The Mediterranean Deep-Sea Ecosystems

Lepidion lepidion( Risso, 1810)
Vincent Fossat, 1879.

Coll. Musum dHistoir e natur elle de Nice.
MHNN.
The Mediterranean
deep-sea ecosystems
An overview of their diversity,
structure, functioning
and anthropogenic impacts,
with a proposal for their conservation
Aristeus antennatus
Alber t I
er
Pr ince de Monaco. Camp. scient. Pnids pl. III. 1908.
EL. Bouvier del, M. Bor r el pinx.
Coll. Muse ocanogr aphique, Monaco.
The Mediterranean
deep-sea ecosystems

.
The Mediterranean
deep-sea ecosystems
This document has been pr epar ed under the coor dination of
Ser gi Tudela
Fisher ies Coor dinator WWF- Mediter r anean Pr ogr amme Ofce
and Fr anois Simar d
Mar ine Pr ogr amme Coor dinator IUCN Centr e for Mediter r anean Cooper ation,
and IUCN Global Mar ine Pr ogr amme
I U C N T h e Wo rld C o n se rva tio n U n io n
N o ve m b e r 2 0 0 4
Published by: IUCN Centr e for Mediter r anean Cooper ation, Mlaga, Spain
and WWF Mediter r anean Pr ogr amme, Rome, Italy
Copyr ight: 2004 Inter national Union for Natur e and Natur al Resour ces
Citation: WWF/ IUCN ( 2004) . The Mediter r anean deep-sea ecosystems: an over view of their diver sity,
str uctur e, functioning and anthr opogenic impacts, with a pr oposal for conser vation. IUCN, Mlaga
and WWF, Rome.
Par t I. Car tes, J.E., F. Maynou, F. Sar d, J.B. Company, D. Llor is and S. Tudela ( 2004) . The
Mediter r anean deep-sea ecosystems: an over view of their diver sity, str uctur e, functioning and
anthr opogenic impacts. In: The Mediterranean deep-sea ecosystems: an overview of their
diversity, structure, functioningand anthropogenic impacts, with a proposal for conservation.
IUCN, Mlaga and WWF, Rome. pp. 9-38.
Par t II. Tudela S., F. Simar d, J. Skinner and P. Guglielmi ( 2004) . The Mediter r anean deep-sea
ecosystems: a pr oposal for their conser vation. In: The Mediterranean deep-sea ecosystems:
an overview of their diversity, structure, functioningand anthropogenic impacts, with a
proposal for conservation. IUCN, Mlaga and WWF, Rome. pp. 39-47.
ISBN: 2-8317-0846-X
Layout and
pr oduction by: Fr anois-Xavier Bouillon, Maximedia SARL, Cagnes-sur -Mer F-06800.
Pr inted by : Per fecta SARL, Villeneuve-Loubet F-06270.
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The text of this book is printed on recycled paper.
The designation of geogr aphical entities in this book, and the pr esentation of the mater ial, do not imply the expr ession
of any opinion whatsoever on the par t of IUCN concer ning the legal status of any countr y, ter r itor y, or ar ea, or of its
author ities, or concer ning the delimitation of its fr ontier s or boundar ies.
The views expr essed in this publication do not necessar ily r eflect those of IUCN.
Cor e suppor t to the activities of the IUCN Mediter r anean office is pr ovided by the Junta de Andalucia, and the
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Acknowledgements
This document has been made possible thanks to the gener ous contr ibution of many exper ts fr om
all ar ound the Mediter r anean and over seas, who have demonstr ated that cor e pr inciple of science
as a tool for r aising the level of human under standing - namely by contr ibuting their knowledge
to the cr eation of a pr oposal to ensur e the long-ter m sustainability of Mediter r anean deep-sea
ecosystems. In this r egar d, the coor dinator s wish to sincer ely thank, in par ticular , Dr Bela Galil,
Dr Helmut Zibr owius, Dr Angelo Tur si, Dr Cinta Por te, Dr Fr ancesco Mastr ototar o, Dr Phil Weaver ,
Dr Nr ia Teixidor , Dr . Giuseppe Notar bar tolo di Sciar a, Dr . Chedly Ras, Dr Maur izio Wr tz, and
Dr Gr aeme Kelleher , for their pr ecious contr ibutions.
Special r ecognition is given to Dr Fr der ic Br iand, Dir ector Gener al of CIESM, and to his team, who
made possible the or ganisation, jointly with IUCN, of a specific r ound table on the pr otection of
the Mediter r anean deep-sea within the fr amewor k of the 2004 CIESM congr ess, held in Bar celona.
The final ver sion of this document, the thr ust of which was outlined ther e by the coor dinator s,
benefited enor mously fr om the ensuing discussion held amongst the top level scientists and legal
exper ts gather ed ther e. The coor dinator s ar e also indebted to the CIESM Wor kshop Monogr aph
n 23 on Mediter r anean deep-sea ecosystems, which inspir ed some aspects of this document.
Dr Alber to Gar ca, Chair man of the GFCM-SAC Subcommittee on the Mar ine Envir onment and
Ecosystems ( SCMEE) , kindly endor sed the pr esentation and later discussion of a dr aft ver sion of
this document dur ing the 2004 meeting of the SCMEE.
Dur ing the last decades a consider able effor t has been made by differ ent exper ts in the knowledge
of deep sea biology of the Mediter r anean. Although we ar e still far to fully under stand the str uctur e
and dynamics of these par ticular ecosystems, the coodinator s want to acknowledge the effor t made
by the author s to summar ize this knowledge in the pr esent document. Their impr essive scientific
exper tise on Mediter r anean deep-sea biology affor ds full scientific legitimacy to the subsidiar y
conser vation pr oposals pr esented her e.
We acknowledge also Jean Jauber t and Anne-Mar ie Damiano fr om the Oceanogr aphic Museum
of Monaco for allowing us to use illustr ations fr om the Campagnes ocanographiques du
Prince Albert I
er
; Br igitte Chamagne-Rollier fr om Nice Natur al Histor y Museum for letting us use
water color s fr om its collections; Michel Huet and Pascale Bouzanquet, Inter national Hydr ogr aphic
Bur eau, Monaco; Geor ge F. Shar man, Mar ine Geology & Geophysics Division National Geophysical
Data Center ; John Campagnoli, NOAA/ NESDIS, National Geophysical Data Center ; Car la J. Moor e,
NOAA National Geophysical Data Center and Wor ld Data Center for Mar ine Geology and Geophysics;
Angelo Tur si, Dipar timento di Zoologia, Univer sit degli Studi, Bar i; Dwight Coleman, Univer sity
of Rhode Island Gr aduate School of Oceanogr aphy and Rosita Stur m, Spr inger ; Fr eder ic Melin,
Joint Resear ch Centr e of the E.C. Institute for Envir onment and Sustainability Inland and Mar ine
Water s; Jean Mascle, Gosciences Azur , Obser vatoir e Ocanologique de Villefr anche, CNRS; Michel
Voisset, Cather ine Satr a-Le Br is and Benot Loubr ieu, Ifr emer and Nelly Cour tay, Editions Ifr emer .
Thank you also to Fr anois-Xavier Bouillon for collecting illustr ations and to Miles Heggadon for
r eviewing the English language.
List of acronyms
CBD Convention on Biological Diver sity
CIESM Inter national Commission for Scientific Explor ation of the Mediter r anean Sea
CMIMA Centr e Mediter r ani dInvestigacions Mar ines i Ambientals
CSIC Consejo Super ior de Investigaciones Cientficas
DHAB Deep Hyper saline Anoxic Basin
EEZ Exclusive Economic Zone
GEBCO Gener al Bathymetr ic Char t of the Oceans
GFCM Gener al Fisher ies Commission for the Mediter r anean
GFCM-SAC GFCM Scientific Advisor y Committee
GFCM-SAC-SCMEE GFCM SAC Sub-Committee on Mar ine Envir onment and Ecosystems
ICES Inter national Council for the Explor ation of the Sea
ISA Inter national Seabed Author ity
IUCN The Wor ld Conser vation Union
MCPA Mar ine and Coastal Pr otected Ar eas
MPA Mar ine Pr otected Ar eas
POM Par ticulate Or ganic Matter
RFMO Regional Fisher ies Management Or ganisation
SPA Pr otocol Specially Pr otected Ar eas
SPAMI Specially Pr otected Ar eas of Mediter r anean Inter est
UNCLOS United Nation Convention on the Law of the Sea
UNICPOLOS United Nations Open-ended Infor mal Consultative Pr ocess on Oceans and the Law of the Sea
UN GA United Nations Gener al Assembly
WSSD Wor ld Summit on Sustainable Development
WWF Wor ld Wide Fund for Natur e
TABLE OF CONTENTS
Foreword .................................................................................................... 8
Part 1
The Mediterranean deep-sea ecosystems
A n o ve rvie w o f th e ir d ive rsity, stru c tu re ,
fu n c tio n in g a n d a n th ro p o g e n ic im p a c ts
1. Introduction ........................................................................................ 10
2. Deep-sea diversity patterns
2.1. Fauna .................................................................................................. 11
2.2. Structure of deep-water mediterranean communities ....................... 19
3. Functioning of deep-sea Mediterranean food webs
3.1. Overview ............................................................................................. 22
3.2. Mesoscale variations in the dynamics of deep-sea ecosystems ....... 23
3.3. Human alteration of trophic webs ...................................................... 27
4. Unique environments of the Mediterranean deep sea
4.1. Cold seeps ......................................................................................... 29
4.2. Brine pools ......................................................................................... 31
4.3. Deep-sea coral mounds ..................................................................... 31
4.4. Seamounts ......................................................................................... 32
5. Anthropogenic impact in the deep Mediterranean
5.1. Fisheries ............................................................................................. 35
5.2. Other anthropogenic threats to the deep-sea Mediterranean ............ 36
Part 2
A p ro p o sa l fo r th e ir c o n se rva tio n
6. Deep-water habitat protection and sheries management
6.1. The particular status of Mediterranean waters ................................... 40
6.2. International policy context ................................................................ 40
6.3. A conservation proposal tailored to the Mediterranean ..................... 42
References .............................................................................................. 48
Glossary of key ecological concepts .................................................. 55
Plates ....................................................................................................... 57
8
THE MEDI TERRANEAN DEEP- SEA ECOSYSTEMS. PART 1 .
The Wor ld Summit on Sustainable Development ( WSSD)
( Johannesbur g, 2002) highlighted the need to pr omote
ocean conser vation, namely:
1. Maintaining the pr oductivity and biodiver sity of im-
por tant and vulner able mar ine and coastal ar eas
( MCPAs) , including ar eas within and beyond na-
tional jur isdiction;
2. Encour aging the application of the ecosystem ap-
pr oach to ocean and fisher ies management by
2010; and
3. Developing and facilitating the use of diver se ap-
pr oaches and tools, including the establishment of
MPAs consistent with inter national law and based
on scientific infor mation, together with r epr esenta-
tive networ ks by 2012.
The last Confer ence of the Par ties of the Convention on
Biological Diver sity ( CBD-CoP7, Kuala Lumpur , 2004)
agr eed to adopt this appr oach for the wor k of the
Convention on mar ine and coastal pr otected ar eas, and
to develop a str ategy to meet this goal.
The CBD acknowledging that: The establishment and
maintenance of marine and coastal protected are-
as that are effectively managed, ecologically based
and contribute to a global network of marine and
coastal protected areas, buildingupon national and
regional systems, includinga range of levels of pro-
tection, where human activities are managed, par-
ticularly through national legislation, regional pro-
grammes and policies, traditional and cultural prac-
tices and international agreements, to maintain the
structure and functioningof the full range of marine
and coastal ecosystems, in order to provide benefits
to both present and future generations, str essed the
impor tance to addr ess the pr otection of the biodiver sity
beyond National jur isdiction. It also agr ees that ther e
is an ur gent need for inter national cooper ation and ac-
tion to impr ove conser vation and sustainable use of bi-
odiver sity in mar ine ar eas beyond the limits of national
jur isdiction, including the establishment of fur ther ma-
r ine pr otected ar eas consistent with inter national law,
and based on scientific infor mation, including ar eas
such as seamounts, hydr other mal vents, cold-water cor -
als and other vulner able ecosystems. The CoP7 invited
the Par ties to r aise their concer ns r egar ding the issue of
conser vation and sustainable use of genetic r esour ces
of the deep seabed beyond limits of national jur isdiction,
and to identify activities and pr ocesses under their jur is-
diction or contr ol which may have significant adver se
impact on deep seabed ecosystems and species.
This WSSD call has been endor sed by the par ticipants
of the Mar ine Theme at the V
th
Wor ld Par ks Congr ess, in
Dur ban, South Afr ica ( 8-17 September 2003) .
The Malaga Wor kshop on High Seas Pr otected Ar eas
( Januar y 2003) str essed, inter alia, that the level of sci-
entific knowledge of the high seas, and the deep-sea in
par ticular , needs to be r aised.
The pr esent document ther efor e ar ises fr om a joint in-
itiative between the WWF Mediter r anean Pr ogr amme
and the IUCN Centr e for Mediter r anean Cooper ation, to
addr ess these issues at a r egional level; it contains, for
the fir st time, a compr ehensive pr oposal for conser va-
tion fir mly based on the scientific infor mation cur r ently
available.
Dr aft ver sions of this document wer e cir culated for
consultation amongst concer ned stakeholder s and dis-
cussed in sever al r elevant meetings ( GFCM-SAC-SCMEE,
CIESM) . This final ver sion has gr eatly benefited fr om
the feedback gener ated by this consultation pr ocess; the
conser vation pr oposals included r eflect a wide consen-
sus among scientists and legal exper ts fr om all ar ound
the Mediter r anean.
Foreword
9
AN OVERVI EW OF THEI R DI VERSI TY, STRUCTURE, FUNCTI ONI NG AND ANTHROPOGENI C I MPACTS.
Part 1
A n o ve rvie w o f th e ir d ive rsity, stru c tu re ,
fu n c tio n in g a n d a n th ro p o g e n ic im p a c ts
J o a n E . C a rte s, F ra n c e sc M a yn o u , F ra n c e sc S a rd , J o a n B . C o m p a n y, D o m in g o L lo ris
Institut de Cincies del Mar, CMIMA-CSIC
Passeig Martim de la Barceloneta 37-49, 08003 Barcelona, Spain
and
S e rg i Tu d e la
WWF Mediterranean Programme
Carrer Canuda 37, 3
er
, 08002 Barcelona, Spain
10
Fi g. 1 . Geogr aphy of the Mediter r anean sea.
Credits: International Bathymetric Chart of theMediterranean, published 1981 by theHead Department of Navigation and Oceanography,
St. Petersburg, Russia, on behalf of theInternational Oceanographic Commission (GEBCO, 2003).
Modified. Se e co l o ur pl ate , p. 5 7 .
1. Introduction
Once consider ed as lifeless domains, deep-sea habitats
ar e at pr esent, fr om a biodiver sity viewpoint, consider ed
exceptional ecosystems that har bour singular tr ophic
webs. This per spective has ar isen fr om pioneer ing r e-
sear ch car r ied out in Ear ths major oceanic ar eas, par -
ticular ly the Pacific and the Atlantic, since the mid 19
th

and dur ing the 20
th
centur ies. A major landmar k was the
discover y in 1977 in the easter n Pacific of ecosystems
based entir ely on chemosynthetic* pr imar y pr oduction,
linked to deep-sea hydr other mal vents* ( Cor liss et al.,
1979) . Mor eover , deep-sea ecosystems ar e now the ul-
timate tar get of industr ial fisher ies wor ldwide, follow-
ing the r elentless depletion of fish communities on the
continental shelves, in a sor t of ( over ) fishing down
the bathymetr ic r ange effect ( Mer r ett and Haedr ich,
1997) .
The fir st data on the pr esence of or ganisms living in
the bathyal* domain was pr esented by Risso ( 1816)
examining specimens deliver ed to him by local fisher -
men in Nice ( Fr ance) , obtained at depths between 600
and 1000 m. Other data collected dur ing the 19
th
cen-
tur y points to the pr esence of life at bathyal and abyss-
al* depths, despite the long-standing extr apolation by
For bes fr om obser vations in the Aegean sea in 1841
claiming the existence of an azoic, or lifeless, domain
deeper than 600 m ( Gage and Tyler , 1991) . The fir st sys-
tematic oceanogr aphic expeditions to the deep-sea wer e
the Challenger expedition ( 1872-1876, Gage and Tyler ,
1991) and, in the Mediter r anean, the Pola( 1890-1893) ,
the Dana ( 1908-1910) and Thor ( 1921-1922) expedi-
tions ( Bas, 2002) .
The Mediter r anean contains sea-beds up to 5000 m
deep. The maximum depth is 5121 m in the Matapan-
Vavilov Deep, off the Souther n coast of Gr eece, with an
aver age depth of 2500 m. The Mediter r anean sea har -
bour s most of the same key geomor phologic str uctur es
such as submar ine canyons, seamounts, mud volca-
noes or deep tr enches that have pr oven to tr anslate
into a distinctive biodiver sity makeup in other r egions
in the wor ld; r ecent findings have indeed confir med
this ( including the pr esence of chemosynthetic tr ophic
webs) . A fur ther unique featur e of the Mediter r anean is
that it har bour s one of the few war m deep-sea basins in
the wor ld, wher e temper atur es r emain lar gely unifor m
at ar ound 12.5-14.5C at all depths, with high salinity
( 38.4-39.0 PSU) and high oxygen levels ( 4.5-5.0 ml l
-1
;
Miller et al., 1970; Hopkins, 1985) . The constant tem-
per atur e and salinity r egime of the Mediter r anean con-
tr asts with the Atlantic at compar able latitudes, wher e
temper atur e decr eases and salinity incr eases with depth.
Another impor tant issue is the r elative isolation of deep-
sea communities, not only with r espect to those of the
Atlantic ( the maximum depth of the sill of Gibr altar is
ca. 300 m) , but also between those in the Easter n and
Wester n Mediter r anean, separ ated by the shallow Sicily
Channel ( ca. 400 m, fig. 1) . All these featur es r einfor ce
the potential for unique deep-sea communities in the
Mediter r anean, and the impor tance of pr ecautionar y ac-
tion to limit the impact of human activities on these fr ag-
ile habitats.
* Key ecological concepts ar e mar ked with an aster isk the r st time they appear , and dened in the Glossar y.
11
The Easter n and Wester n Mediter r anean display im-
por tant geological and biological differ ences. Fir st, the
Easter n Mediter r anean is geologically mor e active, be-
cause it is a contact zone between 3 major tectonic
plates: Afr ican, Eur asian and Ar abian. For this r eason, it
featur es a wide ar r ay of inter esting cases of unique bio-
cenoses*, such as mud volcanoes, seamounts or cold
seeps*. The Wester n Mediter r anean is r elatively featur e-
less in compar ison, although still not devoid of unique
envir onments. Biologically, the Wester n Mediter r anean,
whilst still oligotr ophic* by Nor th Atlantic standar ds, has
r elatively high pr imar y pr oduction, especially in the Gulf
of Lions, due to the r iver Rhone r unoff and wind mixing
( fig. 2) . The Easter n Mediter r anean has ver y low pr ima-
r y pr oduction, par ticular ly in the Levantine sea.
Fi g. 2 . Chlor ophyll amap ( Monthly composite for Apr . 2000) pr o-
duced by the Inland and Mar ine Water s Unit ( JRC-EC) using SeaWiFS
r aw data distr ibuted by NASA-GSFC.
Se e co l o ur pl ate , p. 5 7 .
The continental shelves, wher e most commer cial fish ing
activity is conducted, cover only 30% of the Mediter r a-
nean sea sur face, while the bathyal domain cover s 60%
and the abyssal plain the r emaining 10% ( appr ox.) ( fig.
3) . Continental shelves ar e only extensive near the ma-
jor r iver mouths ( the Rhone in the Gulf of Lions, the Nile
in the Levantine sea) , or on the Adr iatic and Tunisian
coasts. A lar ge par t of the Mediter r anean coast has deep-
water beds ver y near shor e, typically a few hour s away
by commer cial vessel.
Although fishing for continental shelf r esour ces dates
back to ancient times, commer cial fisher ies in the bath-
yal domain star ted only in the ear ly decades of the 20
th

centur y, and have incr eased in impor tance since the
1950s, especially on the Ligur ian, Catalan and Balear ic
coasts ( Sar d et al., 2004a) . The development of deep-
water commer cial fisher ies was linked, as in other seas,
to the development and motor ization of tr awler vessels.
Deep-water fisher ies in the Mediter r anean tar get r ed
shr imps ( Aristeus antennatus and Aristaeomorpha
foliacea) between 400 and 800 m ( spor adically down
to 1000 m.) Beds deeper than 1000 m can be consid-
er ed vir gin fr om the viewpoint of commer cial exploita-
tion of living r esour ces.
Thr oughout the wor ld, commer cial fisher ies, based
mostly on pr edator y fish, ar e suffer ing ser ious depletion
( Myer s and Wor m, 2003) . Given the state of continen-
tal shelf fish r esour ces, the fishing industr y is dir ecting
mor e attention towar ds the potentially exploitable living
r esour ces of the deep-sea. Water s deeper than 1000 m
cover ar ound 60 % of our planet, and their unique eco-
systems have been discover ed only r ecently ( since the
1970s) . Hydr other mal vents, cold seeps, and cold-wa-
ter cor al r eefs can be found in deep water s. Other habi-
tats, such as seamounts ( Roger s, 1994; Koslow, 1997;
Galil and Zibr owius, 1998) and submar ine canyons
( Bouillon et al., 2000; Gili et al., 1998; 2000) , have
been identified as diver sity hotspots. Consider ing their
unique featur es and low tur n-over r ates, deep-sea eco-
systems ar e highly vulner able to commer cial exploita-
tion ( Rober ts, 2002) . Globally, ar ound 40% of tr awling
fishing gr ounds ar e now on water s deeper than the con-
tinental shelf ( Rober ts, 2002) and many seamount fish-
er ies have been depleted in shor t per iods of time ( Lack
et al., 2003) .
Fi g. 3 . Schematic r epr esentation of the mar ine depth zones, with
emphasis on concepts pr esented in this document.
