Professional Documents
Culture Documents
c. C. Emig
Ce ntre d'Ocean olo gie d e Mar se ille (UMR-CNRS 6540) , Station Mari ne d 'Endo um e, Rue de la Batterie-des-Lions.
130 07 Marse ille, France.
ABSTRACT
O n the con tine n tal slop e, th e bath yal can be divide d into two zones, the upper ba thya l and
the mid dl e bath yal, at the shelf break, wh ich represen ts th e bo un dary betwe en th e coastal shelf
envir on men t an d the dee p realm , located at about 100-110 m d ep th. T he up per bathyal, p revi-
o usly co nsidered a transi tional zo ne, is ch ar acteri sed by d istin ct p hysical, geological an d biol ogi-
cal features. Its bath ym en-ic ex ten sio n is dir ectly related to slope p hysiogra p hy, and its lower
bo un dary ge ne ra lly co rresp onds to th e m ud lin e. T h is bel t is gover ne d by sp ecific ab io tic factors
with stee p physical grad ien ts (e.g., hyd ro dynam ics, salin ity, oxyge n , temperat ure , sedirnen ts}.
Major ch ange in tbe ben thi c fau n a is associated with m ajor ch ange in these abi otic factor s. Th e
three main biocoeno ses ar e domin ated by suspen sion-feed in g speci es, which ar e exclusive to th e
Mediter ran ean upp er bat hyal. Dep endi ng on water para m eters, th e limit between th e p hytal and
aphyta l systems ge ne ra lly occu rs with in th e upp er bathyal. The seco n d zo n e, th e middl e bath yal,
is poorl y do cum en ted ; however, it ca n b e divided in to several subzones whos e limi ts ap pe ar to be
re lat ed to water-ma ss characteristics: th e u pper-m idd le slope , the lower-m iddl e slope , an d th e
lower slope . T he lower ba thyal zo ne deve lops into th e bath yal ri se an d plain .
Key words: Medi terranean Sea, con tin en tal slope, bathyal zo nation, bio logical oceanograp hy,
p hysical ocean og rap hy, geolog ical oce an ogra phy.
RESUMEN
L os z ona s btuiales del talud con tinental m edi ter nineo: un en say o
En el talsui continental, el -piso bauai se puede dividi·r en banal superior y batial medio. Sn timue supe-
ri ar (100-110 m) corresponds al reborde del talud, suuada entre dos domisuos muy distintos: el de la plala-
[orma y el prof undo. Aruerumnerue considerada coma «zona de tronsicum» entre etpiso circaluoral y et ba-
tial, el Ltuiol superior se caraaerua 1'01' sus particularidades fisicas, geol6gicas y biol6gi.cas. 511, extension
botimetrica dcpende de la Jisiografia deltalu d, qu e indu ce los efectos de los facnnes abuiticos y asi proooca
una disnibucum. del beruos en d auurones con dominan cia de Los suspensivoros }' la casi ausencia de sedi-
menuuion. El batial supenor comp rende generalmerue el limiie en tre el sistema fital y el ajtal segttn la trams-
parencia de las aguas. El batial medi a empiexa a pm·t.i., de la. lin ea de fa ngo (mud line) qu.e marca un (',(J.t1/.+
bio abrupto de la energia. Aunque faltan estudios daa llados, a partir de Los datos disponibles se pu ede
jJrojJoner un esqlJ,mna de la. umacuni del batiai media: el talud: medic superior, el talud media inferun ; et ta-
lud. inf erior. Por tanto, el boliol inf erior se exuende en la llanu ra batial.
Palabras cla v e: M ar M editemin eo, talud continental, zonacum. batial, oceanografia biologica,
oceanografiafisica; oceanografia geol6gica.
