It is
however very difficult to know to what extent they depend
on the decaying plant substrates, in particular for the
vagrant fauna. Among the decapods, the main groups
repeatedly caught with such substrata are the galatheids (for
example Munidopsis andamanica MacGilchrist, 1905, M.
nitida (A. Milne Edwards, 1880), M. bispinosculata Baba,

462

SUNKEN-wOOD FAUNA

Table1. Diversity of Mollusca observed during the BOA1


cruise (2005).
Tableau 1. Diversit des mollusques observs au cours de la
campagne BOA1 (2005).
Family
POLYPLACOPHORA
NEOLORICATA

GASTROPODA
PATELLOGASTROPODA
VETIGASTROPODA

NERITOPSINA
COCCULINIFORMIA
NEOMPHALINA
CAENOGASTROPODA

HETEROBRANCHIA
BIVALVIA
MYTILOIDA
MyOIDA

TOTAL

Number of
morphospecies

Lepidopleuridae
Ischnochitonidae
Acathochitonidae

3
1
1

Pectinodontidae
Lepetidae
Pseudococculinidae
Trochidae
Turbinidae
Skeneidae
Phenacolepadidae
Cocculinidae
Neomphalidae
Iravadiidae
Vitrinellidae
Buccinidae
Columbellidae
Nassariidae
Orbitestellidae
Pyramidellidae

1
1
1
1
2
5
1
9
1
1
2
1
2
1
1
2

Mytilidae
Pholadidae
Xylophagaidae
Teredinidae

4
1
24
6
72

1988, M. similior Baba, 1988, or Munida rhodonia


Macpherson, 1994), two different families of symmetrical
hermit-crabs (all the species of the genus Xylopagurus and
several species of the genus Pylocheles inhabit centimetresized pieces of wood), thalassinids (e.g. Callianassa
amboinensis De Man, 1888) and alpheids. The same
organisms were also collected in the traps that were sunken
off New Caledonia or Vanuatu. The analysis of the diet of
M. andamanica showed that this species effectively feeds
on wood (Hoyoux et al., 2009).
The examination of isopoda and amphipoda diversity
from several BOA cruises revealed the occurrence of five
main families of amphipods and three families of isopods
(Table 2) living on wood substrates. Among amphipoda, the
most diversified and abundant family was the
Lysianassidae. All the specimens of melitid amphipod have
been identified as belonging to the same genus
Bathyceradocus and probably to new species. Microscopic
and ultra-structural investigations have revealed that these
specimens were wood-feeders (Corbari, pers. com.).
Moreover, the presence of bacterial ectosymbiosis on the
mouthparts of this species suggests an original pathway of
nutrition which could involve bacteria in wood
degradation. Only three species have been described in the
genus Bathyceradocus, one of them, B. wuzzae (Larsen &
Krapp-Schikel, 2007) has been recently discovered on
wood substrates in the vicinity of hydrothermal vents.
Polychaetes
Numerous annelids, sipunculids and echiurian were
collected on, or inside, pieces of wood and other organic
substrates, but one can guest that a lot of the animals on the
surface of the substrates were washed away during
collection. A preliminary identification of the BOA-