2. Deep-sea diversity patterns
2.1. Fauna
2 . 1 . 1 . O ve rvie w
The deep Mediter r anean fauna displays a number of char -
acter istics that differ entiate it fr om other deep-sea faunas
of the wor lds oceans ( Bouchet and Taviani, 1992) : i)
the high degr ee of eur ybathic* species; ii) absence ( or
low r epr esentation) of typical deep-water gr oups, such
as macr oscopic for aminifer a ( Xenophyophor a) , glass
12
sponges ( Hexactinellida) , sea-cucumber s of the or der
Elasipodida, pr imitive stalked sea-lilies ( Cr inoidea) and
tunicates ( sea-squir ts) of the class Sor ber acea ( Pr s,
1985; Monniot and Monniot, 1990) ; and iii) the number
of endemisms ( 26.6% of species in the Mediter r anean
fauna: Ruffo, 1998) declines with incr easing depth,
with compar atively low endemisms below 500 m ( see
also Fr edj and Laubier , 1985) . The number of ende-
misms depends, however , on the taxon and, for instance,
among Amphipoda a high per centage ( 49.2%) of the
bathyal species ar e endemic ( Ruffo, 1998) , even higher
than in coastal zones. In the case of amphipods, the ab-
sence of pelagic fr ee lar vae ( as compar ed to other pe-
r acar id taxa) helps explain this high r ate of deep-sea
endemisms.
The Mediter r anean deep-sea is ver y young com-
par ed to other oceans. Dur ing the Messinian ( Upper
Miocene) the water flow between the Atlantic and the
Mediter r anean was cut off, as a r esult of tectonic move-
ments of the Eur opean and Afr ican plates, pr ecipitating
an almost complete dr ying out of the Mediter r anean
between 5.7 and 5.4 million year s ago ( Messinian sa-
linity cr isis) . The water exchange between the two seas
was r estor ed in the Lower Pliocene, giving r ise to a
Mediter r anean sea fauna r emar kably similar to that of
the Atlantic in the Pliocene and Pleistocene. The cur -
r ent Mediter r anean deep-water fauna is less r elated to
the Atlantic bathyal fauna than it was in the Pleistocene
( Bar r ier et al., 1989) , due to the lack of Atlantic deep
water ( and fauna) enter ing the Mediter r anean ( Salas,
1996) . Bouchet and Taviani ( 1992) postulated that the
impover ishment of the Mediter r anean deep-sea fauna is
r elatively r ecent ( post-glacial Pleistocene) , and that the
onset of cur r ent hydr ological conditions in the Holocene
led to an almost complete exter mination of the r icher
glacial deep-sea fauna, which was mor e similar to the
pr esent Atlantic fauna, par ticular ly affecting the cold
stenohaline* taxa.
Some faunistic gr oups ar e poor ly r epr esented in the
deep Mediter r anean compar ed to the NE Atlantic: fish
( Stefanescu et al., 1992) , decapod cr ustaceans ( Car tes,
1993) , mysids ( Car tes and Sor be, 1995) , echinoder ms
( Tor tonese, 1985) , and gastr opods ( Bouchet and Taviani,
1992) . Fr edj and Laubier ( 1985) r epor ted that ca.2100
species of benthic* macr ofauna* ar e found deeper than
200 m in the Mediter r anean, and only 200 species have
been r ecor ded deeper than 2000 m, although the lower
sampling effor t below 2000 m could cer tainly bias this
r esult. Bouchet and Taviani ( 1992) pointed to the pau-
city of deep Mediter r anean benthic macr ofauna in ter ms
of abundance, species and endemisms.
Systematic deep-sea samplings have shown that the deep
Mediter r anean basin ( along with the Nor wegian and
Car ibbean basins) har bour s impover ished deep Atlantic
fish faunas ( Haedr ich and Mer r ett, 1988) .
Cer tain ar eas of the Mediter r anean deep-sea ar e benthic
diver sity hotspots because they har bour special faunas
r ich in endemic taxa ( see section on unique habitats
below) . Submar ine canyons ar e also consider ed as
hotspots of species diver sity and endemisms ( Gili et al.,
1998; 2000) .
Additionally, a flor o-faunistic impover ishment of the
Easter n Mediter r anean compar ed with the Wester n
Mediter r anean r ichness in species is accepted ( Sar ,
1985) , thr ough a gr adational decr ease fr om west to east,
which is mor e conspicuous for the deep benthos. The
Easter n basin is separ ated fr om the Wester n one by the
Siculo-Tunisian sill ( ca. 400 m) , and the Levantine bath-
yal benthos appear s to be composed of autochthonous
self-sustaining populations of eur ybathic species that
settled after the Messinian event.
2 . 1 . 2 . M e g a fa u n a *
Scientific knowledge of deep megafaunal communities
( mainly fish, cr ustaceans and cephalopods) was limited
to the bathymetr ic r ange exploited by fishing ( down to
800-1000-m) until the ear ly 1980s, when scientific
expeditions began quantitatively sampling the bathyal
gr ounds in the Mediter r anean.
Based on extensive samplings in the Levantine sea,
Klausewitz ( 1989) , Galil and Gor en ( 1994) and Galil
and Zibr owius ( 1998) , r epor ted the scar city of deep-sea
fish fauna in the Mediter r anean easter n basin, but at the
same time r evealed the discover y of new and r ar e taxa.
Dur ing the 1980s and 1990s, a ser ies of sur veys wer e
conducted in the Catalano-Balear ic and Alger ian Basins,
focussing on mor e ecological aspects ( biomass and
size-depth distr ibution, feeding ecology) . These sam-
plings wer e car r ied out in lower -slope bathyal commu-
nities between 1000 and 2300 m in depth, paying spe-
cial attention to the dominant megafaunal gr oups, fish
( Stefanescu et al., 1992, 1993; Mor anta et al., 1998)
and decapod cr ustaceans ( Car tes and Sar d, 1992,
1993; Car tes, 1994, 1998b; Maynou and Car tes, 2000;
Car tes and Car r assn, 2004) .
Standar dised compar isons point to clear differ ences
between Mediter r anean and Atlantic deep-sea demer -
13
sal fish fauna ( Mor anta et al., 2003) . Fish population
densities ar e lower in the Mediter r anean ( fig. 4) , a fea-
tur e that can be r elated to the much lower availability of
or ganic matter * on the Mediter r anean seabed in com-
par ison to the Atlantic, due to the oligotr ophic char ac-
ter istics of the Mediter r anean sea ( fig. 2) . In the same
way, Atlantic fish assemblages ar e composed of a lar g-
er number of species than those of the Mediter r anean.
This might owe to the r ecent or igin of Mediter r anean
deep-sea fauna, after the Messinian salinity event.
Compar ing the Easter n and Wester n Mediter r anean ba-
sins, the Levantine Sea has a par ticular ly low number
of species and low abundance ( Stefanescu et al., 1992;
Kallianotis et al., 2000) , and the assemblage composi-
tion is also qualitatively differ ent. For instance, deep-sea
shar ks ( Centrophorus spp., Etmopterus spinax and
Hexanchus griseus) ar e among the most abundant fish
species in Levantine bathyal communities.
A char acter istic featur e of Mediter r anean deep-sea
megafauna is the numer ical impor tance ( in ter ms of
abundance and number of species) of decapod cr us-
taceans which, together with fish, ar e the dominant
taxa in deep Mediter r anean assemblages. In the deep
Mediter r anean, decapod cr ustaceans ar e the dominant
inver tebr ate taxon, whilst they ar e of little impor tance
in biomass in the Bay of Biscay. Additionally, species of
tr opical or subtr opical or igin ( e.g. Aristeus antenna-
tus, Aristaeomorpha foliacea, Acanthephyra eximia)
dominate in deep Mediter r anean decapod assemblages
( Car tes, 1993) .
Suspension feeder s* ( e.g. hexactinellid sponges, pennat-
ulids) ar e dominant in ter ms of inver tebr ate biomass on
the upper and middle slope ( to 1400 m) in the Atlantic,
whilst echinoder ms ar e impor tant at all depths and
dominant on the middle and lower slope ( Lampitt et al.,
1986) . Suspension feeder s ar e of r elatively little impor -
tance in the Mediter r anean, due to its ologotr ophic wa-
ter s, and they ar e impor tant only locally. Pr obably due
to their low levels of food consumption, cr ustacean de-
capods ar e mor e impor tant in the deep Mediter r anean
( an oligotr ophic r egion, Car tes and Sar d, 1992) than in
the deep Atlantic, wher e echinoder ms dominate ( Billett
et al., 2001) .
Me gafauna abundance and s pe ci e s ri chne s s
de cre as e s wi th de pth, but the re e xi s ts a s e c-
o ndary pe ak o f bi o mas s be twe e n 1 0 0 0 and
1 5 0 0 m.
Do mi nant me gafaunal gro ups i n the de e p
Me di te rrane an are fi s he s and de capo d crus ta-
ce ans
The l e ve l s o f me gafaunal bi o mas s i n the de e p
Me di te rrane an are 1 o rde r o f magni tude l o we r
than i n the Atl anti c, at co mparabl e de pths , due
to the l o w pri mary pro ducti o n o f the Me di te r-
rane an s e a.
Impo rtant di ffe re nce s e xi s t be twe e n the e as t-
e rn and the we s te rn Me di te rrane an, bo th i n
s pe ci e s co mpo s i ti o n and abundance
Fi g. 4 . Compar ison of biomass by depth inter val between the
Wester n Mediter r anean ( Balear ic sea) and the Atlantic ( Rockall
tr ough) , based on Mor anta et al. ( 2003) .
Etmopterus spinax.
Dr awing by M. Wr tz Ar tescienza 1994.
b|omass (kg km
-2
)
14OO-17OO m
8OO-14OO m
4OO-8OO m
western Nediterranean Atlantic
O 1OOO 2OOO 8OOO 4OOO 5OOO
14
Box 1. Species composition of megafauna in the Western Mediterranean
Relative abundance of decapod cr ustaceans with depth,
elabor ated fr om data fr om exper imental samplings in the wester n
Mediter r anean fr om 1988-2001.
Otter tr awl sur veys in the Wester n Mediter r anean
car r ied out between 1988 and 2001 ( Stefanescu et al.,
1994; Car tes, 1993; Sar d et al., 2004b) pr ovide an
accur ate pictur e of the decapod cr ustacean and sh
species composition and biomass distr ibution with
depth.
Decapod crustaceans
Seventy species of decapod cr ustaceans have been r e-
por ted fr om 200 to 4000 m. Fr om these, 20 ar e of
commer cial inter est at pr esent ( including some spe-
cies dwelling pr edominantly in shallow water s) :
- the r ed shr imps: mainly Aristeus antennatus,
but locally also Aristaeomorpha foliacea.
- other r ed shr imps: Acanthephyraeximia
and Acanthephyra pelagica.
- the glass shr imps: Pasiphaeasivado
and Pasiphaea multidentata.
- Pandalid shr imps Plesionikaspp.
( down to 1000 m)
Among these species, only 4 r each signicant densities
( > 0.1 kg/ h in the exper imental sampling gear ) be-
tween 1000 and 1600 m depth: Aristeus antennatus,
Geryon longipes, Acanthephyra eximiaand Munida
tenuimana. Below 1600 m the total biomass of deca-
pod cr ustaceans is ver y low ( < 1 kg/ h) .
The distr ibution of biomass with depth of decapod
cr ustaceans is shown in the gur e below. Ther e is an
impor tant incr ease in biomass fr om 200 to 400 m, fol-
lowed by a shar p decr ease to 1000 m, wher e the deca-
pod biomass is pr actically one tenth of the biomass at
the shelf br eak. A secondar y peak of biomass occur s
between 1300 and 1500 m, though r eaching much
lower levels of biomass. The high abundance of Aris-
teus antennatusat ar ound 1500 m is par tly r esponsi-
ble for this secondar y peak ( Sar d et al., 2003b) .
- the deep water shr imp Parapenaeus
longirostrisand Solenocera membranacea
( down to 400 m only)
- the anomur an cr abs Munidaspp.
- the Nor way lobster Nephrops norvegicus
( pr esent down to 700 m only) and the spiny
lobster Palinurus mauritanicus
( 400 m only)
- cr abs: Geryon longipes, Paromola cuvieri,
Chaceon mediterraneus, and Macropipus
tuberculatus( the latter down to 600 m only) .
Paromola cuvieri.
Dr awing by M. Wr tz
Ar tescienza 1977.
O
2
4
O
8
1O
2
O
O
4
O
O
O
O
O
8
O
O
1
O
O
O
1
2
O
O
1
4
O
O
1
O
O
O
1
8
O
O
2
O
O
O
2
2
O
O
dep h (m)
k
g
l
h
15
Fishes
The sh assemblage fr om 200 to 2250 m ( Stefanescu
et al., 1993; 1994, Mor anta et al., 1998) compr ises
ca.90 species in the wester n Mediter r anean. As shown
in the gur e below, sh biomass str ongly decr eases
fr om 200 m to 800 m ( i.e. ~ 10:1) . Fishes ar e ver y
diver se at these depths, and the commer cial species
ar e r epr esented by Merluccius merlucciusand Phy-
cis blennoides, among other s. Fr om 800 m to 1800 m,
impor tant sh biomasses ar e again pr esent. The domi-
nant species at 800-1100 m ar e the shar ks Dalatias li-
chaand Galeus melastomus, and the teleostean sh-
es Alepocephalus rostratus and Mora moro. The lat-
ter ar e dominant fr om 900 to 1100 m. Fr om 900 m
A. rostratus becomes ver y impor tant, whilst between
1300 and 1700 m it for ms mor e than 70% of the sh
biomass ( see also table below) . It is impor tant to note
that while standing biomass stocks ar e r elatively im-
por tant between 800 and 1500 m, the pr oductivity of
sh at these depths is expected to be ver y low ( see
Chapter 3) .
Stefanescu et al. ( 1992) showed that in the NW Medi-
ter r anean, sh size ( mean sh weight) decr eases for
some dominant species with depth between 1000
and 2250 m. This tr end is especially signicant below
1200 m, due to a faunal shift wher eby lar ge or medi-
um-sized species ( gadifor ms such as M. moro, P. blen-
noides, T. trachyrhynchus or shar ks such as Hexan-
chus griseus) ar e r eplaced by small ones ( such as the
macr our ids Coryphaenoides guentheri and Chalinu-
ra mediterraneaor the chlor ophthalmid Bathypter-
ois mediterraneus) . The author s pointed to the im-
possibility of lar ge or medium-sized shes to satisfy
their ener getic r equir ements in a pr ogr essively mor e
r estr ictive tr ophic envir onment. Similar r esults have
been obtained for the Easter n Mediter r anean down to
4000 m ( DOnghia et al., 2004) .
D e p th in te rva l
( m )
B io m a ss d o m in a n c e
( 3 0 % o f to ta l fish we ig h t)
A b u n d a n c e
( > 3 0 % o f to ta l fish n u m b e r
<1000
1000-1200
1200-1500
1500-2500
>2500
G. melastomus
A. rostratus
A. rostratus
C. clolepis
Ch. mediterranea
T. scabrus
A. rostratus
B. mediterraneus
B. mediterraneus
Ch. mediterranea
O
1O
2O
8O
4O
5O
OO
7O
8O
9O
2OO 8OO 4OO 5OO OOO 7OO 8OO 9OO 1OOO 11OO 12OO 18OO 14OO 15OO 1OOO 17OO 19OO
dep h (m)
k
g
l
h
Nora moro Alepocephal s rostrat s 0ale s melastom s
0ther fishes Nerl cci s merl cci s Dalatias licha Phcis olennoides
Centroscmn s coelolepis lepidion lepidion
Alepocephalus rostratus Risso, 1820
Vincent Fossat, 1879.
Relative abundance
of shes with depth,
extr apolated fr om data
fr om exper imental
samplings in the wester n
Mediter r anean fr om
1988-2001.
16
2 . 1 . 3 . M a c ro fa u n a
In the deep Mediter r anean, macr ofaunal biomass
var ies consider ably acr oss ar eas ( Pr s, 1985) and
also seasonally. Macr ofaunal abundance and biomass
decr ease gener ally with depth, and most likely also,
as in meiofauna, along a west-east gr adient. Close
to submar ine canyons or other ar eas of local high
pr oductivity, macr ofaunal biomass may incr ease.
Macr ofaunal density in the Mediter r anean is about 1/ 10
of the densities r epor ted for the Atlantic at compar able
depths ( Cosson et al., 1997; Flach and Heip, 1996) .
In the Wester n Mediter r anean, Stor a et al., ( 1999) , r e-
por ted values between 2.54 g/ m
2
dr y weight ( at 250 m)
and 0.05 g/ m
2
( at 2000 m) , in the Toulon canyon.
Macr ofaunal biomass differ ed also between the canyon
axis ( 2.54 g/ m
2
) and the canyon flanks ( 0.70 g/ m
2
) at
the same depth. Richness of species decr eased also with
depth, fr om 124 species at 250 m to 31 at 2000 m. A
combination of gr ain size and geochemical composition
of the sediments was suggested as possible explanator y
causes of the patter ns obser ved. Deposit feeder s wer e
the dominant guild in the Toulon canyon instead of sus-
pension feeder s and car nivor es, which wer e dominant
in the upper and middle slope in oceanic r egions.
In the mor e oligotr ophic Cr etan sea, Tselepides et al.
( 2000) r epor ted compar able values of macr ofaunal bio-
mass: 1 g/ m
2
dr y weight at 200 m depth and 0.06 g/ m
2
at
1570 m. Instead, species r ichness was consider ably low-
er in the Cr etan sea than in the NW Mediter r anean, with
ar ound 35 species at 200 m and ar ound 8 at 1570 m.
The small aver age size of species ( most animals between
0.5 and 10 mm) seems to be a char acter istic of the deep
Mediter r anean macr ofauna. As a consequence, the deep
Mediter r anean featur es a mor e mar ked decr ease in the
biomass than in the density of macr obenthos with depth
( Pr s, 1985) .
The diver sity and distr ibutional patter ns of the swim-
ming macr ofauna ( supr abenthos* or hyper benthos*)
have been studied in the wester n ( Car tes and Sor be,
1996, 1999; Car tes et al. 2003) , and, to a lesser extent,
the easter n Basin ( Madur ell and Car tes, 2003) . Differ -
ent taxa of bathyal supr abenthic fauna showed diver si-
ty peaks at mid-bathyal depths in the Catalan sea and in
the Alger ian Basin, with some var iation depending on
the taxon consider ed ( ar ound 600 m for amphipods;
ar ound 1200 m for cumaceans) . Compar isons of su-
pr abenthic assemblages in ter ms of number of species
between the Ionian sea ( easter n Basin) and the Catalan
sea ( wester n Basin) suggest a lower impover ishment in
the Easter n Mediter r anean per acar id assemblages than
would be expected ( Madur ell and Car tes, 2003) .
Regar ding near -bottom zooplankton, Scotto di Car lo
et al. ( 1991) r epor ted the existence of homogenous
copepod assemblages fr om the deep Mediter r anean
( 600-2500 m) , with no differ ences in faunal composi-
tion, and higher copepod biomass in the Wester n ba-
sin than in the Easter n basin. Due to the r elatively war m
temper atur e ( 13-14.5C) of deep Mediter r anean water s,
one might expect low biomass levels of near -bottom zo-
oplankton. For instance, the biomass of near -bottom
deep-sea zooplankton ( 10 m above the sea bed) was
anomalously lower in the deep Red Sea ( wher e the tem-
per atur e of deep water r eaches 22 C) than in the typical
oceanic r egions ( Atlantic and Pacific: Wishner , 1980) , a
tr end attr ibutable to incr eased decomposition r ates of
or ganic matter in a war m water mass.
Macro faunal abundance and s pe ci e s ri chne s s
de cre as e s ge ne ral l y wi th de pth, but l o cal e n-
vi ro nme nts wi th i ncre as e d l e ve l s o f bi o l o gi cal
pro ducti o n may i ncre as e macro faunal abun-
dance and di ve rs i ty.
Macro fauna s pe ci e s are typi cal l y ve ry s mal l i n
the Me di te rrane an, co mpare d to the Atl anti c.
Bathymedon longirostris, Eusirus longipes, Lepechinella manco, Rachotropis grimaldii.
J. E. Car tes. Se e co l o ur pl ate , p. 6 0 .
17
2 . 1 . 4 . M e io fa u n a
While densities of meiofauna* ( or meiobenthos) on
the Mediter r anean continental shelf ar e compar able to
those in other oceans ( Danovar o et al., 2000) , deep-sea
meiobenthos densities ar e one or der of magnitude low-
er in the Mediter r anean than in the nor theast Atlantic
( Bouchet and Taviani, 1992; Danovar o et al., 2000; and
especially below 1700 m: De Bove et al., 1990) .
Meiofaunal abundance ( excluding for aminifer ans) di-
minishes with incr easing depth ( De Bove et al., 1990) ,
along with envir onmental indicator s of food r esour ces.
This patter n is even mor e pr onounced in the Easter n ba-
sin ( Danovar o et al., 2000) ( fig.5) and the extr emely ol-
ogotr ophic easter n Mediter r anean shows one of the low-
est meiofaunal standing stocks in the wor ld ( Tselepides
and Lampadar iou, 2004) .
al., 1999; 2000) ; in the Wester n Mediter r anean it is low-
er ( 2.5 times) . The combination of low pr imar y pr oduc-
tion and bacter ial dominance of secondar y pr oduction
in the east is also of significance as it could account for
the low fisher ies pr oduction ( Tur ley et al., 2000) .
The r apidly diminishing meiofaunal abundance is ex-
plained not only by the oligotr ophic natur e of the Medi-
ter r anean, but also by the r apidity of or ganic matter deg-
r adation in a r elatively war m deep-water envir onment
( De Bove et al., 1990; Bouchet and Taviani, 1992) .
Infor mation on meiobenthos fr om tr enches of the abyss-
al or hadal zones is still limited, but Tselepides and Lam-
padar iou ( 2004) showed that meiofaunal abundance in
Easter n Mediter r anean deep-sea tr enches ( with depths
exceeding 3750 m) was unexpectedly high ( 45-156
ind/ 10 cm
2
) ; higher than in sur r ounding abyssal plains
and compar able to mid-slope ( 1500 m) depths. The ac-
cumulation of or ganic matter in deep sea tr enches could
explain the high meiofaunal abundance r epor ted.