23
c. C. Emig Meduerm nean baih)'at zones
Profile Type I
Profile Type II
I
~
il
Figure 1. Block-d iagram s of th e bathyal bio coe noses d istr ibu tion alo ng th e uppe r con tine n tal slope sec tor accordin g to th e slope gradie nt on th e two main p rofile types, 1 an d ~
n, with the sides sh owing th e sed ime n tatio n prism sinc e th e last Wiirm glaciatio n (hatch ing ) . Fo r profil e location s (P rove nce and Corsica) an d exp lan ation see Emig (1989a, [
1989 b, 1989c) , Em ig and Oarcta-Carrascosa (199 1) . Th e number s in d ica te the depth in me tres. Between do tte d lin es, th e strongest velocity zone of th e near-bo ttom CU lTC n LS
an d th eir main di rectio ns (arrows) . Biocoenos es: (UBM): u pp er barh yal mud ; (SD) : shelf detritic; (BDS): bath yal d eu-itic sa nd ; (MBM): midd le bathyal mud .
.
't
~
""o- ~
c. C. Emig Mediterranean bathyal w nes
1000 ~ ~
"' '''~ '''.J''- ~ '''' ''' "' .... ..... "' ..... "' ,...'"
M editerranean
•
38 .4-38.5 4 .5-4.7
Deep
~
Depth in m P c 5 %. 02 mill Water masses Lower BATHYAL
Figure 2. Diagram o f diffe re nt, parame ters o fth e surro und ing water column in me Bathya l Zo ne (drawn after data from var-
ious au thors}.
bathymetric ex tension, velo city and direction are th e northwestern Mediterran ean Sea (fig ure 2) :
d irec tly relat ed to slope physiog ra p hy (figure 1) . low an n ual tempe rature var iatio n (abo u t 2 DC)
T h e velocity shows a ver tical gradi ent (figure 1) a nd wh ich be co m es co ns tant beyon d a bo ut 200 m
varies fro m abo ut 0.5 up to 2 kn o ts or m ore, an d d e p th (th e Medi terranean h omo th e rm y) , low
some times causes large ripple marks o r undulations salinity increase, and sharp decrease of oAJ'gen
on th e bo ttoms (Reyss and Soyer, 1965; Em ig, 1987) . whi ch, h oweve r, is kn own to be hi g h e r tha n b io-
Conseq ue n tly, sedi mentatio n is absent o r weak in logical consump tion . T h e lowe r b oun dar y co rre-
th e uppe r bathyal, whic h appears to be a resuspen- spon ds to th e lo we r lim it of the fro n tal structure,
sio n lone with minor accum ulation of particles and across whic h the hydrol og ical p ro pe r ties ch a ng e
in wh ich th e fau na is ch aracterise d by high d e nsities sh a r p ly ( H u a n g a n d Su , 199 1), a n d to th e begin-
of suspension-fee ders. At th e lowe r limit o f th e sec- ning of th e oxyge n m ini mu m laye r (O ML) wh ich
to r, th e bo ttom curren t velocity drops off by seve ral is related [ 0 the Levan tin e intermed iate wa ter
te ns of meters, while th e su bstra te becomes m ud dy m ass with a salinity m ax imu m an d h o m o th er my
(Reyss a n d Soye r, 1965; Emig, 1989a; Em ig a nd (fig u re 2) .
Ga rcia-Car rascosa, 1991) : the m ud line is eas ily T he C jN ra tio in th e sed irne n ts dec re ases
de tected in th e case of a ch an ge fro m san dy to fro m 12 to 10 (Carp ine, J970) . T h e ch a nge o f
muddy su bstrates, b ut re main s p ro ble mati cal when sediment oc cur in some tens o f meters, related to
th is lin e occurs sh oreward to th e sh elf edge. the hyd rodyn amics, at bo th lim its of the U p pe r
In th e surro un din g wa ter co lu m n several Bathyal (Em ig, 1989a; E mig a nd Garcia-Carrascosa,
o th e r g ra d ie n ts cha rac te r ise th e u p per bath yal in 199 J ) .
when compared to the ove rlying circalittoral bi o- geographic area, contradicting Bella n-San tini ' s
coenoses; in general, the muddier the su bstrate, (19S3) statemen t that this boundary lies at th e
the lower the richn ess. H owever, the analysis of lower limi t of the circalittoral zone, as well as the
recently p u bli shed d a ta by Albertelli, De lla Croce definition of the lower limi t of the transitional zone
and Frasch etti (1991) , Al bertelli and Fraschetti proposed by Ercegovic (1957) and Peres and
(1992) an d Tse lep ides and Eleftherio u (1992) Picard (1964) .
from m u ddy sh elf-to-u p p er slo pe tr ansects sh ow
that the mac rofauna increases in spec ies and indi-
vid uals, mainly of su sp ension-fe ed ers a nd d etritus- Middle bathyal zone
fe ede rs, in the up pe.r bathyal zone (figu re 3) .