Figure 2. From the left to the right and top down: The traps before being sunken. Colonized pieces of woods from the traps after 20
months at 1100 metres. Pieces of wood pulled up during BOA1 cruise off Santo (Vanuatu). A typical content of a trawl pulled up off Salomon
islands during SALOMONBOA cruise. Adipicola longissima (Mytilidae) settled on a coconut, Munidopsis nitida (Galatheidae) from a trap,
Adipicola sp. (Mytilidae) settled in a nipa nut, small lucinid (Lucinidae) found within the fiber of sunken coconut, Pectinodonta sp.
(Pectinodontidae) on a large piece of wood, chiton (Lepidopleuridae) settled on a coconut, sea-star (Caymanostellinae) attached to a piece
of wood, Xylopagurus sp. (Paguroidea) within a small piece of wood, Idas sp. settled on turtle bones from a trap, limpet (Cocculinidae) and
sea-star (Caymanostellinae) on the same piece of wood, Asterechinus elegans (Echinidae) on a piece of wood, Teredo sp. (Teredinidae)
extracted from a sunken trunk, isopods Limnoria sp. (Limnoridae) and Oceanitis scuticella (Ascomycota) on a piece of sugar cane, limpet
(Cocculinidae) on a leaf.
Figure 2. De gauche droite et de haut en bas : les cages avant dtre coules ; pices de bois colonises dans les cages aprs 20 mois
passs 1100 mtres ; pices de bois remontes au cours de la campagne BOA1 au large de Santo (Vanuatu) ; contenu typique dun dragage remonts du large des Iles Salomon au cours de la campagne SALOMONBOA ; Adipicola longissima (Mytilidae) tabli sur un cocotier ; Munidopsis nitida (Galatheidae) rcolt dune cage ; Adipicola sp. (Mytilidae) tabli dans une noix de nipa ; petit lucinid (Lucinidae)
dans la fibre de coctier coul ; Pectinodonta sp. (Pectinodontidae) sur un grand morceau de bois ; chiton (Lepidopleuridae) tabli sur un
cocotier ; toile de mer (Caymanostellinae) accroch un morceau de bois ; Xylopagurus sp. (Paguroidea) dans un petit morceau de bois ;
Idas sp. tabli sur des os de tortue dans une cage ; gastropode (Cocculinidae) et toile de mer (Caymanostellinae) sur le mme morceau de
bois ; Asterechinus elegans (Echinidae) sur un morceau de bois ; Teredo sp. (Teredinidae) extrait dun bois coul ; isopodes Limnoria sp.
(Limnoridae) et Oceanitis scuticella (Ascomycota) sur un morceau de canne sucre ; gastropode (Cocculinidae) sur une feuille.

S. SAMADI, L. CORBARI, J. LORION, S. HOURDEZ, T. HAGA, J. DUPONT, M.-C. BOISSELIER, B. RICHER DE FORGES

463

464

SUNKEN-wOOD FAUNA

Echinoderms

Table 2. Diversity of the amphipods and isopods collected during


BOA cruises.
Tableau 2. Diversit des crustacs amphipodes et isopodes rcolts
au cours des campagnes BOA.

Ophiurids, sea-urchins and sea-stars were repeatedly


found on sunken wood during BOA cruises. Among
the ophiurids many of the identified species are also
Order
Family
Estimated Depth range (m)
nb
known from other deep-sea environments and are
nb
of different genera stations
thought to be opportunistic. For example, among the
Amphipoda Eusiridae
1
1414-1422
1
specimens sampled during the SalomonBoa cruise, the
1
464-819
2
Epimeridae
very common tropical species Ophiactis definite
Lysianassidae
3
537-1545
17
Koehler, 1922 was abundantly found (OHara, pers.
Melitidae
1
532-1750
8
com.). However Ophiambix sp., described as woodOedicerotidae
1
504-1750
2
eating (Paterson & Baker, 1988), was also found
Isopoda
Cirolanidae
1
537-1160
11
(OHara,
pers. com.). As far as sea-urchins are
Limnoridae
1
300-1500
18
concerned, at least one species was identified in this
Sphaeromatidae 1
450-500
3
collection (Asterechinus elegans Mortensen, 1942).
The analysis of the gut content and the associated
microflora supports that it is indeed a wood-eating
collection of polychaetes revealed a total of 19 families
organism
(Becker et al., 2009). Finally, the most famous
(comprising at least 28 genera). These families cover a
echinoderm
associated with sunken wood is Xyloplax
wide diversity of trophic strategies (Fauchald & Jumars,
turneae
Rowe,
Baker & Clark, 1988, primarily described as
1979), from deposit-feeders to predators, indicating a
a
new
class
of
echinoderms
(Concentrocycloidea) but then
complex ecosystem on, or near, sunken wood piles and
placed
within
the
Asteroidea
(Janies & Mooi, 1998). In
other organic substrates. Some of these families are also
Solomon
Islands,
Vanuatu
or
Philippines we repeatedly
found at hydrothermal vents and cold seeps (Table 3). In
collected
specimens
attributed
to
the Caymanostellidae, a
particular, the phyllodocid genus Protomystides, and the
family
reported
as
associated
with
wood (Fujita et al.,
maldanid genus Nicomache are common in both these
1994).
However,
their
phylogenetic
relationship with
deep-sea environments.
Xyloplacidae is not yet resolved. Preliminary molecular
data (not shown) suggest that the
Caymanostellidae from Solomon Islands are
Table 3. Polychaetes associated with sunken wood collected in BOA cruises.
very diversified in this region.