The abundance o f me i o fauna di mi ni s he s wi th
i ncre as i ng de pth, and i s l o we r i n the e as te rn
Me di te rrane an than i n the we s te rn
Lo cal hi gh abundance o f me i o fauna has be e n
re po rte d fo r ve ry de e p e nvi ro nme nts , s uch as
de e p- s e a tre nche s
The co mbi nati o n o f l o w pri mary pro ducti o n
and bacte ri al do mi nance o f s e co ndary pro duc-
ti o n may acco unt fo r the l o w fi s he ri e s pro duc-
ti o n i n the e as te rn Me di te rrane an
2 . 1 . 5 . E n d e m ism
By its special oceanogr aphic str uctur e and paleoecolo-
gy, the Mediter r anean Sea has a par ticular fauna com-
par ed to the near by Atlantic ocean. Differ ent theor ies
about the or igin and evolution of deep Mediter r anean
fauna have been suggested and documented. Examples
ar e the Tethys hypothesis, which postulates that the or i-
gin of some Mediter r anean endemisms ar e faunal r elicts
sur viving the Messinian cr isis ( Pr s, 1985) , and the
pseudopopulation hypothesis by Bouchet and Taviani
( 1992) , explaining the occur r ence of some species in
the Mediter r anean not as r epr oducing populations, but
as maintained by the per iodical influx of lar vae tr ough
In gener al, meiofaunal abundance is dominated by nem-
atodes, with impor tant fr actions of har pacticoid cope-
pods and polychaetes. In the Wester n Mediter r anean
( Gulf of Lions and Catalan Sea) , the meiofauna is domi-
nated by nematodes ( 92.4%) . In the Catalan Sea meio-
faunal density var ied seasonally fr om 296 ( Septem-
ber ) to 746 ( Mar ch) ind cm-2 ( Car tes et al., 2002)
at ar ound 1200 m depth. Nematode biodiver sity below
500 m in the Easter n Mediter r anean is lower than in
other equally deep sediments wor ldwide, and even low-
er than within a Mediter r anean canyon at 1500 m depth
( Danovar o et al., 1999) .
Below 500 m depth in the Easter n Mediter r anean, the
contr ibution of bacter ia to or ganic matter degr adation
is significant ( bacter ia r epr esent 35.8% of the living bio-
mass, Danovar o et al., 1995) and the bacter ial to meio-
faunal biomass r atio is ver y high ( 20 times, Danovar o et
O
2OO
4OO
OOO
8OO
1OOO
12OO
14OO
1OOO
18OO
O 5OO 1OOO 15OO 2OOO 25OO
dep h (m)
western Nediterranean Eastern Nediterranean
Fi g. 5 . Meiofaunal abundance by depth, combining data
fr om De Bove et al. ( 1990) , Danovar o et al. ( 1999; 2000) .
18
Lepidion lepidion( Risso, 1810)
Lota lepidionRisso,
Moustella deFount Stocoe,
V. Fossat, 1879.
Coll. Musum dHistoir e natur elle
de Nice.
the Str aits of Gibr altar . Most components of the Mediter -
r anean deep-water fauna ( deeper than ca. 200 m) ba-
sically der ive fr om impover ished communities of Atlan-
tic or igin.
It has been estimated that ar ound 26.6% of the total
Mediter r anean mar ine fauna ( 4238 species: Fr edj et al.,
1992) ar e endemic. Though no similar over all balanc-
es exist specifically for the deep-sea fauna, the per cent-
age of endemisms is higher than the mean above cited
for some taxa ( e.g., 49.2% Amphipoda, Ruffo, 1998) .
For some taxa, even endemic gener a can be found ( e.g.,
in decapod cr ustaceans: Levantocaris, Zariquieyon) ,
while in other taxa with a higher number of endemic
species ( amphipods) , no endemic gener a ar e r epor ted.
As explained above for Amphipoda, the pr opor tion of
endemisms by taxon is pr obably closely r elated with life
str ategies. For example, no endemic cephalopods ar e
r epor ted in the deep Mediter r anean ( R. Villanueva, per s.
comm.) , pr obably due to the possession of fr ee-pelagic
lar vae distr ibuted in the entir e water column.
Tor tonese ( 1985) r epor ted 6 endemic fishes below
1000 m: Bathypterois mediterraneus, Paralepis speci-
osa, Rhynchogadus hepaticus, Eretmophorus kleinen-
bergi, Lepidion lepidionand Paraliparis leptochirus.
Only two of these species ar e found both in the easter n
and wester n Mediter r anean.
Among decapod cr ustaceans, Levantocaris hornun-
gae ( Galil and Clar k, 1993) , and the lar ge cr abs Cha-
ceon mediterraneus ( Manning and Holthuis, 1989;
Car tes, 1993) , and Zariquieyon inflatus( Manning and
Holthuis, 1989) ar e endemic. Only 2 of the 28 species
of decapods collected in the Catalan Sea between 862-
2265 m wer e endemic ( Car tes, 1993) . 2 of the 3 endem-
ic decapods ar e found both in the easter n and wester n
Basins, while Z. inflatushas been collected only in the
wester n Mediter r anean.
Tor tonese ( 1985) r epor ted 4 endemic echinoder ms be-
low 200 m, 1 dominant ( Leptometra phalangium) and
3 r ar e species ( Prototrochus meridionalis, Irpa lud-
wigi, Hedningia mediterranea) .
In the case of macr ofauna, although the depth distr ibu-
tion of endemisms is mor e or less acknowledged, it is
har dly known if these endemisms ar e unifor mly distr ib-
uted along all deep Mediter r anean Basins. In the case
of per acar id cr ustaceans, between 552 and 1808 m in
the Catalano-Balear ic Basin, Car tes and Sor be ( 1995)
r epor ted 3 endemic Mysidacea of the total of 16 spe-
cies collected, 4 of the 32 species of Cumacea ( Car tes
and Sor be, 1996) , and 26 of the 82 species of Amphipo-
da ( Car tes and Sor be, 1999) . Ther efor e, 25.4% of bath-
yal per acar ids in the Catalano-Balear ic Basin can be
consider ed as endemics. Pr eviously, a similar pr opor -
19
tion ( 24.2% of the 99 species collected) of bathyal-su-
pr abenthic Per acar ids wer e cited as Mediter r anean en-
demics in the same ar ea ( r ecalculated fr om Car tes and
Sor be, 1993) at similar depths ( fr om 389 to 1859 m) .
In the Alger ian Basin, at depths between 249 and
1622 m, the per centage of endemisms among supr aben-
thic Per acar ids was slightly lower than in the Catalan Sea
( 18.2%: calculated fr om Car tes et al., 2003) .
The number of endemisms is low, par ticular ly in the deep
bathyal domain and below 2500-3000 m. This boundar y
has been suggested by some author s ( Pr s, 1985; Bel-
lan-Santini, 1990) as the upper limit of distr ibution of
the tr ue abyssal fauna. However , the number of Medi-
ter r anean deep-sea endemisms could have been over -
estimated simply by differ ences in sampling between the
deep Mediter r anean and sur r ounding biogeogr aphic ar -
eas; some deep amphipods ( e.g. Rhachotropis caeca,
Lepechinella manco, or Pseudotiron bouvieri) , cata-
logued as endemic Mediter r anean species, have in r e-
cent year s been found in the Bay of Biscay and the Can-
tabr ic Sea ( Bachelet et al., 2003) .
Conver sely, new sampling pr ogr ammes ar e incr easing-
ly r epor ting new endemic taxa in the deep Mediter r ane-
an, and the faunal composition of deep-sea communi-
ties is far fr om being completely r ecognized , especially
for small faunal gr oups.
The numbe r and type o f e nde mi s ms i n the
de e p Me di te rrane an are the re s ul t o f i ts par-
ti cul ar pal e o e co l o gy. The y s ho ul d be an i mpo r-
tant cri te ri o n i n de fi ni ng future MPAs i n de e p
Me di te rrane an e co s ys te ms .
So me fauni s ti c gro ups s uch as amphi po ds
pre s e nt hi gh rate s o f e nde mi s m i n the de e p
Me di te rrane an.
The numbe r o f e nde mi c s pe ci e s amo ng me g-
afaunal taxa i n the de e p Me di te rrane an are 6
fo r fi s he s , 4 fo r de capo d crus tace ans and 4 fo r
e chi no de rms .
2.2. Structure of deep-water
Mediterranean communities
2 . 2 . 1 . C o n tin e n ta l slo p e
Two main biocenoses ( facies) in the bathyal Medi-
ter r anean have classically been defined ( Pr s, 1985) :
i) soft-bottom communities; ii) har d-bottom commu-
nities. Pr s and Picar d ( 1964) established the tr ansi-
tion between the cir calittor al and bathyal domains in the
Mediter r anean at ar ound 180-200 m depth. Depending
on the natur e of the sediment ( which depends in tur n
on factor s such as hydr odynamism and mainland influ-
ence) , thr ee hor izons wer e established by Pr s ( 1985)
in the soft-bottom communities: the upper , middle and
lower slope hor izons. The upper slope hor izon is a tr an-
sition zone between the coastal zone and bathyal do-
mains, compr ising a lar ge shar e of eur ybathic for ms and
extending to 400-500 m deep. The sea pen Funiculi-
na quadrangularisand the cr ustaceans Parapenaeus
longirostris and Nephrops norvegicus ar e char acter -
istic species of this hor izon. The middle slope hor izon,
char acter ized by fir mer and mor e compact muds, is the
zone wher e most taxa achieve maximum diver sity. For
fish and decapods, in the NW Mediter r anean, this ho-
r izon extends to a depth of 1200-1400 m. The cnidar -
ian Isidella elongata, the cr ustaceans Aristeus anten-
natusand Aristaeomorpha foliaceaar e char acter istic
examples. The lower slope hor izon is not so well stud-
ied, but some char acter istic megafaunal species can be
listed: decapods, such as Stereomastis sculpta, Acan-
thephyra eximiaand Nematocarcinus exilis, and fish-
es, such as Bathypterois mediterraneus, Alepochepha-
lus rostratus, Lepidion lepidionand Coryphaenoides
guentheri. Har d bottoms ar e r epr esented below 300 m
and down to 800-1000 m, in ar eas wher e high hydr o-
dynamism pr ecludes sedimentation and exposes bar e
r ock. In the nor ther n Ionian sea ( Santa Mar ia di Leuca)
a unique biocenose of white cor als has been r epor ted
fr om 450-1100 m depth ( Mastr ototar o et al. 2002; Tur -
si et al. 2004) . Live colonies of Madrepora oculataand
Lophelia pertusawer e found, with a r ich epibiont fauna.
These species occur elsewher e in the Mediter r anean as
fossils or subfossils ( Pr s, 1985) , s e e Bo x 6 .
Aristeus antennatus
Alber t I
er
Pr ince de Monaco. Camp. scient. Pnids pl. III.
EL. Bouvier del, M. Bor r el pinx.
Coll. Oceanogr aphic Museum, Monaco.
20
In the mar ine envir onment, depth is assumed as the ma-
jor for cing factor str uctur ing communities: species as-
semblages and communities show mor e var iability ver -
tically, as a function of incr easing depth, than over the
hor izontal dimension ( Rowe and Menzies, 1969; Hae-
dr ich et al., 1975; 1980; Hecker , 1990a; Car tes and
Sar d, 1993; Stefanescu et al., 1994) . This obser vation
suggested the idea of the zonation phenomenon: depth
bands of high faunal homogeneity separ ated by bound-
ar ies of faunal r enewal. Species ar e usually distr ibuted
along ecological gr adients in a bell-shaped cur ve, and
this is also obser ved with diver sity. Ecological theor y and
obser vations show that species ar e distr ibuted along en-
vir onmental gr adients ( see e.g. ter Br aak and Pr entice,
1988) , having a differ ent habitat amplitudeor centre
of gravityin their distr ibution ( Stefanescu et al., 1994,
for a Mediter r anean deep-sea example) . Along a depth
gr adient, species show also a minimum and maximum
depth of occur r ence, with an inter mediate depth, the
Depth of Optimal Abundance ( DOA) , wher e their densi-
ty is the highest. Species may find better living conditions,
i.e. tr ophic r equir ements, at this DOA, as a consequence
of an adaptive success to the envir onment. In deep-Med-
iter r anean fish assemblages, for instance, most fish spe-
cies showed an incr ease in their tr ophic diver sity* asso-
ciated with the DOA ( Car r assn and Car tes, 2002) .
Boundar ies of faunal change have been descr ibed in
deep ecosystems, both at bathyal and abyssal depths.
Most deep-Mediter r anean species have an eur ybath-
ic distr ibution. This is pr obably due to the high ther -
mal ( 13-13.5C in the wester n basin; 14-14.8 C in the
easter n basin) and saline ( 38.5-38.6 PSU) stability of
the water mass below 150 m in the deep Mediter r anean
( Hopkins, 1985) . Pr s ( 1985) alr eady pr oposed a pat-
ter n of change with depth for deep Mediter r anean fau-
na. Although with impor tant local var iations, a faunal
r enewal belt between the upper and middle par t of the
slope appear s r ecur r ently between 300 and 700 m for
megafaunal assemblages, both in oceanic r egions and in
the Mediter r anean, and in the deep Catalan Sea ( nor th-
wester n Mediter r anean) at ca. 500 m depth ( Abell et
al., 1988; Car tes et al., 1994) . A second, deeper , bound-
ar y between 1000 and 1400 m has also been r epor ted
in deep sea ecosystems ( Haedr ich et al., 1980; Wenner
and Boesch, 1979; Car tes and Sar d, 1993) and, in the
deep Mediter r anean, it separ ates the middle and low-
er slope assemblages ( Pr s, 1985; Car tes and Sar d,
1993) . Due to the par ticular paleoecolgical conditions
of Mediter r anean biota, and the lack of a ther mal gr a-
dient below 150 m, the boundar y between bathyal and
abyssal depths has not been clear ly identified. The abyss-
al communities in the deep Mediter r anean ar e extr eme-
ly impover ished in species, and boundar ies ar e masked
by the eur ybathic char acter of deep Mediter r anean spe-
cies; some author s even question the existence of a tr uly
abyssal fauna in the deep Mediter r anean ( Pr s, 1985) .
Each depth assemblage is char acter ized by its own levels
of biomass, pr oduction and faunal diver sity. In the deep-
bathyal Mediter r anean, a decr ease in species r ichness
below 1300 m has been r epor ted to occur in differ ent
taxa ( e.g. in amphipods, Car tes and Sor be, 1999) . Most
studies available ar e descr iptive in natur e, with little data
tr ying to r elate changes in faunal assemblages with en-
vir onmental var iables that could explain possible zona-
tion patter ns. Ther efor e, possible causes for species dis-
tr ibution ar e far fr om being fully established. Some dif-
fer ences in zonation patter ns have been found depend-
ing on the tr ophic level of each taxon. Zonation r ates
r egular ly incr ease with tr ophic level ( Rex, 1977; Car tes
and Car r assn, 2004) , and, in the wester n Mediter r a-
nean, lower r ates of zonation wer e found in per acar ids
( belonging to a low tr ophic level) compar ed to fish and
decapods. Tr ophic and biological causes may explain,
in a ther mally stable envir onment, patter ns of distr ibu-
tion and zonation of deep-sea Mediter r anean species.
The bathymetr ic location of boundar ies of faunal r enew-
al also shows local var iations, accompanied by changes
in the location of peaks of biomass with depth. Concer n-
ing faunal zonation, mid and low bathyal communities
( wher e the dominant taxa ar e fish, decapods and per -
acar ids) ar e distr ibuted deeper in mor e eutr ophic ar e-
as ( the continental side of the Catalan Sea) than in the
insular ar ea of the SW Balear ic Islands, a mor e oligo-
tr ophic zone ( Car tes et al., 2004) . The displacement of
the depth wher e boundar ies ar e located seems a conse-
quence of var iations in tr ophic factor s ( e.g. or ganic mat-
ter enr ichment by r iver dischar ges) . Thus, within the
same community, this tr end is most evident among spe-
cies situated at the lowest tr ophic levels species with
a higher sensitivity to changes in the quality of pr imar y
sour ces of food ( pr imar y pr oduction and detr itus) .
2 . 2 . 2 . S u b m a rin e c a n yo n s
A factor inducing local changes in zonation is the oc-
cur r ence of active submar ine canyons ( fig. 6) , which
pr esent a physical discontinuity of the continental slope.
Active submar ine canyons ar e impor tant pathways chan-
neling advective inputs of mainland or igin, such as r iv-
er outflow, mediated by nepheloid* layer s ( Dur r ieu de
Madr n et al., 1999; Puig et al., 2001) . Canyons have
a higher or ganic car bon content than sur r ounding ar e-
as ( Buscail and Ger main, 1997) , and pr efer ential tr ans-
21
0
2 0 0 0 m
Fi g. 6 . Topogr aphy of a submar ine canyon.
Credits: GRC Geocincies Marines (Universitat de Barcelona)
and RMR Institut deCincies del Mar (CSIC), modified.
Se e co l o ur pl ate s , p. 6 3 - 6 4 .
por t of mater ial within the canyons compar ed with ad-
jacent open slope ar eas has been documented ( Monaco
et al., 1999) . Local sedimentar y events of high intensi-
ty have been r epor ted on the continental slope, displac-
ing lar ge masses of par ticulate mater ial over consider -
able distances. These phenomena ar e the or igin of dis-
tur bance* at small time and space scales ( Cr assous et
al., 1991) . Concer ning the r esponse by or ganisms, high
macr o- and meiofaunal biomass has been r epor ted in
canyons ( Car tes, 1998a) , and they ar e also impor tant to
megafaunal commer cial fisher ies ( e.g. r ed shr imp fish-
er ies in the vicinity of wester n Mediter r anean subma-
r ine canyons: Sar d et al., 1994; Stefanescu et al., 1994;
pr esence of Aristeus antennatusat shallow depths in
the canyons of the Nile delta, B. Galil, per s. comm.) .
In the deep Mediter r anean, submar ine canyons ar e
impor tant for ecosystem str uctur e and functioning be-
cause:
1) Canyons can act as a sour ce and r eser voir of endemic
species. A number of highly specific populations of Hy-
dr omedusae, collected by sediment tr aps, have been r e-
cently descr ibed fr om submar ine canyons in the wester n
Mediter r anean ( Gili et al., 1998; 2000) , with differ ent
species found in each canyon, separ ated by less than
100 km.
2) canyons can act as a pathway for littor al species to
colonize the deep water , when they ar e car r ied down
by advective inputs ( colonization of bathyal depths by
species living in har bour s in the Toulon canyon: Stor a
et al., 1999; littor al species found below 1000 m, e.g.
Psammogammarus caecusin the Catalan Sea: Car tes
and Sor be, 1999) .
3) migr ator y micr onektonic* cr ustaceans ( hyper iids,
euphausiids) tend to accumulate in the canyon head
( Macquar t-Moulin and Patr iti, 1996) . A biomass accu-
mulation of supr abenthos ( Car tes, 1998a) and decapod
cr ustaceans ( Car tes et al., 1993) has also been r epor t-
ed, with mor e or less evident seasonal fluctuations.
4) Species inhabiting canyons or their vicinity ( e.g. the
r ed shr imp Aristeus antennatus, Car tes, 1994; Sar d et
al., 1994) show tempor al changes in abundance, pr ob-
ably as a consequence of the seasonally-var ying accumu-
lation of food r esour ces in canyons. Higher r ecr uitment
of macr o- and megafaunal species into or in the vicin-
ity of canyons has also been documented ( Sar d et al.,
1994; Stefanescu et al., 1994; Car tes and Sor be, 1999) .
5) By changing the vor ticity of under water cur r ents, can-
yons ar e r esponsible for local upwelling, r esulting in an
enr ichment of the water column and pelagic systems.
Impor tant concentr ations of small pelagic fish, ceta-
ceans and bir ds ar e obser ved in the epipelagic* layer
in water s near submar ine canyons ( Palomer a, 1992;
Abell et al., 2003) .
In decapod cr ustacean communities, however , species
composition was similar between canyons and sur -
r ounding ar eas, and only some secondar y species wer e
pr efer entially distr ibuted, or mor e abundant, in canyons
( e.g. Ligur ensiferus, Plesionika edwardsi: Car tes et
al., 1994) .
In conclusion, the biological differ ences between can-
yons and sur r ounding ar eas r esult fr om differ ences in
biomass, r ather than the establishment of distinct com-
munities or species ( excluding per haps the endemic jel-
lyfish species r epor ted by Gili et al., 1998; 2000) , pr ob-
ably as a consequence of the high hydr odynamism of
these type of envir onments.
Physeter catodon.
Dr awing by M. Wr tz.
Ar tescienza.
22
3. Functioning
of deep-sea Mediterranean
food webs
3.1. Overview
Ear ly investigations pointed to a high stability of bathyal
ecosystems, r egar ding both their str uctur e and dynam-
ics ( Gr assle, 1977) . Stability was also assumed at sea-
sonal and inter annual time scales. This view, however ,
has pr ogr essively changed in r ecent decades with the
incr easing evidence of aspects such as the seasonal in-
flux of phytodetr itus to deep sea envir onments ( Hecker ,
1990b) and the non-continuous r epr oduction or peaks
in the r ecr uitment r epor ted for deep sea species ( s e e
Bo x 2 ) . Changes have also been r epor ted on an inter -
annual scale ( e.g. over a per iod of 10 yr in Por cupine
Abyssal Plain: Billett et al., 2001 or in hydr other mal
vents: Lutz and Haymon, 1994) , and it has also been
suggested that climate change is influencing the quanti-
ty and quality of food r eaching deep-sea Mediter r anean
ecosystems ( Danovar o et al., 2001) .