Within the SD an d m iddle bathyal mud (MBM) There is a rapid transitio n over several tens of
bio coe nose s o n the northern Mediterranean coast, mete rs from the upper bathyal zone to the middle
there is a geograph ic decrease in r ich ness fro m the bathyal zo ne, i.e., fro m the u p p er bathyal b io-
west to the east (Ca rp in e, 197 0) , wh ich n ee d s con- coenoses to the MBM . T he su bstrate becomes
firm ation because o pposite to the cyclonic circula- m ud dy, with th e a ppearance of burrowin gs havin g
tion. various diameters (Em ig and Arnau d , 19S8; Emig
Th e extension of circalittoral speci es into the an d Carcia-Carrascosa, 1991) . O ver the co n tinen -
upp er bathyal occurs in general d own to a depth of tal slope in the middle bathyal , with in creasin g
130-140 m , where th eir p re se n ce is prob ably limit- d e p th, a ge neral decrease in the size of the m ac-
ed by abio tic factors, i.e. , ligh t, p ress u re or/ and robenthos is recorded (Carpine, 1970; Peres , 1985;
hydrodynamics, particularly for polychaetes an d Sar d a an d Cartes, 1993; Stefan escu and Cartes,
echinoderrns, wh ile the ex tens ion of 1vfBM species 1992) , as well as a quanti ta tive decrease in th e ma c-
in to th e u pper bathya l is limited by the sub str ate, at robenthos (fig ure 3) and an in crease in d ivers ity
leas t in th e SD and BDS. N ever theless, species related to increased en vironmental stability, associ-
which occur at the same time in the circalittoral ated wi th in cre asin g d e p th (Sanders, 1965; Sa n ders
and b a th yal are generally co nsidered as circ alittoral and Hessler, 1969; H aed rich , Rowe and Po l1on i,
sp ec ies which ex te n d into the bathyal, rare ly th e 19S0; Ab ell6, Vall ad are s and Ca ste1l6n , 19S8) .
contrary. For example, several brachi op od species, Nevertheless, fo ur of th e five grenadier fish specie s
i.e., Megerlia truncata (Linnaeus, 1767) a nd Megathiris occurring in the m id d le bathyal fo llow th e bigger-
detru n cata (Gmelin , 1790) , co nsidered resp ectively as deeper p h e n om e n on (Massu ti, Morales-Nin and
a sh allow-water species and an eurybathic species Stefanescu, 1995).
(Logan, 1979) , h ave their maximum density in There are several faunistic stock changes within
the upp er bath yal, be twee n 100 an d 150 m (Em ig, th e mi d dle b athyal zone, which consequen tly can
19S8). be divided in to severa l d istributional zones (figure
4) , based m ai nly on recent data from crustacean
and fish distribution . However, these zo nes can be
Phytal-aphytal boundary rel ated to several previous proposals, su ch as th a t
of Per es (19S5) , who d ivided th e up p er slope into
T his boundary can be estimated by the percent- two zones below the transitio nal one: fro m 200 to
age of the Gryphus vi/reus shells infested with th e 500 m a nd from 50 0 to 1 000 m. Car p ine (19 70)
green alga Ostreobiu m sp . Bornet and Flahau t. proposed the followi ng limits: transi tional zone,
Occurring generally within th e upp e r bathyal zo ne, 15 0-250 m ; upper bathyal , 250 -550 m ; m idd le
i ts dep th varies in relation to the water transparen- bathyal , 550-2000 m ; an d lower bathyal, I 500-
cy, i.e ., betwee n LSO an d 210 m in normal water 3000 m .