* indicates families also found at hydrothermal vents and cold seeps.


Tableau 3. Polychtes associs aux bois couls rcolts au cours des
campagnes BOA. * dsigne les familles galement trouves au niveau des
sources hydrothermales et des suintements froids.
Family

Amphinomidae*
Aphroditidae
Capitellidae*
Chaetopteridae*
Chrysopetalidae*
Cirratulidae
Eunicidae*
Euphrosinidae*
Glyceridae*
Goniadidae
Hesionidae*
Lumbrineridae*
Maldanidae*
Nereididae*
Onuphiidae
Phyllodocidae*
Polynoidae*
Sigalionidae*
Terebellidae*

Estimated nb
of genera
2
1
2
1
1
1
2
2
1
1
1
1
2
2
1
1
3
2
1

Depth range
(m)
275
721
256
1230
441
548
381
414
381
447
256
420
256
83
721
173
190
190
431

883
773
900
1306
441
560
1306
1750
1034
503
354
741
1054
1285
773
680
1285
553
445

Nb
Nb ind.
stations Examined
8
2
2
1
1
1
22
12
3
1
2
3
8
14
1
4
23
6
1

36
2
12
1
1
2
118
16
3
1
3
4
9
63
1
20
64
6
1

Fungi
Recent molecular ecology studies suggested
a high fungal diversity in the deep-sea and
one can guest that fungi play an important
role in the decomposition of wood also in the
deep-sea (see references in Dupont et al.,
2009). However very few data are available
and the diversity identified using molecular
ecology approaches need to be completed by
evidences that the corresponding organisms
are truly living at the deep-sea bottom.
During BOA cruises, pieces of wood were
examined to find sporulating fungi. From
Vanuatu and Solomon Islands two different
species of deep-sea ascomycetes were identified Oceanitis scuticella Kohlmeyer, 1977
and Alisea longicolla Kohlmeyer, 1977
(Dupont et al., 2009). Oceanitis scuticella
has been described from sunken wood and is
among the only four species that have been
described from such substrates at depths
below 1600 m. This species was sampled in

S. SAMADI, L. CORBARI, J. LORION, S. HOURDEZ, T. HAGA, J. DUPONT, M.-C. BOISSELIER, B. RICHER DE FORGES

this area at depths ranging from 600 to 1300 m. Alisea


longicolla is a new species from a new genus that was
sampled at depths ranging from 600 to 800 m. Both species
fall into the main group of marine ascomycetes
(Halosphaeriaceae). while O. scutella is close to shallow
water species, the position of A. longicolla remains
unresolved within the Halosphaeriacea.
Perspectives
This preliminary review of the results obtained from this
series of cruises confirms the originality and the diversity
of the fauna associated with the organic remains sampled at
the deep-sea floor. Some of the organisms (e.g. mytilids)
have close relatives in other chemosynthetic environments
but others (e.g. Xylophagaidae) belong to lineages
specialized on sunken wood. The available data suggest
that many organisms feed on wood and that bacterial
associations are often involved. There are also evidences of
association to sulfur-oxidizing bacteria for some taxa.
However, much of the diversity at the specific level
remains to be described as well as the phylogenetic
relationships with organisms from others chemosynthetic
environments.
Acknowledgments
we would like to thank the crew of RV ALIS; P. Bouchet
and the malacological staff in Paris; Nadia Amziane and
the staff of the marine invertebrates collection at the
MNHN; all the taxonomists involved in the Tropical Deep
Sea Benthos Network, and more particularly H. Kano, E.
Macpherson, B. Marshall, R. T. OHara, J. Sigwart, R. von
Cosel, and A. waren; F. Gaill, P. Compere, C. de Ridder, M.
Zbinden, O. Gros, S. Dupperon, P. Becker C. Hoyoux, N.
Le Bris who are involved in the BOA program and in the
European network Diwood; all other participants to the
BOA cruises (S. Tillier, F. Rousseau, P. Lopez).

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