Globally, the Mediter r anean sea is consider ed an oligo-
tr ophic r egion. Based on satellite imager y data ( SeaW-
iFS, fig. 2) , the mean annual sur face pr imar y pr oduc-
tion, as indicated by the Chlor ophyll-a pigment, r anges
between 1 and 2 mg / m
3
/ yr in the most pr oductive ar e-
as, such as the Gulf of Lions or the Albor an Sea, and 0.1-
0.2 mg/ m
3
/ yr in the SW Balear ic Islands, 0.05 mg/ m
3
/ yr
in the Tyr r henian sea, and only 0.02-0.03 mg/ m
3
/ yr in
the most oligotr ophic ar eas in the Levantine sea, to the
south of Cr ete and Cypr us. The pr imar y pr oduction, the
flux of par ticles along the water column ( total par ticle
flux between 32.9 and 8.1 g/ m
2
/ yr at 80 and 1000 m, r e-
spectively: Miquel et al., 1994 in the Ligur ian Sea) and
the concentr ation of ( total or labile) or ganic matter on
deep-sea bottoms follow a seasonal patter n with peaks
that seem to be coupled with the spr ing peak of pr ima-
r y pr oduction ( Car pine, 1970; Danovar o et al., 1999;
Car tes et al., 2002) . Sur face pr imar y pr oduction often
peaks ar ound Apr il ( Car tes et al., 2002) , while or gan-
ic matter flux is at a maximum in June ( Miquel et al.,
1994) in the Ligur ian sea. For or ganics in deep sedi-
ments, ther e is only discontinuous data available ( Car tes
et al., 2002) . Peaks seem to occur also ar ound June
( Meder nach, 2000; Car tes et al., 2002) . Pr imar y pr o-
duction, phytoplankton pigment concentr ation and flux
of par ticles ar e, for instance, lower in the Catalan sea
( NW Mediter r anean) than in the Bay of Biscay ( NE At-
lantic) , located at similar latitude ( Buscail et al., 1990,
summar ized by Car tes et al., 2001b) . Par ticulate or -
ganic matter ( POM) in deep bottoms, although var ying
depending on local conditions, is also low in the deep
Mediter r anean, par ticular ly in the Easter n Basin ( Dano-
var o et al., 1999) .
Danovar o et al. ( 1999) r epor ted that mass fluxes at
equal depths ar e up to two or der s of magnitude high-
er in the Wester n Mediter r anean ( Gulf of Lions) than
in the Easter n Mediter r anean ( Cr etan sea) . 10% of the
car bon in sur face water s is expor ted to 1000 m depth in
the Wester n Mediter r anean, but only 2-3% in the East-
er n Mediter r anean, whilst bacter ial densities ar e 4 times
higher in the for mer than in the latter . The same author s
also r epor ted differ ent efficiencies in the tr ansfer of or -
ganic matter to the deep sea between the west and the
east 10% and 1% r espectively. This has pr ofound im-
plications in ter ms of bentho-pelagic coupling.
The high ther mal stability of the deep Mediter r ane-
an pr obably contr ibutes to a r apid degr adation of the
POM r eaching the deep bottoms, r esulting in poor qual-
ity ( or r efr actor y) food for the benthos. In the absence
of changes in abiotic factor s such as temper atur e, food
availability and local pr oductivity r egimes might explain
impor tant biological featur es ( e.g. species composition
and size distr ibution) of the Mediter r anean fauna. The
tr ophic level of species and the dietar y diver sity explain
patter ns of distr ibution over lap ( i.e. possible compe-
tence between species) , and the depth r ange occupied
by Mediter r anean top pr edator s below 1000 m ( Car tes
and Car r assn, 2004) . In the context of a gener ally ol-
igotr ophic r egion, such as the deep Mediter r anean, it
is of par ticular significance to under stand the function-
ing of ar eas of compar atively higher pr oduction, such as
canyons and upwelling r egions.
With the exception of some extr eme envir onments ( cold
seeps) , most deep ecosystems in the Mediter r anean ar e
allocthonous ( i.e., they depend on POM or iginating in
the photic zone or of ter r igenous or igin) . At pr esent,
most of the infor mation on the str uctur e and functioning
of tr ophic webs concer ns megafauna dwelling on mud-
dy bottoms. Detailed infor mation on diets and r esour ce
par titioning of fish and decapod cr ustaceans available
fr om the Catalano-Balear ic Basin ( wester n Mediter r a-
nean) indicates:
1) Diets ar e gener ally highly diver sified ( Shannon index
r eaching 5.3 bits in the diet of the r ed shr imp Aristeus
antennatus: Car tes, 1994) ;
2) Fish and decapods ar e or ganized in thr ee differ ent
tr ophic levels, as r epor ted fr om isotopic analyses ( Pol-
unin et al., 2001) ;
23
3) The occur r ence, both among decapods and fish, of 3
basic tr ophic guilds ( functional gr oups) , composed of
meso-bathypelagic*, benthopelagic* and benthic feed-
er s, which r espectively pr ey on macr oplankton, supr ab-
enthos and macr obenthos compar tments;
4) Ther e is a high degr ee of r esour ce par titioning* ( low
dietar y over lap) among deep-sea fish and decapod cr us-
taceans ( Macpher son, 1979; Car tes, 1998b; Car r assn
and Car tes, 2002) . Pr ey size and pr ey mobility ar e the
two main factor s explaining this tr end;
5) Decapods ar e not only detr itivor es or scavenger s, as
pr eviously assumed, but active pr edator s selecting, as
fish do, the size of their pr ey ( Car tes, 1993; Fanelli and
Car tes, 2004) . Among decapod cr ustaceans, however ,
ther e is an incr ease of detr itivor e habits below 1200 m
coinciding with the r ar efaction of some impor tant pr ey
( euphausiids, macr ofauna such as Calocaris macan-
dreae) at that depth, and with a boundar y of community
change at that same depth level ( Car tes and Sar d, 1993;
Car tes, 1998b) ; and
6) Ther e is a tr end to a decr ease of feeding intensity
with depth ( between 500-2200 m) both for decapods
( Car tes, 1998b) and fish ( Car r assn and Car tes, 2002) ,
which suggests a r eduction of metabolic activity with in-
cr easing depth ( in connexion with the extensive wor k
by Childr ess ( e.g., 1995) , showing the same tr end for
oxygen consumption data) . This gener al patter n, how-
ever , may var y depending on seasonality ( pr ey availabil-
ity) and the r epr oductive state of species ( Maynou and
Car tes, 1998) . As a gener al conclusion, in deep envir on-
ments tr ophic r elationships ar e complex, indicating that
we ar e wor king mor e within the str uctur e of a tr ophic
web than in a tr ophic chain ( as occur s, for instance, in
epipelagic envir onments) .
A similar degr ee of r esour ce par titioning is mor e diffi-
cult to establish for benthic macr o- or meiofauna. It has
been documented for macr ofauna ( e.g. supr abenthos:
Car tes et al., 2001a; 2001b) and even for meiofauna
( for aminifer ans: Gooday et al., 1992) with dir ect ( e.g.
consuming labile/ r efr actor y or ganic matter ) , or indi-
r ect ( e.g. via consumption of for aminifer ans by isopo-
ds) r esponses to POM fluxes pr obably depending on the
tr ophic level occupied by each species or taxon.
Surface pri mary pro ducti o n i s l o w i n the Me d-
i te rrane an s e a and e xtre me l y l o w i n the mo re
o l i go tro phi c are as , s uch as the Cre tan s e a.
Se as o nal pe aks i n pri mary pro ducti o n are re -
fl e cte d i n s e as o nal pe aks o f s e co ndary pro duc-
ti o n i n the de e p s e a, as we l l as o the r s e as o n-
al change s o f fo o d co ns umpti o n and re pro duc-
ti o n.
The l o w fo o d i nput to the de e p- s e a re s ul ts i n
s carce fo o d re s o urce s , hi gh fo o d parti ti o n-
i ng, hi ghl y di ve rs i fi e d di e ts , and ve ry co mpl e x
tro phi c we bs .
3.2. Mesoscale* variations
in the dynamics of deep-sea
ecosystems
In addition to tempor al changes, deep-sea ecosystems
also var y on a spatial scale. Differ ences between the
easter n and wester n Mediter r anean ar e mor e evident
for fish: cer tain dominant species ar e found only in the
wester n basin ( e.g. Centroscymnus coelolepis, Polya-
canthonotus rissoanus, Chalinura mediterranea, Le-
pidion lepidion, Alepocephalus rostratus, Nemichthys
scolopaceus, Nettastoma melanurum; Whitehead et
al., 1989; DOnghia et al., 2004; Llor is, per s. obs.) . In
other cases, it has been suggested that the small differ -
ences between the easter n and wester n basins should be
simply attr ibuted to differ ences in sampling effor t ( Bel-
lan-Santini, 1990) . At mesoscale, deep Mediter r anean
fauna r egular ly show a high similar ity between neigh-
bour ing ar eas. Thus, only 11.6 % of the hyper benthic
per acar id cr ustaceans found in the SW Balear ic slope
had not been r epor ted pr eviously 350 km away in the
Catalan Sea ( Car tes and Sor be 1993, 1995, 1999) .
In addition to the biogeogr aphical var iations cited,
changes in the functioning of food webs have been r e-
por ted at mesoscale ( 350 km) in the wester n basin. De-
pending on the distance to the coast ( and thus, to the
influence of advective fluxes of mainland or igin) , the
tr ophic r esour ces of deep sea fauna ar e based mor e on
the benthic or pelagic compar tments. For instance, on
the middle slope of the continental side of the Catalan
Sea, benthos was the main contr ibutor to the food sup-
ply for megafauna ( cr ustaceans and fish) , after a study
on the food consumption-food supply balance ( Car tes
and Maynou, 1998) . In contr ast, in a mor e open-sea
ar ea ( e.g. the SW Balear ic Islands) bathyal fish r elied
mor e on pelagic pr ey as a food r esour ce. A wester n-
easter n incr ease in oligotr ophic conditions in the Medi-
ter r anean seems to have similar consequences, and fish
assemblages in the Ionian Sea exploit pr efer entially pe-
lagic pr ey, a featur e consistent with the low abundance
24
Box 2. Reproductive strategies of deep-water species:
recruitment patterns in productivity hotspots
Although most studies wer e or iginally focused on com-
mer cial species of fish and decapod cr ustaceans ( e.g.
the r ed shr imp Aristeus antennatus) , ther e is now de-
tailed infor mation on the biological cycles of a number
of deep Mediter r anean species. Supr abenthic per acar -
id cr ustaceans ( Car tes and Sor be, 1999; Car tes et al.,
2001b) , and non-commer cial, though ecologically
impor tant, fish and decapods have been the topic of
these studies ( e.g. Macr our idae: Massut et al., 1995;
DOnghia et al., 1999; Pandalidae: Company and Sar d,
2000; Pasiphaeidae: Company et al., 2001; Polycheli-
dae: Abell and Car tes, 1992) . Both continuous and
non-continuous r epr oductive patter ns have been doc-
umented, with some tr end towar ds seasonality, or an-
nual r estr icted per iods of matur ity, in mid-slope dwell-
ing species with incr easing depth, both among Cuma-
cea ( Car tes and Sor be, 1996) , and Pandalidae ( Com-
pany and Sar d, 2000) . Biological tr ends with depth
have also been studied in the last decade ( Stefanescu
et al., 1992; Sar d and Car tes, 1993) . Some dominant
species follow a bigger -deeper and smaller -shallower
tr end ( e.g. Phycis blennoides, Mora moro, Plesioni-
ka martia) pr efer entially on the upper and middle
slope, while some show a smaller -deeper tr end ( e.g.
Aristeus antennatus) , on the lower slope, and some
species do not show any significant size-r elated tr end
with depth at all ( Mor ales-Nin et al., 2003) . Pr oba-
bly the depth of r ecr uitment and r epr oductive aggr e-
gations by deep-water species depend on a number of
physical and tr ophic factor s which may var y between
species. A link has been suggested, for example, be-
tween phytodetr itus* deposition and r ecr uitment
for small supr abenthic species ( Car tes et al., 2001a,
2001b) . In megafauna, a link between r ecr uitment and
the occur r ence of nepheloid layer s ( Puig et al., 2001)
has been shown for pandalid shr imps.
Puig et al. ( 2001) showed that the lar val and r ecr uit-
ment pr ocesses of thr ee deep-water pandalid shr imps
( genus Plesionika) wer e r elated to nepheloid de-
tachments at the continental mar gins, along a nar r ow
bathymetr ic r ange of ca. 400 m depth. The existence
of a nepheloid layer is the r esult of the over all water
mass cir culation in the NW Mediter r anean ar ea ( fr on-
tal str uctur e) . Fr ontal str uctur es located at ar ound
400 m depth ar e widespr ead in the oceans wor ldwide
( Puig et al., 2001) and suggest the existence of a pr ef-
er ential r ecr uitment habitat for some continental slope
species.
/
m
O
O 1
O 2
O 8
O
O 1
O 2
O 8
O
O 1
O 2
O 8
O 4
2 2 O 1 O 1 4
h P
O
O 1
O 2
O 8
8 2 2 2 O 1 O 1
e P
O
O 1
O 2
O 8
O 4
2 2 O 1 O 1 4
g P
O
O 1
O 2
O 8
O
O 1
O 2
O 8
O
O 1
O 2
O 8
2 2 O 1 O 1 4
a P
7 O = n
8 2 4 = n
O 2 2 = n
8 7 = n O 8 1 = n
7 8 = n O O 1 = n
8 8 = n O 9 = n 7 9 = n
7 O = n O O 1 = n 9 O = n
8 O = n
4 9 1 = n
8 9 = n
9 2 = n
4 2 8 1 2 1 O
m P
O
O 1
O 2
O 8
8 2 1 = n
O
O 1
O 2
O 8
7
2 2 O 1 O 1 4
j m m ( l C
s e l i n e J
s t l d A
8 . O <
8 . O <
O 25 5O 75 1OO
1OO
5OO
1OOO
28.4
nm
Foo (1988)
P|g (2001)
8i e freq enc histograms
s.
Partic late matter concentration (mg/lj
d ring April 1998

t
25
Another impor tant aspect of the population dynamics
of deep-water species is the pr esence of seasonal r e-
pr oduction peaks, despite the appar ent stability of ec-
ological conditions in the deep sea. A study on the du-
r ation of the r epr oductive per iods of 17 deep-water
decapod cr ustaceans ( some of them dwelling below
1000 m) showed that their r epr oductive per iods wer e
mor e clear ly seasonal than those of species dwelling
along the shelf and upper -slope continental mar gins
( Company et al., 2003) . The shorter r epr oductive pe-
r iods shown by deep-sea dwelling species ( Fishelson
and Galil, 2001; Company et al., 2003) ar e an aspect
of their life histor ies that might affect their commer cial
exploitation. The r epr oductive output of deep-sea spe-
cies could be compr omised if any exploitation is un-
der taken dur ing their shor t r epr oductive per iod.
Box 3. Diversity, biodiversity and species richness
Gr aphical synthesis of the thr ee for ms to under stand the ter m biodiversity.
Diversity
2 3 8 5 in d ivid u a ls
1 1 6 k ilo g ra m m e s
3 sp e c ie s
Boops boops
1195 individuals
60 kilogrammes
Diplodus annularis
760 individuals
30 kilogrammes
Pagellus acarne
430 individuals
26 kilogrammes
Diversity
Di ve rs i ty
r efer s to theabsolutenumber of species
present, or any other measurement
that incorporates thenumber of species
as well as its relativeabundance
( Nor se et al., 1980 inWilson, 1988) .
Biodiversity
<
1 F a m ily
1 G e n u s
3 sp e c ie s
Scorpaena notata
Scorpaena porcus
Scorpaena scrofa
1 F a m ily
3 G e n u s
3 sp e c ie s
Boops boops
Diplodus annularis
Pagellus acarne
Biodiversity
<
Species richness Species richness
M e d ite rra n e a n sh e s
to ta l c a ta lo g u e d
6 5 7 sp e c ie s
169 pelagic species
488 Demer sal species
E x c lu sive ly
in so ft b o tto m s
( m u d & sa n d )
257 species
H a rd & S o ft
b o tto m s
107 species
E x c lu sive ly
in h a rd b o tto m s
124 species
Bi o di ve rs i ty
r efer s to thevariety of organisms considered at
all thelevels, fromthegenetic variants
pertainingto thesamespecies, to varieties,
species, genera, families and even superior
taxonomic levels, includingthevariety
of ecosystems, that includes thecommunities
of organisms that livein certain habitats as
well as thephysical trainingconditions under
which they live( Wilson, 1992) .
Spe ci e s ri chne s s
r efer s to a list of species
at a given location or community
( Puttman, 1994) .
26
of macr obenthos and meiobenthos biomass in the east-
er n Mediter r anean below 400 m ( Tselepides and Elefth-
er iou, 1992; Danovar o et al., 1999) .
As a r esult of these ecological patter ns, it is impor tant
to note that:
( i) Fish assemblages can change depending upon the
tr ophic char acter istics of the ar ea they inhabit.
Thus, dominant species in the easter n Ionian Sea,
mainly depending on pelagic r esour ces ( e.g. Ho-
plostehtus mediterraneus and Chlorophthalmus
agassizi) , ar e r ar e or absent in mor e eutr ophic ar -
eas, such as the continental side of the Catalan Sea
( Madur ell et al., 2004) ; and
( ii) a compar ison between ar eas under mainland influ-
ence and far fr om this influence ( insular ar eas) in
the deep Mediter r anean evidenced local changes in
the bathymetr ic distr ibution and zonation of deep
sea fauna ( Maynou and Car tes, 2000; Car tes et al.,
2004) .
In conclusion, an incr ease in oligotr ophic conditions ( in
ter ms of pr imar y pr oduction in sur face water s) favour s
the exploitation of pelagic r esour ces, instead of benthos,
by deep sea top pr edator s. In these oligotr ophic ar eas
( e.g. the Ionian Sea) , fish tend towar ds a str ategy of ca-
lor ic maximization consuming pr ey fr om the mesope-
lagic compar tment, mor e ener gy-r ich than benthos.
Food availability decr eases even mor e below 1000 m in
the deep Mediter r anean, despite the biomass incr ease
r epor ted at ar ound 1200 m in the NW Mediter r anean
for fish. This peak of biomass ( s e e Bo x 1 ) , fir st de-
scr ibed as a gener al tr end in the Nor th Atlantic ( Gor -
don and Duncan, 1985) was also found in the wester n
Mediter r anean ( Stefanescu et al., 1993; Mor anta et al.,
1998) , though it has not conclusively r epor ted for the
Easter n Basin. Some species r esponsible for this peak
that ar e r ecor ded in the wester n Mediter r anean ar e ab-
sent ( e.g. Alepocephalus rostratus) in the Easter n Ba-
sin, wher e studies of megafaunal biomass distr ibution
ar e par ticular ly scar ce ( Kallianotis et al., 2000; Politou
et al., 2003) , r eaching only 1000 m depth. On the Cr e-
tan slope, megafaunal biomass showed a peak at 500 m
( Kallianotis et al., 2000) and did not show a significant
decr ease with depth between the two deepest str ata sam-
pled ( 800 and 1000 m) , even showing a non-significant
biomass incr ease at 1000 m. Some fish species r espon-
sible for the biomass peak in the wester n Basin ar e still
dominant in fish assemblages ( e.g. Mora moro, Galeus
melastomus: Kallianotis et al., 2000) at 1000 m depth
in the Easter n basin. In the wester n Mediter r anean, be-
low 1000 m, dominant species such as Alepocepha-
lus rostratusbase their diet on macr oplankton, adopt-
ing the str ategy of species inhabiting low pr oductive r e-
gions. Consistently, benthic biomass also decr eases at
those depths ( Car tes et al., 2002) . Fur ther , a number of
fish species ( e.g. Phycis blennoides, Mora moro, Tra-
chyrhynchus trachyrhynchus) ar e r epr esented ar ound
these depths almost exclusively by lar ge specimens, with
a clear bigger -deeper tr end in their size distr ibutions
( Stefanescu et al., 1993) , and with a pr ogr essive de-
pendence on benthopelagic r esour ces with incr easing
size. Food consumption also decr eases with incr eas-
ing fish size, and the bathymetr ic aggr egation of these
lar ge sizes for these species ar ound a nar r ow depth
r ange ar ound 1200 m pr obably indicates the incr eas-
ing scar city of food r esour ces with incr easing depth be-
low 1000 m.
The gener al pr inciple that it is essential to know the car -
r ying capacity of ecosystems befor e any exploitation be-
gins, is even mor e cr ucial in the case of deep-water , low
pr oductive, ecosystems. Tempor al studies, following not
only the dynamics of commer cial species, but the whole
functioning of the ecosystem, ar e essential. In par ticular
must be addr essed: ( 1) the pr oductive capacity of the
lowest tr ophic levels macr ofauna, including the swim-
ming compar tment ( supr abenthos, macr ozooplank-
ton) ; and ( 2) the complexity of food webs number of
tr ophic levels involved and the food consumption of
top pr edator s.
Depth-distr ibution balances based on pr oduction r ath-
er than biomass constitute a mor e r ealistic appr oach to
better under standing the dynamics of deep-water eco-
systems. The evidence pr esently available points to the
Mora moro ( Risso, 1810)
Mota mediterranea,
Mora,
V. Fossat, 1878.
Coll.
Musum dHistoir e
natur elle de Nice.
27
conclusion that in the Mediter r anean, depths below
1000 m ( or most appr opr iately, we should r efer to the
lower slope assemblages: Pr s, 1985) ar e par ticular ly
poor and especially sensitive to futur e human inter ven-
tion, for instance fishing.
De pths be l o w 1 0 0 0 m i n the Me di te rrane an
are parti cul arl y fragi l e and e s pe ci al l y s e ns i -
ti ve to future human i nte rve nti o n, fo r i ns tance
fi s hi ng.
The ge ne ral pri nci pl e that i t i s e s s e nti al to
kno w the carryi ng capaci ty o f e co s ys te ms be -
fo re any e xpl o i tati o n be gi ns , i s e ve n mo re cru-
ci al i n the cas e o f de e p- wate r, l o w pro ducti ve ,
e co s ys te ms .