co nditi ons in Corsica, but it can be restricted to The cr us ta cean assemblages occurring o n the
125-135 m u nder turbid wa ter con d itions (Emig , up p er an d on th e lower- middle slo pe correspond
19S9c ). Along the P rove n cal coast, be can se o n ly a to hi ghly d iffere n tia te d faun is tic communities,
small number of shells is infested with Ostreobium whose d ifferences can be a ttr ibu te d to hyd r ol ogi ca l
sp. , this boundary is estima te d at abou t 150 m difference s. The upper community is associated
de pth . Such resu lts in d icat e that the phytal -ap hytal with the warmer intermediate waters, and the
boundary is inde pendent of benthic zo natio n , and lower zone with the deep -water mass, which has a
that its depth can flu ctu a te eve n with in a restricte d fairly co nstan t te m p e rature throughout the year
Cl
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~ I ,... ji 8 ~ species i
to
,., ..- 1 200 ru u~
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Vl
e---e Numb er IIf s p~cic slm~ ~ wi th biornass
:--;'l 0---0 Mean num ber of species/O. ! 10:
~§ 1000 50 o---a M ean abundance
~ .c. <:>-- -OOi OlllllS$
<
'"'" 800 +40
Biornass dryWt glm2
600 + 30
0.4
' K_~ 0 CS)
'.
\,
400 +20
\,
0.2
0 /I \
0---
200+10 ~~~ - - - ~ - ~ - ~ - ~ ~~ ------------- ---------
e;j
I I I I .,--
Depth in m 200 400 600 800 1000 1200 1400 1600 1800 2 000
;,.
f~
Figure 3. (A): Abu n da nce , species rich n ess a n d biom ass of the macro fauu a on the muddy bo ttom s in the shelf-to-slope sector of th e G ulf o f Ge noa (da m fro m AJbc n elli, Dclla [
Croce an d Frasch eu.i, I99 J; AJbene lli an d Frasch eu.i, 1992) . (8 ) ; On a slo pe tran sect in th e Cre ta n Sea (Tsele pid cs an d Eleftheriou , J992) . (C): Biomass o f d em er sal fish co m- '"a.
N
muni ties o n. the slope of Catalo nia (da ta fro m Stefa nescu , Llo ris an d Ruca bad o , 1993 ) .
~
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c. C. Emig Mediterranean. bathj'al zones
too poorly d ocumented to all ow com par iso n (see T h is also seem s to be the ca se in th e Me d i-
Monaco et al., 1990; Bo urcier, Stora an d Cerino, ter ra nean uppe r bathyal, as su gge sted by th e d at a
1993; Gerino et aI., 1995) . T hese valleys are geo- of Alb er te lli an d Cattaneo (198 5) , Albertelli, Dclla
mo rph ol ogically important in concen tratin g and Croce an d Frasch e tti (.1 991) , Alberte lli and
transp or ting se dime n t to d eepe r areas (Mo naco et Frasch e tti (.1 992) and Tse le pid es and Eleftherio u
al., 1990), with h igh dynamic water circulation (1992) on mudd y bottoms (fig u re 3) . H owever, th e
(Mille t, 1990). T heir biol ogica l im p ortance is du e sp ecies and individ ua l ri chness glo ba lly decreases
to the input of b igh prod uctivity with abundant . from circalittoral to bathyal in. the Mediterranean,
re sources, and the occur rence of recru itm ent areas as a lso established by Sh errnan et aL (.1988) in the
co n ce n tr atin g juve nil es of certain species, p articu- n ortheast Atlantic for the d ensity and biornass of
larly in th e middle ba th yal zo n e. Fa un a b iom ass four m ajor ta xonomic groups.
a n d ab un dance are higher than those re corded o n Th e distributional zones in th e Medite rranean
th e sur rou n d ing slope areas. mi d dle b athyal (figure 4) can be com pared with
th e d epth boundar y on th e Atlan tic co n tin ental
slo pe at 400, 700 an d 1200 m , wh ich a re con sid ered
DISCU SSION n early un ive rsal by Hecker (1990) . Nevertheless ,
this author indica tes th e fo llo wing zo n es : upper
T he zone h erein defined as th e upper bathyal slo pe 180-600 m , upp e r-middle slo pe 60 0-1 300 m ,
",..a s previo usly considered a transitional zon e. no t lower-mi ddle slop e 1400-1 60 0 m , lower slo pe .1 600-
only in th e Mediterranean but in the ""orld O cean 2000 m , an d a con tin e n tal rise fr om a bou t 2000 m .