3.3. Human alteration of trophic webs
Anthr opogenic impacts on deep Mediter r anean commu-
nities ( e.g. fisher ies, tr awling, and waste disposal) may
have a str ong influence on the dynamics of such fr agile
ecosystems, although the number of studies addr essing
this issue is still limited. Pr esent knowledge of tr ophic
dynamics available for the deep Mediter r anean shows
that:
1. Gor gonian communities ( e.g. Isidella elongata) and
other sessile or ganisms ar e immediately r emoved fr om
soft bottoms after tr awling, changing the associated bio-
coenosis, which may compr ise species of commer cial
inter est ( such as r ed shr imps) in the case of Isidella
biocenose. Fr om a tr ophic per spective, communities
may change fr om those dominated by filter feeder s ( e.g.
sponges, br achiopods such as Gryphus vitreus) and
low-tr ophic level pr edator s ( gor gonians) towar ds oth-
er communities dominated by deposit feeder s and top
pr edator s. Tr awling may also have a negative impact
on colonies of other har d-bottom dwelling cor als ( e.g.
Lophelia pertusaand Madrepora oculata) , some r ar e,
living in the deep Mediter r anean.
2. Tr awling may also alter the secondar y pr oduction of
benthic communities, with incr easing dominance of
species of higher P/ B r atio, by decr easing the mean in-
dividual size. Additionally, a decr ease in the mean tr oph-
ic level of food webs is also expected.
3. The influence of discar ds and the damage induced on
benthic species has r eceived less attention, but it may
change the food consumption patter ns of species that
ar e, only potentially, oppor tunists and scavenger s. Dis-
car ds of non-commer cial species ar e a non-tr ivial par t
of the catch biomass in fisher ies of the r ed shr imp Aris-
teus antennatus( 27%: Car bonell et al., 1998; Bozzano
and Sar d, 2002) . Ther efor e, the feeding behaviour of
species and the whole dynamic of such low-pr oductive
ecosystems can also be alter ed.
4. Mar ine pollution can be channelled to the deep-sea
by submar ine canyons, as r epor ted for the Toulon Can-
yon in the wester n Mediter r anean ( Stor a et al., 1999) .
As a r esult of dr edge spoil dumping at the canyon heads,
str ong concentr ations of heavy metals and or ganic mat-
ter have accumulated deeper . This enr ichment in or gan-
ic matter even favour s the colonization of bathyal depths
by some species living in har bour s, in shallow water . Or -
ganochlor inated compounds ar e another sour ce of pol-
lution of the deep Mediter r anean, evidenced by the en-
zymatic xenobiotic activities detected in deep-sea fish
living between 1500-1800 m depth, which indicate the
ability of such or ganisms to cope with these pollutants
( Por te et al., 2000) . Differ ences between the r espons-
es by species wer e suggested in r elation to the feeding
behaviour of each species. Accumulation of solid waste
( plastics, etc.) is impor tant in the Catalano-Balear ic Ba-
sin, pr obably because it is channelled by the numer ous
canyons occur r ing in the continental par t of this basin.
The influence on the r ecr uitment and incor por ation of
cer tain mater ials ( e.g. tar ) to the tr ophic webs is un-
known.
5. Finally, climate change induced by human activity in-
fluences the quantity and quality of food r eaching the
deep-sea Mediter r anean ecosystems. Contr ar y to what
might be expected, deep-sea ecosystems would r espond
quickly to climate change ( Danovar o et al., 2001) .
The de e p Me di te rrane an bi o l o gi cal co mmu-
ni ti e s are adapte d to a ge ne ral o l i go tro phi c
e nvi ro nme nt, wi th l o cal are as o f hi ghe r pro -
ducti vi ty and bi o di ve rs i ty ho ts po ts .
The s e bi o l o gi cal co mmuni ti e s are ve ry s e ns i -
ti ve to human mo di fi cati o n i n the fo rm o f
de e p- s e a trawl i ng, was te di s po s al o r che mi cal
po l l uti o n.
Human mo di fi cati o n i mpacts di re ctl y o n the
co mmuni ti e s by s e l e cti ve l y affe cti ng s o me o f
the i r co mpo ne nts ( e . g. re mo val o f to p pre da-
to rs by fi s hi ng, de s tructi o n o f s us pe ns i o n fe e d-
e rs s uch as go rgo ni ans o r co ral re e fs ) , o r i ndi -
re ctl y by changi ng the s tructure o f a co mpl e x
tro phi c we b, mo di fyi ng s e co ndary pro ducti o n
patte rns o r the e ffe ct o f cl i mate change .
28
Box 4. Mass fluxes in deep-water ecosystems
A theor etical scheme of mass fluxes thr ough deep-
water ecosystems inhabiting the Benthic Boundar y
Layer ( the layer close to the sea floor wher e an
impor tant incr ease of living biomass is r ecor ded) .
Two main pathways ar e involved ( the advective flux
of mater ial of ter r igenous or igin and the ver tical flux
of mater ial of pelagic or igin) . Fluxes of par ticulate
or ganic matter ( POM) ar e consumed pr imar ily by
benthos and supr abenthos close to the bottom, and by
zooplankton-micr onekton along the water column by
means of chained ver tical migr ations. These or ganisms
constitute the basis of the diet of benthopelagic
megafauna ( fish and lar ge decapod cr ustaceans) . The
ver tical and advective inputs var y depending on local
pr oductivity ( r iver dischar ges, upwelling ar eas linked
to eddies and oceanogr aphic cir culation) , favour ing
the density of benthos-supr abenthos/ zooplankton, and
r esulting in food webs wher e top pr edator s pr ey on
pelagic or benthic or ganisms.
Ter r igenous
mater ial
Advective
flux
BBL
De tri tus
Faecal pellets
Plant r emains
Cetacean car casses
Pr imar y pr oduction
phytoplankton
Supr abenthos
Infauna
Phytodetr itus
Megafauna
( fishes, decapod
cr ustaceans)
Vertical flux
Ver tical
migr ations
Scheme of energy fluxes in the BBL
Micr onekton
29
Cr edits: GEBCODigital Atlas, 2003, Modified.
4. Unique environments
of the Mediterranean deep-sea
4.1. Cold seeps
Upwar d seepage of cold fluids enr iched in methane oc-
cur s wor ldwide along cer tain deep-sea tectonic featur es
such as mud volcanoes ( Henr y et al., 1996) . These ar e
the home of unique chemosynthesis-based communities
( not r elying on photosynthetic pr oduction) dominat-
ed by bacter ial mats and par ticular species of bivalves
and tubewor ms, that ar e associated with endosymbiot-
ic* chemo-autotr ophic* bacter ia. Specially evolved bac-
ter ia able to oxidize the methane ar e at the basis of this
unique food web. Impor tant cold seep ar eas hosting
such unique benthic communities wer e fir st descr ibed
in the Atlantic and in the Easter n and Wester n Pacific
Oceans ( Kennicut II et al., 1985) .
Initial evidence of benthic communities based on chem-
osynthesis in the Mediter r anean r efer r ed to the Napo-
li mud volcano, on the top of the Napoli dome on the
Mediter r anean Ridge, at depths of 1900-m ( Cor selli and
Basso, 1996) . Cold seep biological communities r ely-
ing on methane and associated to mud volcanoes and
faults have r ecently been discover ed in the southeast-
er n Mediter r anean Sea, south of Cr ete and Tur key ( in
the Olimpic field and Anaximander mountains, r espec-
tively; MEDINAUT/ MEDINETH Shipboar d Scientific Par -
ties, 2000) , at depths of between 1700-2000 m, as well
as nor th of Egypt near the Nile delta. In the latter local-
ity ( near Egypt and the Gaza Str ip) , living communities
of polychaetes and bivalves have been found at depths of
500-800 m ( Coleman and Ballar d, 2001) .
The isolation of the Mediter r anean deep-sea seeps and
vent habitats fr om the Atlantic Ocean has r esulted in the
development of unique communities, as illustr ated by
the specific bivalve populations associated to Mediter -
r anean cold seeps, with species much smaller in size
than those dominating in other seep sites. The biologi-
cal community inhabiting the Anaximander mountains
field, SW Tur key, is dominated by small bivalves belong-
ing to 4 families ( Lucinidae, Vesicomydae, Mytilidae and
Thyasir idae) and tube wor ms ( both pogonophor an and
vestimentifer an species) , all of which contain bacter ial
endosymbionts. Wher eas wor ms, vesycomid and lucinid
bivalves r ely thr ough their sulfur -oxydizing-type sym-
bionts on the sulphide issued fr om micr obial sulphate
r eduction pr ocesses in the super ficial sediment, the my-
tilid symbionts ar e able to use either sulphides fr om the
sediment or , dir ectly, the methane expelled in the seeps
( Fiala-Mdioni, 2003) .
Unlike cold seeps, which ar e well r epr esented along the
Mediter r anean Ridge, active deep-sea hydr other malism
seems to be absent fr om the Mediter r anean. Known hy-
dr other mal vents in the r egion occur in shallow water s
( < 100-m) , associated to volcanic ar cs such as the
Helenic Volcanic Ar c. In these ar eas, tr ophic webs ar e
mostly based on photosynthetic pr imar y pr oduction and
the associated macr o-epibenthic biological assemblag-
es ar e not distinct fr om the sur r ounding ar eas ( Cocito
et al., 2000) .
Cr edit: Coleman and Ballar d ( 2001) . Spr inger .
Anaximander
Mountains Olimpi
Mud Volcanoe
Field
Napoli Dome
Gaz Bubbles
30
Recent geological cr uises involving multibeam mapping
and use of the side scan sonar have uncover ed the ex-
istence of numer ous mud-volcanoes and anoxic basins
in the Easter n Mediter r anean ( Desbr uyr es, 2003) .
Evidence of pr eviously unknown mud-volcanoes in the
Easter n Mediter r anean opens the door to an incr eased
occur r ence of cold-seep type communities in the r egion.
The r ecent discover y of pockmar ks ( geological str uc-
tur es consisting of shallow cr ater s typically 30-40 m in
diameter and 2-3 m deep) ar ound the Balear ic Islands
( Acosta et al., 2001) also points to the existence of these
cold-seep type communities in the Wester n Mediter r a-
nean. In the major ity of cases, the mechanism of pock-
mar ks for mation is attr ibutable to gas dischar ges.
Co l d- s e e ps harbo ur uni que bi o ce no s e s bas e d
o n the o xi dati o n o f me thane as the pri mary car-
bo n s o urce ( i . e . , no t bas e d o n pho to s ynthe ti c
pro ducti o n as i n mo s t mari ne e nvi ro nme nts ) ,
do mi nate d by bacte ri al mats and co mmuni ti e s
o f s pe ci al i s e d bi val ve s and tube wo rms .
Box 5: Mud volcanoes
The ter m mud-volcano is gener ally applied to a mor e
or less violent er uption or sur face extr usion of water y
mud or clay which is almost invar iably accompanied by
methane gas, and which commonly tends to build up a
solid mud or clay deposit ar ound its or ifice which may
have a conical or volcano-like shape. Mud volcanoes
also commonly appear to be r elated to lines of fr actur e,
faulting, or shar p folding.
In the Mediter r anean, mud volcanoes ar e found along
the Mediter r anean Ridge, in the South Easter n Mediter -
r anean ( i.e. Napoli Dome, Olimpi mud volcano) .
The motivating for ce r esponsible for a mud volcano is,
in par t, simply the weight of r ock over bur den bor ne by
the fluid content of under compacted shales. However ,
mud volcanoes all over the wor ld ar e associated so in-
var iably with quietly or explosively escaping methane
gas, that it is r easonable to conclude that the pr esence
of methane gas in the subsur face is also an essential
featur e of the phenomenon. The mud of the volcanoes
is a mixtur e of clay and salt water which is kept in the
state of a slur r y by the boiling or chur ning activity of
escaping methane gas. Some liquid oil is often, but not
always, associated with the hydr ocar bon gases of mud
volcanoes.
Commonly, the activity of a mud volcano is simply a
mild sur face upwelling of muddy and usually saline wa-
ter accompanied by gas bubbles.
Sour ce: Geological Society of Tr inidad and Tobago.
Cr edit:
Loubr ieu B., Satr a C., 2001. Car togr aphie par sondeur multifais-
ceaux de la Ride Mditer r anenne et des domaines voisins. Comit
Franais deCartographie, n 168, pp. 15-21.
31
4.2. Brine pools
The easter n Mediter r anean Sea hosts a unique envi-
r onment that r anges among the mor e extr eme ever de-
scr ibed in r elation to its compatibility with life. Five deep
hyper saline anoxic basins ( DHABs) wer e r ecently dis-
cover ed on bottoms below 3300 m depth, i.e. in an envi-
r onment with a complete absence of light and subject to
high pr essur es ( Lampadar iou et al., 2003) . DHABs ar e
char acter ised by a salinity value higher than 30%, oxy-
gen depletion and elevated methane and sulphide con-
centr ations ( De Lange et al., 1990) ; the Ur ania basin
pr esents the highest concentr ation of sulphide among
the Ear th aquatic envir onments. These unique envir on-
ments have been isolated fr om the global ocean for mil-
lions of year s. Sur face of known DHABs r ange fr om 5
to 20 km
2
, and the str ong differ ence in density between
the br ines and the sur r ounding seawater ensur es a com-
plete lack of mixing ther eof. Newly descr ibed pr okar y-
otic gr oups have been found in the inter face ar ea that
har bour s impor tant populations of Bacter ia and Ar -
chaea. Fir st genetic evidence point to DHABs as har -
bour ing a much higher diver sity of unknown extr emo-
philic bacter ia than other hyper saline anoxic basins in
the wor ld. DHABs ar e toxic to megafauna and macr ofau-
na, so they ar e only able to suppor t pr otistan and meio-
faunal communities that likely benefit fr om symbiotic
associations with pr okar yotes. Such meiofaunal assem-
blages ( composed of nematodes, copepods, for aminif-
er a, etc.) could even r each biomass values much higher
than those found outside the br ine pool, and many of
the species involved ar e thought to be new for science
( Lampadar iou et al., 2003) .
Bri ne po o l s ( o r De e p Hype rs al i ne Ano xi c Ba-
s i ns , DHABS) harbo ur uni que faunal as s e m-
bl age s , adapte d to wi ths tand hi gh s al i ni ty l e v-
e l s ( 3 0 PSU) , o xyge n de pl e ti o n and hi gh co n-
ce ntrati o n o f me thane and s ul phi de .
4.3. Deep-sea coral mounds
Although most scler actinian r eef-for ming cor als oc-
cur in tr opical r egions and in shallow water , ther e is
a gr oup of scler actinian cor als which can exist in wa-
ter between 4 and 12 C, and at depths fr om ca. 50 m
to over 2000 m wher e they typically settle escar pments,
seamounts and over hangs in total dar kness. These cor -
als do not have symbiotic algae, but ar e still able to for m
a har d skeleton. They for m colonies and can aggr egate
into patches and banks which may be descr ibed as r eefs.
The most common cold water cor al is Lophelia pertu-
sawhich has a global distr ibution but is most common
in the nor th-east Atlantic. It for ms extensive buildups in
the Atlantic Ocean between ca. 300-800 m, often in as-
sociation with Madrepora oculataand Desmophyllum
cristagalli. A study of L. pertusacor al r eefs off the Fae-
r oes indicated that the diver sity of L. pertusacor al r eefs
is of a similar magnitude to that of some tr opical, shal-
low water her matypic cor als. The over all faunal diver si-
ty and the number of species within many faunal gr oups
( for aminifer a, por ifer a, polychaetes, echinoder ms and
br yozoans) wer e found to be similar . The diver sity of
the taxa associated with the L. pertusar eefs is ar ound
thr ee times as high as that of the sur r ounding soft sedi-
ment seabed, indicating that these r eefs cr eate biodiver -
sity hotspots and incr eased densities of associated spe-
cies ( Tur si et al., 2004) .
So-called cold-water cor al r eefs, which ar e cur r ently
the object of str ong conser vation effor ts in the NE At-
lantic ( Gubbay, 2003) , ar e also pr esent in the Mediter -
r anean. Though most of such occur r ences ar e subfossil
and date back to the last glacial age, witnessing cooler
seawater and better food availability, some living r elict
r eefs have been found ( Mastr ototar o et al., 2002; Tur -
si et al., 2004) . Dur ing the coldest phases of the Pleis-
tocene, these deep-sea cor al banks wer e equally com-
mon in the Mediter r anean basin, as pr oven by their r e-
cur r ence within outcr opping and submer ged fossil as-
semblages found on the tops and flanks of submar ine
canyons, seamounts and banks of the whole Mediter -
r anean at water depths in excess of ca. 300 m ( Pr s,
1985) . The postglacial onset of compar ably war m, ho-
mother mic, nutr ient-poor deep water s in the Mediter r a-
nean basin has been advocated as the major factor con-
tr olling the decline and almost total disappear ance of
these deep-sea cool water cor als, although an indir ect
human impact cannot be r uled out. Pr s ( 1985) points
Lophelia pertusa. Cr edit: Angelo Tur si. Se e co l o ur pl ate , p. 6 2 .
32
out that many subfossil white cor al mounds ar e cover ed
by a fine layer of sediment, possibly accumulated since
histor ic times, due to pr ogr essive for est destr uction by
man. The most impor tant fr ame-builder Lopheliahas
been identified as par ticular ly vulner able to the ocea-
nogr aphic changes linked to the glacial-postglacial tr an-
sition, whilst viable Madreporamounds still occur in
some spotty ar eas of the Mediter r anean basin. Deep-wa-
ter cor als ar e passive suspension feeder s, and they ar e
pr efer entially distr ibuted on topogr aphic ir r egular ities
( seamounts, pr omontor ies, canyons) . Communities of
white cor als in the deep Mediter r anean ar e disper sed
elsewher e ( e.g., Blanes canyon, Lacaze-Duthier can-
yon, Albor an Sea; Zibr owius, 1980; Zabala et al., 1993) .
Consider ed as r elict communities, they ar e mainly com-
posed of dead br anches, or with only a few ter minal por -
tions r emaining alive. These residual colonies ar e often
associated with sites r eceiving lar ger food inputs ( e.g. in
or near submar ine canyons) , and it has been postulat-
ed that the decline of these cor als in the Mediter r anean
was linked with a decr ease in tr ophic r esour ces cor r e-
lated with a water temper atur e incr ease ( Delibr ias and
Taviani, 1984, for the Albor an sea) . Recently, a healthy
and well developed Lophelia-Madreporadeep-sea cor -
al mound has been discover ed in the Ionian Sea ( Nor th
of Calabr ian Ar c) , offshor e fr om the Apulian platfor m, at
between 300 and 1000 m depth ( Giuliano et al., 2003) .
As a gener al r ule, it has been pr oposed that the last, r el-
ict, deep-water Mediter r anean cor al banks would set-
tle ar eas char acter ized by local peculiar oceanogr aph-
ic conditions, enhancing nutr ient availability which has
so far pr evented their total er adication fr om this basin,
wher e pr esent conditions ar e not adequate for them.
Though dir ect tr awling ( or other fishing methods) on
cor al r eefs is the main obvious thr eat to the r emaining
Mediter r anean deep-water cor al r eefs, tr awling in the
neighbour ing bathyal mud bottoms could be equally
deleter ious on these suspension feeder s, due to the ef-
fects of sediment r esuspension and r elated incr eased
sedimentation, even at depths well beyond the ones
tr awled. A r ecent study showed evidence of how sedi-
ment r esuspension fr om tr awler s wor king at 600-800 m
depth r eached a depth of 1200 m ( Palanques et al.,
2004) .
Co l d- wate r co ral re e fs fo rme d by l i ve co l o ni e s
o f the s cl e racti ni ans Lophelia pertusa and
Madrepora oculataare as s o ci ate d wi th hi ghl y
pro ducti ve e nvi ro nme nts , harbo ur hi gh l e ve l s
o f di ve rs i ty and are thre ate ne d ( di re ctl y and
i ndi re ctl y) by fi s hi ng.
4.4. Seamounts
Rising up fr om the sea floor , seamounts ar e steep-sided
submer ged mountains that pr ovide unique habitats in
oceans all over the wor ld. Str ictly speaking, seamounts
ar e discr ete ar eas of topogr aphic elevation higher than
1000 m above the sur r ounding seafloor ( Inter national
Hydr ogr aphic Bur eau, 2001) . The tops of seamounts
can r ange fr om near the sur face to sever al thousand
metr es below it.
While our knowledge of seamounts in the global oceans
is far fr om complete, enough sampling has been done
to show that seamounts suppor t biologically unique and
valuable habitats, being highly pr oductive, and with high
r ates of biodiver sity and endemisms ( Richer de For ges
et al., 2000) . They may act as r efuges for r elict popu-
lations or become centr es of speciation ( Galil and Zi-
br owius, 1998) .
Though not compar able to cer tain Atlantic or Pacific
ar eas, the Mediter r anean sea har bour s some impr es-
sive seamounts whose biodiver sity values ar e still poor -
ly known, in the Gulf of Lions, in the Albor an sea, in
Desmophyllumcristagalli.
Cr edit: Angelo Tur si. Se e co l o ur pl ate , p. 6 2 .
Nile Fan
Eratosthene Seamount
Cr edit: GEBCODigital Atlas, 2003, Modified.
33
Box 6. The discovery of a deep-water coral reef in the Ionian Sea
Santa Maria
di Leuca
Cr edit: GEBCODigital Atlas, 2003, Modified.
In August 2000 a scientific cr uise car r ied out by a team
of the Univer sity of Bar i r ediscover ed a living deep-sea
cor al r eef 20-25 miles off Cape of Santa Mar ia di Leuca,
souther n Italy, in the Ionian Sea. The r eef alr eady
r epor ted by Mar enzeller in 1893 is dominated by
Lophelia pertusaand Madrepora oculata, though two
fur ther species of scler actinian cor als wer e also found
( Desmophyllumcristagalli and Stenocyathus ver-
miformis) . The most impor tant r eef building species,
Lophelia and Madrepora, occur r ed in samples taken
fr om 425 to 1110 m depth.
A total 58 taxa ( including 12 species of bone fish and
5 condr ichthyans) wer e r ecover ed and identified as
char acter istic species of the r eef, associated species,
accompanying species and co-occur r ing species, de-
pending on their degr ee of r eliance on the r eef. Ap-
par ently, the ver tical flux of or ganic matter in the ar ea
is r emar kably high, which could r esult in a high avail-
ability of food to suspension feeder s like Lopheliaand
Madrepora.
Accor ding to data r ecover ed fr om the Ionian Sea r eef,
Tur si et al. ( 2004) postulate that the habitat pr ovided
by the biocoenosis of deep-water cor al in the Mediter -
r anean Sea acts as an oasis in the deser t ( biodiver -
sity hot-spot) . It cr eates a thr ee-dimensional str uctur e
that pr ovides ecological niches to a diver sity of spe-
cies, including deep-sea char acter istic species such
as the Mediter r anean or ange r oughy ( Hoplostethus
mediterraneus) and species of economic inter est
such as the r ose shr imp ( Aristaeomorphafoliacea)
or the conger ( Conger conger) . These r eefs, being a
natur al deter r ent to tr awling, ar e thought to pr oduce
a positive spill-over effect on the deep-water demer -
sal r esour ces intensively fished on the neighbour ing
muddy bottoms.
Aristeomorpha foliacea
Alber t I
er
Pr ince de Monaco. Camp. scient. Pnids pl. III. EL. Bouvier del, M. Bor r el pinx.
Madrepora oculata. Cr edit: Angelo Tur si.
34
Box 7. The Mediterranean Ridge
The Mediter r anean Ridge consists of a mor e than 1500 km long tecto-sedimentar y-accr etionar y pr ism, which r e-
sults fr om the offscr aping and piling up of thick sedimentar y sections, and which r uns fr om the Ionan basin, to
the west, to the Cypr us ar c to the east. This complex was cr eated by subduction of the Afr ican plate beneath the
Eur asian plate to the nor th. It is an extensive fold-fault system cor r esponding to r ecent uplift and folding of past
abyssal plains.
the easter n Tyr r henian basin, to the south of the Ionian
abyssal plain and in the Levantine seas. Located off the
south coast of Cypr us and west of Isr ael lies the mas-
sive Er atosthenes Seamount, 120 km in diameter at the
base, and extending fr om the seafloor to within 800 m
of the sea sur face. Dir ectly adjacent to the Er atosthenes
Seamount is a deep ( appr oximately 2750 m) depr es-
sion, par t of the Her odotus abyssal plain.
Galil and Zibr owius ( 1998) obtained benthos samples
fr om the Er atosthenes Seamount and found that the
fauna living on the seamount was r ich and diver se, in-
cluding two scler actinian cor als ( Caryophyllia calveri,
Desmophyllumcristagalli) , and a lar ge ar r ay of other
inver tebr ates in higher densities than sites of compar a-
ble depth in the Levantine basin. Red shr imps of com-
mer cial inter est ( Aristaeomorpha foliacea, Aristeus
antennatusand Plesionika martia) wer e also caught
by the beam tr awl sampler .
Commer cial tr awling on seamounts, pr imar ily tar geting
the or ange r oughy in the South Pacific, has consider able
impact on the benthos of these communities ( Koslow et
al., 2000) .

Se amo unts are s ubme rge d mo untai ns that
ri s e 1 0 0 0 m o r mo re abo ve the s urro undi ng
s e afl o o r and pro vi de uni que habi tats wi th a
wi de array o f i nve rte brate s , i ncl udi ng i nte r-
e s ti ng co ral s ( Caryophyllia calveri, Desmo-
phyllum cristagalli) . The bi o l o gi cal co mmu-
ni ti e s o f s e amo unts are thre ate ne d by fi s hi ng
i n s o me are as o f the wo rl d ( e . g. s o uth Paci fi c
s e amo unts ) and the s e fragi l e co mmuni ti e s re -
qui re urge nt pro te cti o n.
Cr edits:
Sar dou O. and Mascle J., 2003. Car togr aphie par sondeur multifais-
ceaux du Delta sous mar in du Nil et des domaines voisins. Publica-
tion spciale CIESM/ Gosciences-Azur , sr ie Car te et Atlas.
Loubr ieu B. and Satr a C., 2001. Car togr aphie par sondeur multifais-
ceaux de la Ride Mditer r anenne et des domaines voisins. Comit
Franais deCartographie, n 168, pp. 15-21.
Cr edit:
Loubr ieu B. and Satr a C.,
2001.
Se e co l o ur pl ate ,
p. 6 3 .
35
5. Anthropogenic impact
in the deep Mediterranean
5.1. Fisheries
Deep-sea fishing
1
is a r elatively new phenomenon, and
expanding in par t of the wor ld. In the Wester n Medi-
ter r anean it has become r elatively impor tant since the
1940-50s due to the high commer cial value of deep sea
Ar isteid shr imps ( mainly Aristeus antennatusand Aris-
taeomorpha foliacea) .
Most oceanic fisher ies have been concentr ated in the
upper r egions of the oceans, either on the continental
shelves for demer sal fish, or the epipelagic r egions for
fish such as tuna, billfish or shar ks ( Haedr ich, 1996) .
Now ther e is a pr onounced shift of fisher ies fr om shal-
low to much deeper r egions ( Hopper , 1995, Mer r ett
and Haedr ich, 1997) , especially in the demer sal fisher -
ies, motivated by the gr owing number of collapsed fish
stocks on the continental shelves, and in the Mediter -
r anean, by the high value of deep-sea ar isteid shr imps.
Globally, lar ge fish appear near the bottom in the deep
sea; in the Mediter r anean, fish ar e complemented by
lar ge shoals of ar isteid shr imps that make a tr adition-
al ( since the 1940s) deep-sea r esour ce in many ar eas.
Deep-sea fisher ies now occur in par ts of the wor ld down
to 1800 m depth ( Haedr ich, 1996) . However , due to
the slow gr owth of typical deep-sea fishes, it is unlikely
that demer sal deep-sea fisher ies conducted by bottom
tr awl will ever be impor tant in the long ter m because
r eplacement r ates ar e higher than cur r ent har vest r ates
( Haedr ich, 1996) . Once a deep-sea stock is depleted, it
will not r etur n within a r easonable time span ( Haedr ich,
1996, Lack et al., 2003; Koslow et al., 2000) . Sever -
al deep-water stocks ar e heavily exploited, or have col-
lapsed, in the wor lds oceans: r edfish ( Sebastesspp.)
and gr enadier ( Coryphaenoides rupestris) stocks in
the Nor th Atlantic; or ange r oughy ( Hoplostethus atlan-
ticus) in the South Pacific.
Cur r ently, the Mediter r anean fisher ies below 200 m
depth tar get decapod cr ustacean r esour ces. In the
upper slope ( down to ca. 500 m) , the deep-water
shr imp Parapenaeus longirostrisand the Nor way lob-
ster Nephrops norvegicusr epr esent impor tant fisher ies
in cer tain ar eas. These fisher ies have impor tant quanti-
ties of other commer cial species, such as Merluccius
merluccius, Micromesistius poutassou, Conger con-
ger, Phycis blennoidesand, to a lesser extent, monk-
fish ( Lophiusspp.) and the cephalopod Todarodes sag-
ittatus. The by-catch of other decapod cr ustaceans is
incr easingly commer cialised: glass shr imp Pasipahea
spp., Acanthephyra eximia, Plesionikaspp., Geryon
longipes, Paromola cuvieri.
Deeper fisher ies ( appr ox. 400 m to 800 m) tar get al-
most exclusively ar isteid shr imps ( Aristaeomorpha fo-
liaceaand Aristeus antennatus) , though some hake is
also har vested by tr awler s and bottom longliner s. Other
deep sea fisher ies exist in the Mediter r anean, but on a
smaller scale: longliner s tar geting hake in the Gulf of Li-
ons and the Italian Ionian sea, and longlines tar geting
the deep-sea shar k Hexanchus griseusin the souther n
Aegean sea ( 600-1500 m, Sar d et al., 2004a) .
In the Gr eek Ionian sea, A. foliaceais mor e abundant
than A antennatus. However , deep-water fisher ies
( > 500 m) ar e not yet well-developed in Gr eece ( Mytili-
neou and Politou, 1997; Politou et al. 2003) .
Even with a r elatively shor t histor y of fishing, r ed shr imp
stocks ar e alr eady showing symptoms of over exploita-
tion. Some A. antennatusstocks have collapsed in r e-
cent decades ( late 1970s - ear ly 1980s in Ligur ia, Or si
Relini and Relini, 1988) , or show signs of over exploita-
tion ( Car bonell et al., 1999) , while some stocks ( Demes-
Hexanchus griseus ( Bonnater r e, 1788)
Notidanus griseus Cuv.
(Hexanchus cinereus Risso) . Moungegris.V. Fossat, 1879.
Coll. Musum dHistoir e natur elle de Nice. Se e co l o ur pl ate , p. 5 8 .
1
Ther e is no accepted definition of what constitutes a deep-sea fish-
er y. The Deep Sea 2003confer ence consider ed deep-sea fisher ies as
those oper ating beyond the continental shelf br eak ( i.e. deeper than
200 m, Lack et al., 2003) . The ICES ( ICES, 2003) has defined deep-
sea fisher ies as those deeper than 400 m, while Koslow et al. ( 2000)
consider deep-sea fisher ies as those occur r ing deeper than 500 m.
In this document, although we have gener ally r epor ted infor mation
available fr om 200 m, we will focus on deep-water fisher ies deeper
than 400 m, i.e. following the ICES defintion.
36
tr e and Lleonar t, 1993; Bianchini and Ragonese, 1994)
seem to be under exploited to fully exploited. A. foliacea
has decr eased significantly fr om the commer cial catch-
es in many ar eas ( Gulf of Lions: Campillo, 1994; Catalan
sea: Bas et al., 2003, Tyr r henian sea: Fior entino et al.,
1998) and is consider ed over exploited in Italian water s
( Matar r ese et al., 1997; DOnghia et al., 1998) .
The potential fishing inter est of the cur r ently unexploit-
ed bottoms below 1000 m depth in the Wester n Medi-
ter r anean is ver y limited. This is so because the over all
abundance of cr ustacean species is consider ably lower ,
and fish communities ar e lar gely dominated by fish ei-
ther of non-commer cial inter est ( like the smooth head
Alepocephalus rostratus) or of a small size ( such as the
Mediter r anean gr enadier Coryphenoides guentheri) . If
these species ever become of economic inter est, the ec-
osystem effects of fishing could be ver y impor tant. Given
the impor tance of depths below 1000 m for the juveniles
of r ed shr imp ( s e e Bo x 9 ) and for the r epr oduction
of many fish species, the exploitation of these bottoms
would pr obably entail negative impacts on shallower
ecosystems, beyond the r apid depletion of par ticular ly
vulner able deep-sea megafauna communities.
In their r eview, Koslow et al. ( 2000) pointed out that
deep sea fishes follow a highly conser vative ecological
str ategy; the low fecundity and the low metabolic r ates
in a stable envir onment like the deep sea imply a high
vulner ability for their populations.
De e p- wate r e co s ys te ms are hi ghl y vul ne rabl e
to co mme rci al e xpl o i tati o n due to the l o w
turno ve r rate s o f the s pe ci e s adapte d to the s e
e nvi ro nme nts .
Co mme rci al e xpl o i tati o n bas e d o n de e p- s e a
trawl i ng has be co me i ncre as i ngl y i mpo rtant
s i nce the 1 9 5 0 s i n the Me di te rrane an, tar-
ge ti ng de e p- wate r s hri mp s pe ci e s ( Aristeus
antennatus and Aristaeomorpha foliacea)
do wn to 1 0 0 0 m de pth.
Trawl i ng has an e xtre me l y i mpo rtant di re ct
i mpact o n s e a- bo tto ms , as de mo ns trate d i n
co nti ne ntal s he l ve s thro ugho ut the wo rl d, o r
i n the s o uth Paci fi c o range ro ughy s e amo unt
fi s he ri e s .
Gi ve n the s tate o f s hal l o w wate r fi s he ri e s ,
whe re mo s t s to cks are ful l y to o ve r- e xpl o i te d,
and the hi gh e co no mi c val ue o f de e p- wate r
s hri mps , i ncre as i ng pre s s ure to fi s h i n de e p
wate r i s to be e xpe cte d i n the ne ar future .
5.2. Other anthropogenic threats
to the deep-sea Mediterranean
Anthr opogenic thr eats ar e not limited to fishing. Gr assle
( 1991) identified other sour ces of man-mediated im-
pacts that may thr eaten the conser vation of the diver -
sity, str uctur e and functioning of deep-water ecosystems.
Among these, we can cite waste disposal ( solid tr ash and
other toxic compounds) , pollution ( Haedr ich, 1996) ,
oil explor ation/ pipelines, or , mor e indir ectly, climate
change ( Danovar o et al., 2001) .
Kr ess et al., ( 1993) r epor ted the r esults of a monitor -
ing study of the disposal of coal fly ash in the Easter n
Mediter r anean sea. The coal fly ash was dumped 70 km
offshor e fr om the coast of Isr ael, at 1400 m depth. A
compar ison of the benthic fauna at the centr e of the dis-
posal site with that of a contr ol ar ea indicated a sever e
impover ishment of the benthos in the affected ar ea.
Dur ing a monthly cr uise of the Meteor in the Easter n
Mediter r anean in 1993 between 194 and 4614 m depth,
the litter r etained in a beam tr awl net included solid
waste such as plastic and glass bottles, metal cans, nylon
r ope and plastic sheeting ( Galil et al., 1995) . Although
the disposal of all litter ( except food waste) is pr ohib-
ited in the Mediter r anean, the study pr esented evidence
that this r egulation is r outinely ignor ed: 70% of the tr awl
hauls contained litter ( Galil et al., 1995) . Refuse gener -
ated by vessels is a major sour ce of litter in the Mediter -
r anean.
Toxicological studies have found that PCB levels in deep-
water fishes ( Alepocephalus rostratus, Bathypterois
mediterraneus, Coryphaenoides guentheri and Lepid-
ion lepidion) wer e lower than in coastal fishes, close to
Tr ash r ecover ed fr om 2000 m depth in the Easter n Mediter r anean.
37
the pollution sour ces, but much higher than in shelf to
upper slope fishes ( Micromesistius poutassou, Phycis
blennoidesand Lepidorhombus boscii) and within the
same r ange as a top pr edator , such as Thunnus thyn-
nus( Fig. 7, Por te et al., 2000; Sol et al., 2001) . The
levels of TPT ( tr iphenyltin) wer e higher in two bathyal
species ( Mora moroand Lepidion lepidion) than in
bivalves and fishes inhabiting har bour s and the coastal
ar ea ( Fig. 8, Bor ghi and Por te, 2002) . These r esults
point to a differ ential accumulation of PCBs and TPTs
by deep-water fishes, suggesting that bathyal and abyssal
food chains may alr eady be affected by human activities
on land.
Box 8. Increasing catches,
diminishing catch rates
The r epor ted landings of ar isteid shr imps in the
Mediter r anean have steadily incr eased over the last
decade, with a maximum of 6,637 t in 2000 ( GFCM
captur e pr oduction 1970-2002) . However , a detailed
analysis of the r ed shr imp ( Aristeus antennatus)
fisher y, conducted by Bas et al. ( 2003) in the por t
of Blanes ( Catalonia) , showed that catch r ates ( in
kg/ HP) have decr eased dr amatically between the
beginning of the fisher y in the 1950s and the pr esent
( 1997-2000) : fr om ar ound 0.4 kg/ HP to ar ound
0.04 kg/ HP. The catches ar e incr easing, but at a ver y
high ener getic cost: the engine power r equir ed to
pr oduce one unit of r ed shr imp is ten times what
it was 50 year s ago. The high pr ices fetched by
r ed shr imps ( typically above 30 / kg on fir st sale)
explain the incr eased effor t towar ds this deep-water
living r esour ce, r aising concer n about its long-ter m
sustainability.
N. oaroatus
N. galloprovincialis
R. decussata
Haroors
Coastal areas
Deep-sea areas
l. aurata
B. orandaris
N. moro
T. haemastoma
A. rostratus
B. mediterraneus
The deep-sea fish st died o the scientists at 8pain's lnstit te of Chemical and
Enironmental Research sho that TPT is persisting in the enironment and
s oject to long-range transport. The leels of PCBs, dioins, and TPT in these
deap-see fish are loer tha the leels of those contaminants fo nd in organisms
inhaoiting haroors and coastal areas, o t the TPT leels are m ch higher.
C. guentheri
l. lepidion
TPT |e e|s |o orgao|sms Irom he oor hes Ned| erraoeao
oglg .. as So
O 2O 4O OO 8O 1OO 12O 14O 1OO 18O 148O
Coastal fishes.
close to poll tion
so rce
8helf/slope fishes.
far from so rce
Top predator
Bathal fishes
Nar. Ecol. Prog. 8er. 2OOO, 192. 259-2OO, Deep-8ea Res. 2OO1, 48(2j. 495-518
N. oaroatus
8. caorilla
8. porcus
N. poutassou
l. crocodilus
P. olennoides
l. ooscii
T. th nus
A. rostratus
B. mediterraneus
C. guentheri
l. lepidion
O 25 5O 75 1OO
S
p
e
c
|
e
s
P08s (oglg ..)
Fi g. 7 . Concentr ation of PCB in fish tissue ( fr om Por te et al., 2000;
Sol et al., 2001) .
O
1OOO
2OOO
8OOO
4OOO
5OOO
OOOO
7OOO
1
9
5
O
1
9
5
9
1
9
O
2
1
9
O
5
1
9
O
8
1
9
7
1
1
9
7
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Human thre ats to the de e p s e a i ncl ude was te
di s po s al o f che mi cal and s o l i d was te , che mi cal
po l l uti o n by l and- bas e d acti vi ti e s and the l o ng-
rangi ng e ffe ct o f cl i mate change .
Fi g 8 . Tr iphenyltin concentr ation in Mediter r anean fish ( fr om
Bor ghi and Por te 2002) .
The effect of land-based human activity on deep-sea
ecosystems has a long histor y in the Mediter r anean,
and dates to pr e-industr ial times if we accept the sug-
gestion by Pr s ( 1985) that one explanation for the
often encounter ed sub-fossil white cor al assemblages
( cover ed by a fine layer of sediment) is the pr ogr essive
for est destr uction by man since the times of the ear ly
Mediter r anean civilisations.
38
Box 9. An important deep-sea living resource in the Mediterranean,
the red shrimp Aristeus antennatus
A. antennatusis pr esent in the entir e Mediter r anean
sea ( except the Adr iatic and the Black sea) and the At-
lantic fr om the nor th Iber ian peninsula to Angola. It is
found, locally, fr om 100-200 m depth ( Alger ia, Italy,
Egypt) to 3300 m ( Sar d et al., 2003a) . However , its
abundance is high only fr om 600 to 800 m, wher e the
fisher y takes place.
The gr owth r ates of A. antennatus( k r anging fr om
0.25 to 0.3 yr
-1
, Demestr e and Mar tn, 1993) ar e much
lower than those of other penaeids ( kbetween 1.8-3.6
yr
-1
) . Estimates of natur al mor tality ( M) ar e also much
lower , ar ound 0.5 yr
-1
compar ed to M= 1-4 yr
-1
in other
penaeids ( Demestr e and Mar tn, 1993) . Hence, it is a
typical deep-sea species of low pr oductivity.
The population compr ises a high pr opor tion of adult
females at depths shallower than 1000 m, with high
density. Females for m aggr egations in spr ing and sum-
mer . Juveniles and males ar e abundant below 1000 m
depth, with low population density. The smallest juve-
niles ( CL < 15 mm) ar e r ecr uited almost exclusively
below 1000 m, pr obably after ca.1 year of lar val / post-
lar val development ( Car tes and Demestr e, 2003) . Ju-
veniles ar e pr esent, seasonally, in submar ine canyons.
Sar d et al. ( 1994) r epor ted that the stock of Aris-
teus antennatus in the NW Mediter r anean appear s
to r emain constant at appr oximately optimum levels
of exploitation because par t of it is unexploited below
1000 m depth. The study pr ovided a r ationale for inter -
pr eting the seasonal catch fluctuations obser ved in the
r ed shr imp fisher y. Females contr ibute to most of the
catches shallower than 1000 m thr oughout the year ,
and the catch levels var y seasonally. Catch of males var -
ied seasonally and by depth. Juveniles wer e pr esent in
catches fr om autumn to spr ing, and var ied significant-
ly by depth and season. The r ole of local topogr aphic
featur es, viz. submar ine canyons, in the r ecr uitment of
this species was impor tant.
The fisher y shows impor tant fluctuations over the
shor t and mid-ter m ( cycles of minimum catches ever y
8-10 year s, Car bonell et al., 1999) and seasonal fluc-
tuations ( summer fishing occur s usually in deeper wa-
ter s and in autumn-winter ar ound 400 m) . Addition-
ally, a gener al decr ease in catches in the Spanish Medi-
ter r anean since the ear ly 1950s is noted: fr om a peak
of ~ 900 t in 1953 to 200-400 t in the 1980s ( 350 t in
Demestr e and Mar tn, 1993) . This decr easing tr end in
the catches has been accompanied by i) an impor tant
incr ease in the engine power of the tr awler s involved
in the fisher y; and ii) an impor tant incr ease in the eco-
nomic value of this species.
The evolution of exploitation in the Balear ic Islands
shows an over all incr easing tr end ( Car bonell et al.,
1999) , with a per iod of ver y low catches at the begin-
ning of the 1980s. This decr ease in catches coincided
with that obser ved in other shr imp fishing gr ounds in
the wester n Mediter r anean ( Ligur ian Sea, Gulf of Li-
ons) . Fr om 1983 a r ecover y in catches began, and in
r ecent year s the annual catch has stabilised at a level
similar to that attained dur ing the 1970s.
Landings of this r esour ce showed high fluctuations
in all Italian Seas. In the Ligur ian Sea, A. antennatus
was an impor tant r esour ce until 1979. In the per iod
1976-1980 the population decr eased pr ogr essively un-
til commer cial fishing in the ar ea was suspended. As of
1987 the catches incr eased and, at pr esent, the quan-
tities ar e of commer cial inter est, although ver y var i-
able fr om year to year ( Relini and Or si Relini, 1987;
Or si Relini and Relini, 1994) . Or si Relini and Relini
( 1985) for mulated some hypotheses to explain this
var iability, such as envir onmental decay, over fishing,
pathologic causes, hydr ological changes, and failur e
of r ecr uitment due to pr edation upon juveniles. In the
Ionian Sea, absence of A. antennatuswas fr equently
obser ved in concomitance with high catches of Micro-
mesistius poutassouand Phycis blennoides ( Matar -
r ese et al., 1997) .