(se e Zenkevitch, 1963; Peres, 1982; Blake and Acco r ding to Van n ey an d Stan ley (.1 983 ), the
Doyle , 1983; Zez in a, 1994 ) . However, thi s zo ne sh elf-ed ge ap p ears a s a boun d ary of importan ce
be gin s at the con tinen tal sh e lf-edge (geomorpho- co mparable to the coastline , se par a ting the coastal-
logi ca l sh e lf-break) which is ch aracte r ised b y bi o- shelf environmen t from the deep realms; it is the
logi cal, physical a n d ge ological expression, and point wh e re th e first maj o r change occurs in the
beyo nd which th e upper bathyal d evelop s as a b el t p hysical gradie n t, and co nsequ en tly, in th e bio logi-
whose b athyrnetric exte n sion is di re ctl y rela te d to cal g ra d ien t, wh en entering the deep se a . T h is
slo p e physiograph y (figure 1) . Con se q ue n tly, u p p er lim it of th e d eep-se a zone, the bathyal,
future stu d ies of bathyal fa u na shou ld always indi- dependin g bathym etrically o n th e de p th of th e
ca te th e slo pe profile from whi ch the d a ta h ave sh elf-edge, is expresse d by biological , physical and
b e en re por te d; rhi s is also valid for th e middle geomorph ol ogi cal criter ia wh ich have to coin cid e
bathya]. In the bathyal , th e major chan ge in fa una with eac h other and with th e lower boundar y of the
is directly associa te d with major e nviron m e ntal circalittoral zo ne.
chan ges, wh ich pl ay a key role in d eter min a tio n of T h e tr ansi tion from the upper bathyal to the
the fauna ac ross the shelf-break. T h e biological middle bathyal, whose bathym etric lim it depends
ex te nsion alo ng the slope is gi ven by th e bio- o n loc al environmental factors govern e d by phys-
co e noses occu rring in th e u pper b a th yal , dominat- iography, al so follows su ch multidisciplinary crite-
ed by hi gh d ensities of ex clusive su spen sio n -fee de rs ri a. It co r respon ds in genera l to the mud lin e ,
wh os e distributional belts sho uld provide informa- lo cated between ab o u t 165 and 300 111, depending
tion regar d ing flo w str uctures in th e shelf-to-slo pe o n th e slo p e physiograph y, in the n orthwest
co n text. Med ite r r a n ean, where this line al so in d icates a
The vali d ity of th e bathyal biocoenoses needs to d rastic ch a nge in th e en ergy leve l, as reported in
be esta blish e d with n ew d at a, usin g fa u nistic den sity other ocean areas by Stan ley, A ddy and Behrens
and bio m ass an d th e effe cts of almos t abi otic char- (1983) .
acters, to d efine cir ca litto ral and b aihyal sp ec ie s As a result of stee p gra d ie n ts in physical con di -
and the exclusive sp e cies of a biocoenosis, whi ch tions, the upper bathyal zo ne re p rese n ts a n unde r-
can only be stenotop ic sp e cies (Ar n a ud and Emig, e stimated barrier fr o m shelf to d e ep sea, occu r rin g
1987). T he dis tr ibutional zo n es of fauna de ter- in the World O ce an. In the Me diterran e an, su ch a
mined b y abi otic fa ctors n ee d a careful bathyrnetric p hysical barrier m ay partly explain th e sh arp fauna
and sp atial choice of sampling alon g th e slope. d e crea se in th e middle bathyal and the bi ol ogical
In th e Atlan tic , in fa u n al density incre ases in the pove r ty of the Med ite r ranea n deep sea. On the
u ppe r m ost slo pe zo ne (Blake and Doyle, 1983). o the r hand, a lo n g-botto m tran sport p assing
through the canyon axes a p pear as p a th ways across Canes, J. E., J. C. Sorbe and F. Sarda. 1994. Spatial dis-
th e sh e lf break to th e Mediterranean deep sea tribution of deep-sea decapods an d eusphausiids near
(Courp and Monaco, 1990) . the bo ttom in th e northwes tern Med ite rrane an.
J. E"/J. Mar Bioi. £CoL 179: 131-144.
Courp , T. and A. Monaco. 1990. Sedimen t dispersal and
accumulation on the co n tine n tal margin of the Gulf
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