Many autho rs co i nci de i n s tre s s i ng that the ab-
s e nce o f fi s hi ng be l o w 1 0 0 0 m he l ps e xpl ai n the
hi gh l e ve l s o f e xpl o i tati o n s us tai ne d by the s pe -
ci e s . Mo s t o f the po pul ati o n fi s he d are adul t fe -
mal e s , whi l e the mal e s and juve ni l e fracti o n o f
the po pul ati o n, l i vi ng de e pe r than 1 0 0 0 m, may
he l p re pl e ni s h the e xpl o i te d s to ck. Ho we ve r, an
i ncre as e i n the e xpl o i tati o n rate s o f adul t fe -
mal e s mi ght unde rmi ne the re pro ducti ve capa-
bi l i ti e s o f the s to ck. Addi ti o nal l y, e xpl o i ti ng the
juve ni l e fracti o n ( de e pe r than 1 0 0 0 m) mi ght
l e ad to re crui tme nt o ve rfi s hi ng i n the future .
39
A PROPOSAL FOR THEI R CONSERVATI ON
Part 2
A p ro p o sa l fo r th e ir c o n se rva tio n
S e rg i Tu d e la
1
, F ra n o is S im a rd
2
, J a m ie S k in n e r
2
a n d P a o lo G u g lie lm i
1

1
WWF Mediterranean Programme Office
Via Po, 25/c, 00198 Rome, Italy
2
IUCN Centre for Mediterranean Cooperation
Parque Tecnolgico de Andaluca, Calle Mara Curie, 35,
Campanillas, 29590 Mlaga, Spain
The first part of this document presents that scientific information which is currently avail-
able to guide policy and decision makers in deep-sea-related issues. To translate this into ac-
tion regardingthe management of anthropogenic activities in the various sites, it is impor-
tant to understand the current legal situation with respect to these areas, as well as to con-
sider the international policy context and the current commitments of the UNand of partners
to the relevant international conventions. Part 2 seeks to describe this background, and then
to make proposals for conservation action.
40
6. Deep-water habitat protection
and fisheries management
6.1. The particular legal status
of Mediterranean waters
Unlike in other r egions of the wor ld, Mediter r anean
coastal states have gener ally r enounced their r ight to
extend national jur isdiction acr oss tr ue 200-mile wide
Economic Exclusive Zones ( EEZs) as pr ovided for by
the United Nations Convention on the Law of the Sea
( UNCLOS) . The semi-enclosed natur e of the Mediter r a-
nean and the lar ge number of coastal states explain this
cautious appr oach, aimed at avoiding ter r itor ial fr ic-
tions.
However , in r ecent year s, some countr ies have tr ied to
adopt tailor -made, sui generis solutions to enlar ge their
fisher ies jur isdiction beyond the 6-12 mile ter r itor ial
water s, whilst avoiding a for mal EEZ declar ation. This
would be the case for the so-called fishing zone ( zone
de pche rserve) in Alger ia ( 1994) or the fisher ies
pr otection zones unilater ally declar ed by Spain ( 1997)
and Cr oatia ( ecological and fishing pr otection zone;
2003) . Malta, in tur n, has had its 25-mile management
zone r ecognized after its r ecent accession to the Eur o-
pean Union. The final declar ation of the Minister ial Con-
fer ence on the Sustainable Development of Fisher ies in
the Mediter r anean, held in Venice in November 2003,
acknowledges the beneficial r ole to be played by extend-
ing fisher ies pr otection zones for fisher ies management
in the r egion, and calls for the need to follow a concer t-
ed appr oach in their declar ation.
As for unilater al extensions of jur isdiction for pur pos-
es other than fishing, Fr ance has enacted a law in Apr il
2004 cr eating an ecological pr otection zone in the Med-
iter r anean beyond its ter r itor ial water s, mainly aimed
at implementing contr ols to fight pollution. Also, Italian
legislation pr ovides for the establishment of zones of bi-
ological pr otection beyond the Italian ter r itor ial sea on
the high seas; use of this pr ovision was made to cr eate a
zone of biological pr otection cover ing a str etch of water
in the vicinity of the island of Lampedusa.
Never theless, in spite of the occasional attempts by some
countr ies to par tly apply UNCLOS pr ovisions r efer r ed to
above, about 80% of the Mediter r anean still lies in the
High Seas, which poses specific pr oblems for the gov-
er nance of fisher ies management and biodiver sity con-
ser vation in this r egion. This doesnt mean, however ,
that ther e ar e no legal instr uments in place allowing for
the pr otection of the Mediter r anean deep-sea biodiver -
sity beyond national jur isdiction. In this r egar d, it must
be pointed out that the par ticular situation discussed
above r elates to the lack of EEZs, which mostly deal
with the use of biological r esour ces found within the
water column. Indeed, the use of those found on the
seabed beyond national jur isdiction is specifically cov-
er ed by the Inter national Seabed Author ity ( ISA) , which
establishes that coastal states have exclusive r ights over
seabed r esour ces on the continental shelf, a jur idical
concept that encompasses all the seafloor in the Med-
iter r anean basin. Sedentar y species, legally defined in
Ar ticle 77 of UNCLOS, ar e fully subject to this legisla-
tion. Also, the Bar celona Convention Pr otocol r elative to
Specially Pr otected Ar eas and Biological Diver sity in the
Mediter r anean ( SPA Pr otocol) pr ovides for the cr eation
of mar ine pr otected ar eas beyond ter r itor ial water s, as
exemplified by the Ligur ian Sea Cetacean Sanctuar y. This
pr otected ar ea was or iginally established in Mediter r a-
nean High Seas by the gover nments of Fr ance, Monaco
and Italy, being then listed as a Specially Pr otected Ar ea
of Mediter r anean Impor tance ( SPAMI) under the Bar -
celona Convention.
6.2. International policy context
The Plan of Implementation r esulting fr om the Wor ld
Summit on Sustainable Development ( WSSD) held in
Johannesbur g in 2002 includes, among other commit-
ments, the maintenance of the pr oductivity and biodi-
ver sity of impor tant and vulner able mar ine and coast-
al ar eas, including ar eas within and beyond national
jurisdiction; the establishment by 2012 of r epr esent-
ative networ ks of mar ine pr otected ar eas; and the de-
velopment of national, r egional and inter national pr o-
gr ammes for halting the loss of mar ine biodiver sity.
Decision VII/ 5, adopted by the 7
th
Confer ence of the Par -
ties to the Convention on Biological Diver sity ( CBD COP-
7) , in Kuala Lumpur in Febr uar y 2004, endor sed the
Johannesbur g Plan of Implementation mentioned above.
Indeed, it set up a biodiver sity management fr amewor k
to be adopted by Par ties ( appendix 3 to annex I) that
aims to deliver on a set of pr eviously agr eed objectives,
among them:
- To enhance the conservation and sustainable use
of biological diversity of marine livingresources
in areas beyond the limits of national jurisdic-
tion.
This entails a) to identify threats to biological di-
versity in areas beyond national jurisdiction, in
41
particular areas with seamounts, hydrothermal
vents and cold-waters corals, and certain other
underwater features;and b) to urgently take the
necessary measures to eliminate/avoid destruc-
tive practices, consistent with international law,
on a scientific basis, includingthe application of
precaution, for example, consideration, on a case
by case basis, of interimprohibition of destructive
practices adversely impactingthe marine biologi-
cal diversity associated with marine areas beyond
the limits of national jurisdiction, in particular
areas with seamounts, hydrothermal vents, and
cold-water corals, other vulnerable ecosystems
and certain other underwater features.
- To establish and strengthen national and region-
al systems of marine and coastal protected areas
integrated into a global network and as a contri-
bution to globally agreed goals.
This objective includes the setting up of r epr esenta-
tive mar ine pr otected ar eas wher e extr active uses
ar e excluded, and other human impacts ar e mini-
mized or r emoved, to enable the integr ity, str uctur e
and functioning of ecosystems to be maintained or
r ecover ed.
The Decision adopted by the CBD COP-7 includes a call
on the utilization of the pr ecautionar y and ecosystem
appr oaches when addr essing the conser vation of bio-
logical diver sity beyond national jur isdiction, and pr o-
poses a gener al two-level appr oach for conser vation in
which the marine and coastal protected areas net-
work would be sittingwithin a framework of sustain-
able-management practices over the wider marine
and coastal environment. This means combining a
site-based appr oach ( networ ks of pr otected ar eas) with
gener al r estr ictions that would apply to the entir e ar ea
( e.gmeasures to eliminate/avoid destructive prac-
tices, consistent with international law, on scientif-
ic basis, includingthe application of precaution, for
example, consideration on a case by case basis, of
interimprohibition of destructive practices CBD De-
cision VII/ 5-60) .
Resolution 58/ 240 of the United Nations Gener al Assem-
bly, appr oved ear lier in December 2003, alr eady called
for the ur gent management of r isks to the mar ine biodi-
ver sity of seamounts, cold water s cor al r eefs and cer tain
other under water featur es, and invited the r elevant r e-
gions and r egional bodies to addr ess the conser vation of
vulner able and thr eatened mar ine ecosystems and bio-
diver sity in ar eas beyond national jur isdiction. It also r e-
affir med the WSSD commitment to establish r epr esent-
ative networ ks of mar ine pr otected ar eas by 2012. In
addition, it r ecommended that the fifth meeting of the
Open-ended Infor mal Consultative Pr ocess on Oceans
and the Law of the Sea ( UNICPOLOS) , scheduled in June
2004, included in its agenda the conservation and
management of the biological diversity of the seabed
in areas beyond national jurisdiction.
UNICPOLOS, in tur n, welcomed CBD decision VII/ 5 on
mar ine and coastal biodiver sity and encour aged the
adoption of r estr ictive measur es on a r egional basis,
and within existing r egional fisher ies management or -
ganizations. It r eaffir med the concer n over the ineffec-
tive conser vation and management of seabed biodiver -
sity beyond national jur isdiction and, among other as-
pects, pr oposed to the UN Gener al Assembly ( UN GA)
to encour age Regional Fisher ies Management Or gani-
zations ( RFMOs) with a mandate to r egulate deep sea
bottom fisher ies to addr ess the impact of bottom tr awl-
ing on vulner able mar ine ecosystems, as well as to ur ge
States, either by themselves or though RFMOs, to consid-
er on a case-by-case basis, including the application of
pr ecaution, the inter im pr ohibition of destr uctive fish-
ing pr actices that have an adver se impact on vulner a-
ble mar ine ecosystems in ar eas beyond national jur is-
diction, including hydr other mal vents, cold water cor als
and seamounts.
Dur ing the meeting, delegates fr om sever al countr ies,
along with NGOs, suppor ted a mor ator ium on bottom
tr awling in the High Seas. A few months befor e, in August
2003, deep-sea biologists fr om ar ound the wor ld met in
Coos Bay ( Or egon, USA) , and launched a statement of
concer n to the UN Gener al Assembly r ecommending im-
mediate measur es for the pr otection of biodiver sity of
the deep sea on the High Seas, as well as within ar eas of
national jur isdiction. Inter alia, consistent with the
precautionary approach, concer ned scientists:
- called on the UN GA to adopt a moratoriumon
deep-sea bottomtrawl fishingon the High Seas, ef-
fective immediately, and
- r ecommended the development of representa-
tive networks of [ deep-sea] marine protected ar-
eas (MPAs), as called for by the World Summit on
Sustainable Development and endorsed by the UN
General Assembly.
UNICPOLOS also welcomed decision VII/ 28 of CBD COP-
7 in the sense of explor ing options for cooper ation for
the establishment of MPAs beyond national jur isdiction,
consistent with inter national law and on the basis of the
best available scientific infor mation.
42
With r espect to the Mediter r anean basin, the issue of
the pr otection of the local deep-sea biodiver sity fr om
impacting fishing pr actices was dealt with in the 2004
meeting of the Sub-Committee on Mar ine Envir onment
and Ecosystems ( SCMEE) of the Scientific and Advisor y
Committee ( SAC) of the Gener al Fisher ies Commission
for the Mediter r anean ( GFCM) , the RFMO with the man-
date for the Mediter r anean r egion. The SCMEE conclud-
ed in its r epor t that:
Current scientific advice does not support any ex-
pansion in the range of depths at which fishing
takes place. There is a strongopinion fromsome
scientists fromthe NWMediterranean that fish-
ingat depths greater than 1000 mshould not take
place, based on a precautionary approach. The
Subcommittee recommends that SAC (the Scien-
tific and Advisory Committee of the GFCM) should
analyze the conservation benefits to be gained
fromsettinglimits to the depths at which fishing
takes place and balance these against the cost to
fishermen.
6.3. A conservation proposal
tailored to the Mediterranean
6 . 3 . 1 . O ve rvie w
Accor ding to the detailed analysis pr esented in this r e-
por t, the gener al two-level appr oach embedded in the
CBD outcomes summar ized above that combines a) the
inter im pr ohibition of destr uctive pr actices adver sely
impacting the mar ine biological diver sity ( a general ap-
proach) , with b) the establishment of national and r e-
gional systems of mar ine and coastal pr otected ar eas in-
tegr ated into a global networ k ( a site-based approach) ,
is fully valid also for the Mediter r anean r egion.
However , a successful str ategy for the pr otection and
conser vation of the Mediter r anean deep-sea biodiver -
sity, including the over ar ching ecosystems and the r e-
lated biological and ecological pr ocesses, should be de-
signed so as to take into account the r egional geogr aphi-
cal, socioeconomic and political specificities. Such spe-
cificities include the pr edominantly nar r ow continental
shelves, the vast expanses of High Seas, and the exist-
ence of r egional bodies with a mandate for mar ine bio-
diver sity conser vation and fisher ies management, such
as the Bar celona Convention, the GFCM and, to some
extent, the EU.
The two-level appr oach detailed her e was fir st pr esent-
ed for discussion to a specialized scientific audience on
the topic, by means of a pr esentation - and later debate
- to the r ound table Potential Mediter r anean Pr otected
ar eas in High Sea and Deep Sea, held on June 11th
2004 in Bar celona, within the fr amewor k of the 37th
CIESM Congr ess. The pr oposal r eceived wide suppor t
fr om the Mediter r anean scientific community, being
based ar ound the following elements:
-a gener al appr oach, based on pr eventing an exten-
sion of fishing pr actices beyond 1000 m depth as a
pr ecautionar y measur e, seeking the agr eement of
stakeholder s and implementing the CBD r ecommen-
dations; and
-a site based appr oach aiming at the cr eation of a Med-
iter r anean r epr esentative networ k of deep-sea pr o-
tected ar eas, including the following habitats: subma-
r ine canyons, deep-sea chemosynthetic communities
( associated to cold seeps in mud volcanoes) , cold-
water cor als, seamounts and deep-sea br ine pools.
Both elements ar e complementar y ( i.e. not all biologi-
cally valuable deep sea ecosystems ar e found below the
1000 m isobath, wher eas all communities found at those
deeper gr ounds ar e assumed to be poor ly r esilient and
vulner able) and should be developed in par allel, using
adequate policy instr uments.
We need to str ess that most of the unique deep sea en-
vir onments her e pr oposed as futur e MPAs in the Medi-
ter r anean have been discover ed only r ecently, enhanced
by the use of newly available techniques ( deep submer s-
ibles, ROVs) . A fur ther incr ease in our knowledge on
Mediter r anean deep-sea ecosystems, paying special at-
tention to their dynamics and to the influence of anthr o-
pogenic impacts on their functioning and str uctur e, is
necessar y for their effective management.
6 . 3 . 2 . C o n se rvin g d e e p se a e c o syste m s
in th e M e d ite rra n e a n :
E le m e n t 1 . I n te rim p re c a u tio n a ry
p ro h ib itio n o f d e e p -se a fish e rie s
> 1 0 0 0 m in d e p th
Rationale
As discussed ear lier , notwithstanding the potential ef-
fects of other anthr opogenic per tur bations, such as glo-
bal war ming and pollution, fishing constitutes the most
immediate, for eseeable shor t-ter m thr eat to Mediter r a-
nean deep-sea ecosystems. As happens thr oughout the
wor ld, the pr ogr essive depletion of tr aditional fishing
gr ounds, together with technological cr eep, is push-
43
ing fishing activities towar ds offshor e ar eas, on much
deeper gr ounds. Medium-scale bottom tr awler s ( some
with a desk length of bar ely 15 m) tar geting deep-sea
shr imps alr eady r each fishing gr ounds at a depth of 800
m in some Mediter r anean ar eas on a r egular basis ( de-
pending on the season) , and ar e incr easingly appr oach-
ing the 1000 m isobath, par ticular ly when shr imp avail-
ability is low. However , such fisher y has not yet affected
those impor tant fish communities occur r ing at a depth
of ar ound 1000-1500 m.
The scientific basis suppor ting this measur e has been
extensively discussed ear lier in this document, on the
basis of specific Mediter r anean studies. Indeed, deep-
sea biological communities have an intr insic higher vul-
ner ability to exter nal per tur bations ( which may be ex-
tr eme in the case of par ticular ly impacting fishing sys-
tems, such as bottom tr awling) . Also, the few fish r e-
sour ces abundant enough to suppor t commer cial fish-
er ies ther e in the Mediter r anean ar e cur r ently non-mar -
ketable species. Fur ther mor e, any eventual exploitation
of these populations would lead to a r apid depletion of
the stocks, would str ongly disr upt the par ticular tr ophic
webs ther e, and would cause a deep impact on the str uc-
tur e and functioning of Mediter r anean deep-sea ecosys-
tems, which have star ted being scientifically explor ed
at a significant scale only in r ecent decades. In addi-
tion, as scientific studies point out, a fur ther expansion
of fisher ies down the bathymetr ic r ange would thr eaten
the sustainability of cur r ent deep-sea fisher ies in the r e-
gion, namely those of deep-water ar isteid shr imps, for
which deeper gr ounds cur r ently play the r ole of natu-
r al r efuges and nur ser y gr ounds for the juvenile popu-
lation fr action.
Implementation
It is impor tant to point out that limiting the maximum
depth in the Mediter r anean susceptible to exploitation
by fishing to the isobath of 1000 m would be a pr ecau-
tionar y measur e suppor ted by sound scientific evidence,
in the line r ecommended by Decision VII/ 5 adopted by
CBD COP-7.
Fur ther mor e, the pr oposed deep-sea fishing limitation
should be consider ed mor e a r estr iction on potential
fishing development than a tr ue fishing ban, since no
r egular fisher ies ar e taking place at those depths at
pr esent. In this sense, it is impor tant that stakeholder s
fully under stand the r ationale suppor ting this measur e
that clear ly benefits the sustainability of fishing activity
and suppor t the pr oposal as well.
Accor ding to the par ticular legal status of Mediter r anean
water s discussed above, it appear s that the adoption and
implementation of this measur e should be done thr ough
a binding r esolution, aimed both at the domestic level of
coastal states ( including the Eur opean Union thr ough
the new r egulation on Mediter r anean fisher ies manage-
ment) , and also at the level of the r elevant Regional Fish-
er ies Management Or ganization, that is the Gener al Fish-
er ies Commission for the Mediter r anean.
6 . 3 . 3 . C o n se rvin g d e e p se a e c o syste m s
in th e M e d ite rra n e a n :
E le m e n t 2 .
R e p re se n ta tive n e two rk o f d e e p -se a
p ro te c te d a re a s
Rationale
As descr ibed in detail in this r epor t, ther e ar e a number
of specific habitats with par ticular biodiver sity impor -
tance in the Mediter r anean deep-sea. These include sub-
mar ine canyons, cold seeps associated to mud volcanoes
( har bour ing chemosynthetic communities) , cold water
cor al r eefs, seamounts and br ine pools. Also, ther e is
incr easing consensus on the impor tance of the abyssal
ar eas cur r ently poor ly studied to biodiver sity in the
r egion. Based on our pr esent knowledge, the locations
of these habitats ar e r epr esented in fig. 9, but this figur e
should not exclude pr otection of other unique sites of
similar char acter istics that we expect might be discov-
er ed in the futur e.
The implementation of a gener al limitation on deep-
sea fisher ies below the isobath of 1000 m as pr oposed
above would eliminate the r isk of a pr ofound anthr opic
impact due to the lar ge-scale r emoval of biomass fr om
the ecosystems, str ongly alter ing their str uctur e and
functioning, and would pr event the intr oduction of dam-
aging fishing techniques like bottom tr awling in sensitive
deep-sea envir onments. However , this gener al measur e
alone needs to be completed with a par allel site-based
appr oach to ensur ing adequate pr otection of the mor e
biologically valuable deep-sea Mediter r anean habitats.
Indeed, some of the habitats listed above lie totally or
par tially above the 1000 m isobath, as with Lophelia
r eefs in the Ionian Sea, some chemosynthetic commu-
nities, and submar ine canyons and seamounts. Besides,
even in those habitats fully cover ed by the 1000 m fishing
ban, fishing is not the only possible thr eat to be avoided
or alleviated: pollution, seafloor dr illing, etc. would also
mer it specific attention in any conser vation scheme.
44
For these r easons, the setting up of a r epr esentative
networ k of deep-sea pr otected ar eas, endowed with a
str ict management of major anthr opic impacts, should
be a second key element of a r egional policy for the
conser vation of deep-sea biodiver sity, in full agr eement
with decisions adopted at the CBD COP-7. Implementa-
tion of this measur e would cer tainly benefit fr om a wide
consensus among stakeholder s, including all potential
sector s with an influence in the Mediter r anean sea.
Submarine canyons
As descr ibed ear lier in this r epor t, submar ine canyons
ar e geological str uctur es key to biological and ecologi-
cal pr ocesses in the entir e Mediter r anean basin. Indeed,
canyons ar e cr ucial in channelling ener gy and matter
fr om coastal ar eas to the deep-sea, and by changing
the vor ticity of cur r ents cr eating local upwellings that
ar e cr ucial to the life-cycle biology of cer tain species,
including pelagic fish and mar ine mammals. Besides,
impor tant biocoenoses such as those linked to cold-wa-
ter cor als ar e associated with their flanks.
Canyons ar e also significant for Mediter r anean fisher ies,
being tightly linked to the life cycle of ar isteid shr imps,
natur al r epr oductive r efuges for over exploited demer sal
species, and impor tant spawning gr ounds for anchovy.
Due to the thr ee-dimensional impor tance of canyons, af-
fecting both benthic and pelagic biodiver sity, pr otection
should be affor ded at least to the entir e canyon system,
i.e. the sea floor and the entir e water column associated
with it.
Scientific liter atur e shows that canyons ar e r elevant to
Mediter r anean biodiver sity and biological pr ocesses all
ar ound the r egion, though they ar e especially impor tant
in the NW Mediter r anean ( Catalonia, Gulf of Lions and
Ligur ian coast) , as well as in cer tain par ts of the Easter n
Basin ( i.e. those associated with the Nile fan, in the Le-
vantine basin) . A complete and fully r epr esentative net-
wor k of deep-sea pr otected ar eas in the Region should
contain pr otected canyons systems fr om at least these
two ar eas.
Cold seeps (chemosynthetic communities)
Deep-sea cold seeps ar e of maximum ecological and bi-
odiver sity impor tance in the Mediter r anean since, in the
absence of deep-sea hydr other mal vents, cold seeps ar e
the only fully chemosynthetic ecosystems in the basin.
Candidate sites for pr otection include those cold seeps
wher e live chemosynthetic communities have been
identified to date: the Olympic field ( South of Cr ete) , the
Anaximander mountains ( South of Tur key) and a lim-
ited ar ea off Egypt and Gaza.
Munida rugosa.
Pr ince Alber t de Monaco. Camp. scient. Crustacs podopht. pl. VII.
J. Het del.
45
It is likely that fur ther r esear ch will demonstr ate the
existence of mor e live chemosynthetic communities in
other ar eas har bour ing mud volcanoes and cold seeps.
Accor dingly, the networ k of deep-sea pr otected ar eas
could, on a pr ecautionar y basis, also include some of
the ar eas wher e cold seeps have been detected, and the
pr esence of chemosynthetic communities is pr esumed.
Cold water coral reefs
Cold water cor als r eefs or mounds ar e of high biodi-
ver sity value in the Mediter r anean. Only few r elictual
colonies of the main cold water r eef-builder Lophelia
pertusar emain in the Mediter r anean, wher e it appear s
that cur r ent envir onmental conditions ar e consider ably
below the optimum for this species. Madrepora ocu-
latais another impor tant r eef-building species in the
Mediter r anean.
Cold water r eefs or mounds ar e biodiver sity hotspots,
since the associated thr ee-dimensional str uctur e pr o-
vides ecological niches to Mediter r anean deep-sea
species such as the Mediter r anean or ange r oughy ( Ho-
plostethus mediterraneus) and other s.
Obvious candidate sites for pr otection include the only
known live Lopheliamounds in the Mediter r anean the
site found 20-25 miles offshor e Santa Mar ia di Leuca
( Italy) , wher e this biocoenosis has been found at a
depth r ange of 425-1110 m and a few scatter ed ar eas
in the Albor an Sea, the Gulf of Lions and the Catalan Sea
( associated to canyons) wher e the species has been con-
fir med to occur . Other ar eas of impor tant Madrepora
oculata pr esence could also be included. Incr eased
pr ospection will pr obably lead to the discover y of mor e
live Lopheliacolonies in the Mediter r anean.
Seamounts
Seamounts ar e par ticular mar ine geomor phological
str uctur es that ar e consider ed of gr eat biodiver sity im-
por tance wor ldwide. In ar eas of the Indian and Pacific
oceans, par ticular ly, the discover y of significant bio-
masses of commer cial fish r esour ces occur r ing ar ound
the slopes of seamounts has given way in r ecent year s
to the development of new deep-sea fisher ies. These
have systematically r esulted in a str ong decline of fish
biomass after only a few year s of exploitation, due to
the ver y high vulner ability that the r elated deep-sea fish
communities face to commer cial exploitation.
Although the biodiver sity of Mediter r anean seamounts
has seldom been explor ed, the fir st benthos samples
r ecover ed fr om the top of the Er atosthenes Seamount,
in the Easter n Mediter r anean, r evealed a r elatively r ich
and diver se fauna that included the fir st occur r ences of
two scler actinian species in the Levant Sea, both of them
constituting the deepest r ecor ds in the whole Mediter -
r anean. This and other scatter ed evidence suggest that
Mediter r anean seamounts ar e not an exception in r ela-
tion to its biodiver sity impor tance.
Even if a gener al limitation on deep-sea fisher ies as
pr oposed ear lier could entail the par tial pr otection of
some Mediter r anean seamounts, in most cases the sum-
mit and upper slopes would r emain unpr otected faced
with potentially har mful fishing techniques such as bot-
tom tr awling. Consequently, a site-based appr oach for
the pr otection of Mediter r anean seamounts appear s to
be necessar y. In this sense, a Mediter r anean r epr esenta-
tive networ k of deep-sea pr otected ar eas should include
the best known Mediter r anean seamounts, that is, Er a-
tosthenes, as well as, on a pr ecautionar y basis, those of
pr esumed biodiver sity impor tance most thr eatened by
the potential development of anthr opic activities ( nota-
bly deep-sea fisher ies) .
Brine pools (deep hypersaline habitats)
Deep hyper saline str uctur es known as br ine pools ar e
deep-sea habitats of high biodiver sity impor tance, par -
ticular ly to extr emophilic bacter ia and metazoan meio-
faunal assemblages.
Repor ted sites in the Mediter r anean ar e r estr icted to
the Easter n Basin, at beds below 3000-m. All illustr ated
deep hyper saline anoxic basins should be included in a
Mediter r anean r epr esentative networ k of deep-sea pr o-
tected ar eas, to be pr eser ved fr om potential anthr opic
impacts, such as seafloor dr illing or waste disposal.
46
Implementation of a Mediterranean network
of deep-sea protected areas
Differ ent legal possibilities exist for the establishment of
deep-sea pr otected ar eas in the Mediter r anean, r anging
fr om national ( including EU) legislation, to inter nation-
al tr eaties with competence on the Mediter r anean High
Seas, either dealing with integr al conser vation aspects
such as the Bar celona Convention or r elating to a
par ticular sector such as GFCM on fisher ies.
Given this complex pictur e, dur ing the discussion held
at the CIESM thematic r ound table on the pr otection of
the Mediter r anean deep-sea in June 2004, a need was
clear ly identified by those exper ts pr esent. A coor dinat-
ing platfor m should be cr eated gather ing together all
r elevant tr eaties with mandates for the pr otection of the
Mediter r anean and the management of human activities
in this sea, to move towar ds a har monized appr oach for
the establishment of a r epr esentative networ k of deep-
sea pr otected ar eas in the Mediter r anean Sea.
This said, and given the usual slow pace of institutional
pr ocesses, the pr inciple outlined above should not pr e-
vent selected Mediter r anean deep-sea habitats, such as
those pr oposed in the sections above ( see map below) ,
alr eady being cr eated under the most appr opr iate legal
fr amewor k, to secur e immediate pr otection. In this
sense, it is wor th r efer r ing to the Ligur ian Sea Cetacean
Sanctuar y, cr eated in 1999 by Fr ance, Italy and Monaco
on the Mediter r anean High Seas, that was then desig-
nated as one of the fir st Specially Pr otected Ar eas of
Mediter r anean Impor tance ( SPAMI) under the Bar ce-
lona Convention Pr otocol r elative to Specially Pr otected
Ar eas and Biological Diver sity in the Mediter r anean.
Fi g. 9 . Pr esently known distr ibution of deep-sea unique biocenoses in the Mediter r anean and adjacent Atlantic water s.
Cr edit: Her mes ( Hotspot Ecosystem Resear ch on the Mar gins of Eur opean Seas) , VI FP Eur opean Commission Pr oject;
and An InteractiveGlobal Map of Sea Floor Topography Based on SatelliteAltimetry &Ship Depth Soundings.
Meghan Miller , Walter H.F. Smith, John Kuhn, & David T. Sandwell. NOAA Labor ator y for Satellite Altimetr y.
http:/ / ibis.gr dl.noaa.gov. Modified.
0|bra|tar
Stra|ght
S|c||y
0hanne|
0u|f
of L|ons
0u|f
of L|ons
Levant|ne Sea
Santa Nar|a
d| Leuca
0||mp| Nud Vo|canoe F|e|d
Anax|mander
Nounta|ns
h||e Fan
Fratosthenes
Seamount
A|boran Sea
8a|ear|c
|s|ands
Uran|a 8as|n
hapo|| 0ome
Coral
Cold seep sites
47
0ar|o Noods
2OO
2OO
2OO
1OOO
1OOO
1OOO
1OOO
5w
OOh
he r les from
28.O8.2OO4
C rrent emergenc
meas res
Rona ls.
Heorides
Areas here bo om ra||og
o|d be proh|b| ed
Nadeira
and the Canar lslands
Aores
Box 10. Community initiatives for the protection of deep sea biodiversity
in Atlantic waters
Recent initiatives under taken at Eur opean level have
ser iously addr essed the pr otection of fr agile deep-sea
habitats fr om tr awling in the NE Atlantic. In Mar ch
2004, the Eur opean Council agr eed to give per manent
pr otection to Scotlands unique cold-water cor al r eefs,
the Dar win mounds, by banning deep-water bottom
tr awling in the ar ea.
Only discover ed in 1998, the Dar win Mounds ( see
below) ar e a unique collection of cold-water cor al
mounds ( Lophelia pertusa) at a depth of 1000 m,
about 185km nor thwest of Scotland. They ar e made
up of hundr eds of cor al r eefs up to 5m high and 100m
wide cover ing an ar ea of appr oximately 100 sq km.
The r eefs suppor t a wide diver sity of mar ine life, such
as sponges, star fish, sea ur chins, cr abs and deep-sea
fish including the blue ling, r ound-nosed gr enadier
and or ange r oughy.
Ear lier , in Febr uar y 2004, the Eur opean Commission
tabled a pr oposal to ban the use of bottom-tr awled
fishing gear ar ound the Azor es, Madeir a and the Ca-
nar y Islands ( see map below) . The declar ed aim is to
eliminate the r isk of the lasting or ir r epar able damage
that such gear can cause to highly sensitive deep-water
habitats found in these ar eas. A r estr ictive access r e-
gime has pr evented until now the activity of deep-sea
tr awling in these ar eas. The continental shelf ar ound
the islands concer ned by the pr oposed measur es is
ver y nar r ow or vir tually non-existent. Sever al habitats
ar e to be found at the bottom of these deep water s.
These include deep-water cor al aggr egations, ther mal
vents and car bonate mounds, which give shelter and
food to a highly diver sified fauna and flor a. Scientif-
ic evidence, including r epor ts fr om the Inter national
Council for the Explor ation of the Sea ( ICES) , shows
that habitats such as those found ar ound the islands
concer ned by the pr oposal ar e in need of special pr o-
tection, especially against the physical damage caused
by bottom tr awls and similar fishing gear .
48
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55
GLOSSARY
Abys s al : Depth zone usually below 3000 m cor r espond-
ing to the abyssal plain ( down to 5000 m in the Medi-
ter r anean) .
Bathyal : Depth zone between 200 and 3000 m, cor r e-
sponding to the continental slope.
Bathy- pe l agi c: open water s between 1000 and 3000 m
depth, wher e no sunlight is detected.
Be nthi c Bo undary Laye r ( BBL) : the near -bottom r e-
gion ( ca. 50-100 m above the bottom) wher e an in-
cr ease in the r esuspended matter and the biomass of
living or ganisms is r ecor ded.
Be ntho s : Or ganisms living on( epifauna) or in( infau-
na) the sea-floor . Adj. benthic.
Be ntho pe l agi c: swimming or dr ifting animals of the
Benthic Boundar y Layer , which may spend var ying
amounts of time on the seabed. Applied to megafau-
na ( fishes and cr ustaceans) , the ter m is often synony-
mous with demer sal.
Bi o ce no s e : ecologically inter acting or ganisms living to-
gether in a specific habitat.
Che mo s ynthe ti c ( Che mo - auto tro phi c) : Or ganisms
( usually bacter ia) that pr oduce car bohydr ates fr om
simple inor ganic compounds, by a metabolic pr oc-
ess involving the oxidation of r educed or ganic mate-
r ial and chemical compounds as an ener gy sour ce;
as opposed to photosynthesis, wher e plants make en-
er gy to fix car bon fr om sunlight.
Co l d s e e p: locations wher e natur al gases and noxious
water s, with low oxygen and high sulphur contents,
emer ge fr om bur ied sediments, salt domes or petr o-
leum deposits.
De tri ti vo ry ( de tri ti vo re ) : feeding str ategy of or gan-
isms based on consumption of dead plants ( phyto-
plankton) and animal r emains.
Di s turbance : an event or cir cumstance that inter r upts
the usual r elationship between or ganism and envi-
r onment.
Endo s ymbi o thi c ( e ndo s ymbi o nts ) : or ganisms ( usu-
ally bacter ia) that live within the cells of another or -
ganism.
Epi pe l agi c: applied to the water layer fr om 0 to about
200 m depth wher e ther e is enough light for photo-
synthesis.
Eurybathi c: applied to fauna that live acr oss wide
depth r anges.
Fi l te r fe e di ng ( fi l te r fe e de rs ) : feeding str ategy of
those animals consuming small par ticulate or ganic
matter suspended in the water column.
Gui l d: gr oups of animals that shar e a common way of
life; e.g., feeding guilds gr oup of animals consuming
similar food r esour ces.
Hadal : ocean depths usually below 6000 m, cor r e-
sponding to the oceanic tr enches.
Hydro the rmal ve nt: Geyser s of the sea-floor , wher e
water laden with chemicals is expelled at ver y high
temper atur es. The chemical composition of the wa-
ter s allows the establishment of unique animal com-
munities, based on chemosynthesis.
Macro be ntho s : Benthic or ganisms lar ger than 0.5 mm.
Polychaetes, bivalves and gastr opods ar e typical fau-
nal gr oups of macr obenthos in the deep-sea.
Macro fauna: A ter m mainly employed for benthic or -
ganisms to define those animals sieved thr ough 0.3-1
mm mesh size ( usually 0.5 mm) . Dominated in the
deep-sea by Polychaetes and per acar id cr ustaceans,
and to a lesser extent, molluscs ( bivalves, gastr o-
pods) .
Glossary of key ecological concepts
56
Macro pl ankto n ( = macro z o o pl ankto n) : planktonic
or ganisms between 1-200 mm in size, usually col-
lected with plankton nets of a mesh size between 0.75
and 4 mm.
Me gafauna: lar ge fauna, they ar e obser ved by subma-
r ine photogr aphy or collected by bottom tr awls, e.g.
fishes, echinoder ms, decapod cr ustaceans and ce-
phalopods for the vagilemegafauna, and sponges,
jellyfish, sea-pens and deep-sea cor als for the ses-
silemegafauna.
Me i o fauna ( = me i o be ntho s ) : Fauna sieved thr ough
0.062 mm mesh, smaller than macr ofauna, inhab-
iting the inter stitial space of sediments. Dominated
by Nematodes in the deep-sea. Despite the fact that
lar ge, single-celled pr otozoans ( For aminifer ans) ar e
technically meiofauna they ar e excluded fr om meio-
fauna studies ( and the ter m metazoan meiofauna is
mor e appr opr iate) .
Me s o - bathype l agi c: open water s fr om 200 to about
1000 m depth. Although some light r eaches these
depths it is insufficient for photosynthesis. Typical or -
ganisms of the meso-bathypelagic zone include mac-
r o-zooplankton that car r y day-night migr ations to
sur face water s.
Me s o s cal e : r elated to ocenogr aphic pr ocesses at spa-
tial scales between 10 and 1000 km.
Mi cro ne kto n: compr ises the size fr action of nekton
immediately above or similar in size to that of mac-
r oplankton.
Ne kto n: The actively swimming pelagic fauna able to
move independently of water cur r ents, usually lar ger
than 20 mm.
Ne ri ti c: r efer r ed to or ganisms living in water s over lying
the continental shelf.
Ne phe l o i d l aye r: tur bid layer s of water s car r ying fine
r esuspended matter usually located close to the oce-
anic bottoms.
Ol i go tro phi c: habitats that ar e poor in nutr ients and
plant life, and usually r ich in oxygen.
Organi c Matte r: matter in the water column or the sea
floor der ived fr om living or ganisms.
Pe l agi c: par t of the sea compr ising the water column,
i.e. all except the coast and the sea floor .
Phyto de tri tus : detr itus gener ated by the fall of dead
phytoplankton on the sea floor .
Re s o urce parti ti o ni ng: Shar ing by or ganisms of the
food r esour ces available; i.e. differ ent species ( or
par ts of the life cycle of one species) specialize in dif-
fer ent par ts of the food r esour ce.
Se co ndary pro ducti o n: the amount of mass ( or
weight) pr oduced per ar ea and unit time by an or -
ganism, a population or an animal community.
Ste no hal i ne : applied to an or ganism living in r estr ict-
ed r anges of salinity.
Suprabe ntho s ( = hype rbe ntho s ) : The fr action of
benthopelagic fauna collected with plankton nets of
0.5-1 mm mesh size. Size between 1-20 mm. Mainly
composed of Per acar id cr ustaceans.
Sus pe ns i o n fe e di ng ( s us pe ns i o n fe e de rs ) : feed-
ing str ategy based on tr apping ( usually by specially
adapted or gans) par ticle-bound or ganic matter .
Tro phi c di ve rs i ty: the var iety of food items in the diet
of an or ganism.
Tro phi c l e ve l : Hier ar chy of an or ganism in the food
chain. Each step or level within a food chain.
57
PLATES
Fig. 1. Geogr aphy of the Mediter r anean sea.
Credits: International Bathymetric Chart of theMediterranean, published 1981 by theHead Department of Navigation
and Oceanography, St. Petersburg, Russia, on behalf of theInternational Oceanographic Commission (GEBCO, 2003).
Modified.
Fig. 2. Chlor ophyll amap ( Monthly composite for Apr. 2000) pr oduced by the Inland and Mar ine Water s Unit ( JRC-EC)
using SeaWiFS r aw data distr ibuted by NASA-GSFC.
Fig.9. Pr esently known distr ibution of deep-sea unique biocenoses in the Mediter r anean and adjacent Atlantic water s.
Cr edit: Her mes ( Hotspot Ecosystem Resear ch on the Mar gins of Eur opean Seas) , VI FP Eur opean Commission Pr oject;
and An InteractiveGlobal Map of Sea Floor Topography Based on SatelliteAltimetry &Ship Depth Soundings.
Meghan Miller, Walter H.F. Smith, John Kuhn, & David T. Sandwell. NOAA Labor ator y for Satellite Altimetr y.
http:/ / ibis.gr dl.noaa.gov. Modified.
0|bra|tar
Stra|ght
S|c||y
0hanne|
0u|f
of L|ons
0u|f
of L|ons
Levant|ne Sea
Santa Nar|a
d| Leuca
Anax|mander
Nounta|ns
h||e Fan
Fratosthenes
Seamount
A|boran Sea
8a|ear|c
|s|ands
Uran|a 8as|n
hapo|| 0ome
Coral
Cold seep sites
58
Lepidion lepidion( Risso, 1810) . Lota lepidionRisso, Moustella deFount Stocoe, V. Fossat, 1879. Scale 1/ 2.
Hexanchus griseus ( Bonnater r e, 1788) . Notidanus griseus Cuv. (Hexanchus cinereus Risso) . Moungegris.V. Fossat, 1879. Scale 1/ 4.
Alepocephalus rostratus Risso, 1820. Vincent Fossat, 1879. Scale 1/ 2.
Mora moro ( Risso, 1810) . Mota mediterranea, Mora. V. Fossat, 1878. Scale 1/ 2.
59
PLATES
Aristeomorpha folicea and Aristeus antennatus
Alber t I
er
Pr ince de Monaco. Camp. scient. Pnids pl. III.
EL. Bouvier del., M. Bor r el pinx.
Scale 3/ 4.
60
Bathymedon longirostris, Eusirus longipes, Lepechinella manco, Rachotropis grimaldii.
J. E. Car tes
O
1O
2O
8O
4O
5O
OO
7O
8O
9O
2OO 8OO 4OO 5OO OOO 7OO 8OO 9OO 1OOO 11OO 12OO 18OO 14OO 15OO 1OOO 17OO 19OO
depth (m)
k
g
l
h
Nora moro
Alepocephal s rostrat s
0ale s melastom s
0ther fishes
Nerl cci s merl cci s
Dalatias licha
Phcis olennoides
Centroscmn s coelolepis
uepidion lepidion
Relative abundance
of shes with depth,
extr apolated fr om data
fr om exper imental
samplings in the wester n
Mediter r anean fr om
1988-2001.
61
PLATES
Etmopterus spinax,
Paromola cuvieri,
Physeter catodon.
Dr awings by M. Wr tz.
Ar tescienza.
62
Cr edit: Coleman and Ballar d ( 2001) .
Desmophyllumcristagalli, Lophelia pertusa and Madrepora oculata.
Cr edit: Angelo Tur si.
Gaz Bubbles
63
PLATES
Fig. 6. Topogr aphy of a submar ine canyon.
Credits: GRC Geocincies Marines (Universitat deBarcelona) and RMR Institut deCincies del Mar (CSIC), modified.
Cr edits:
Sar dou O. and Mascle J., 2003. Car togr aphie par sondeur multifaisceaux du Delta sous mar in du Nil et des domaines voisins.
Publication spciale CIESM/ Gosciences-Azur, sr ie Car te et Atlas.
Loubr ieu B. and Satr a C., 2001. Car togr aphie par sondeur multifaisceaux de la Ride Mditer r anenne et des domaines voisins.
ComitFranais deCartographie, n 168, pp. 15-21.
64
0|bra|tar
Stra|ght
S|c||y
0hanne|
0u|f
of L|ons
0u|f
of L|ons
Levant|ne Sea
Santa Nar|a
d| Leuca
Anax|mander
Nounta|ns
h||e Fan
Fratosthenes
Seamount
A|boran Sea
8a|ear|c
|s|ands
Uran|a 8as|n
hapo|| 0ome
Coral
Cold seep sites

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