Depósi o de in es igación de la Uni e sidad de Se illa
h ps://idus.us.es/
“This e sion o he a icle has been accep ed o publica ion, a e pee e iew
(when applicable) and is subjec o Sp inge Na u e’s AM e ms o use, bu is
no he Ve sion o Reco d and does no e lec pos -accep ance imp o emen s,
o any co ec ions. The Ve sion o Reco d is a ailable online a :
h p://dx.doi.o g/10.1007/s10152-012-0292-5”
1
SOFT BOTTOM CRUSTACEAN ASSEMBLAGES IN MEDITERRANEAN
MARINE CAVES. THE CAVE OF CERRO GORDO (GRANADA, SPAIN) AS
CASE STUDY
C. Na a o-Ba anco1,2, J.M. Gue a-Ga cía1, L. Sánchez-Tocino2, J.C. Ga cía-Gómez1
1Labo a o io de Biología Ma ina, Depa amen o de Fisiología y Zoología, Facul ad de
Biología, Uni e sidad de Se illa, A da. Reina Me cedes 6, 41012 Se illa, Spain. E-
mail: [email p o ec ed] (Au ho o co espondence)
2Depa amen o de Biología Animal, Facul ad de Ciencias, Uni e sidad de G anada,
18071, G anada, Spain
Co esponding au ho :
Tel: (+34) 954556229
E-mail: ca losna a [email protected]
Abs ac
Al hough ma ine ca es a e p io i y conse a ion a eas acco ding he Di ec i e
92/43/CEE o he Eu opean Communi y, he e is a lack o s udies dealing wi h hei so
bo om communi ies. We selec ed he Ce o Go do ca e, a 15 m deep and wi h h ee
di e en zones: a semi da k en ance 25 m long, a da k in e media e a ea o 35 m and
he inal zone a 90 m om he en ance. Sedimen samples we e collec ed by SCUBA
di ing a ou si es: one ou side he ca e (as a con ol) and he o he h ee inside he
ca e coinciding wi h he h ee zones o he ca e. Six ec angula co es o 10x25 cm2
we e collec ed in each si e o mac o aunal s udy and h ee mo e eplica es we e aken
o physico-chemical pa ame e s. The g anulome y showed a clea g adien om
medium sands ou side he ca e o sil and clay in he inne zone. Measu emen s o he
c us acean assemblages showed ha he numbe o species and abundance we e
signi ican ly highe ou side he ca e (30-40 species, >4000 ind m-2) han inside (5-10
species, <1000 ind m-2). Mul i a ia e analyses showed a clea di e ence in species
composi ion be ween ou side and inside he ca e. Cap ellids, anaids, cumaceans and
decapods, we e only ound ou side he ca e and disappea ed inside he ca e, while
gamma ids and isopods we e p esen bo h ou side and inside he ca e. The gamma id
Siphonoece es saba ie i and he anaid Apseudes la eilli we e he dominan species
2
ou side he ca e, while he gamma ids Ha pinia pec ina a, H. c enula a and H. ala we e
dominan inside. The p esen s udy ep esen s an inc ease in deep ange and geog aphic
dis ibu ion o Kupellonu a medi e anea and Monoculodes packa di. This is he i s
ime ha so bo om amphipod communi ies om subma ine ca es o Sou he n Spain
a e desc ibed.
Keywo ds C us aceans, Pe aca ids, Amphipods, ma ine ca es, Medi e anean, So
bo oms
In oduc ion
Ma ine ca e assemblages ha e gene a ed g ea in e es in he scien i ic
communi y o e he las decades (Benede i-Cecchi e al. 1996). This en i onmen is
a ac i e o axonomis s and ecologis s o se e al easons. I is a simpli ied and
oligo ophic sys em, depending solely on ene gy inpu s om he su ounding p oduc i e
coas al a ea. The e a e s ong discon inui ies in he dis ibu ion o o ganisms which
e lec ma ked g adien s in he en i onmen al condi ions. These special condi ions o
da kness, oligo ophy and low hyd odynamism enable he p esence o deepe li ing
species in ha shallow en i onmen . Mo eo e , he in es iga ion o ma ine ca es o en
gi es he possibili y o disco e new and endemic species, due o i s isola ion condi ions
(O and S oboda 1976, Ha melin e al. 1985, Ros e al. 1989). Fo all hese easons,
subma ine ca es a e unique and ulne able ecosys ems (Sa à 1976) p o ec ed by he
Eu opean Communi y (Habi a Di ec i e 92/43 EEC). Howe e , he s udy o ma ine
ca e communi ies has ocused p ima ily on he ben hic communi ies ha inhabi ha d
subs a e, while e y li le e o has been de o ed o he s udy o so bo om
communi ies. Di icul ies in ca ying ou esea ch in such con ined habi a s may help o
explain he lack o exhaus i e s udies on his opic. Unde wa e ca es o en ha e muddy
sedimen a he bo om, whose al e a ion and suspension a e wo o he g ea es dange s
o di ing in o hem, since hey imply an almos comple e loss o isibili y.
All ca e s udies ha e e ealed a ma ked ho izon al zona ion wi hin he animal
communi ies o ha d bo oms inhabi ing he walls (Labo el and Vacele 1958, Sa à
1961, Riedl 1966, Pé ès 1967, Cinelli e al. 1978, Balduzzi e al. 1989, Bibiloni and
Gili. 1982, Ha melin e al. 1985, Gili e al. 1986, Zabala e al. 1989, Gili and
Mcphe son 1987, Bibiloni e al. 1989, Fichez 1990, Benede i-Cecchi e al 1996,
Ha melin 1997, Benede i-Cecchi e al 1998, Bell 2002, Busso i e al. 2002, Ma i e al.
3
2004a, Ma i e al. 2004b, Denni o and Licciano 2006, Busso i e al 2006, Deni o e al
2007, Mosca ello and Belmon e 2007, Busso i and Guide i 2009). A common ea u e
in hese s udies is a dec ease in species ichness, biomass and co e age o ben hic
o ganisms om he ou e mos o he inne mos pa o he ca e. The p oposed
explana ions o hese ea u es a e he physical s ess g adien s inside he ca e (ligh ,
oxygen, salini y, e c), ophic supply g adien and he limi ed capaci y o he la ae o
he dispe sion and se lemen (Ha melin 1985, Zabala e al. 1989, Fichez 1990). All
hese ac o s may also apply o so bo om communi ies, so i would be expec ed o
ind simila pa e ns in hese en i onmen s. Howe e , un il now he a ia ion in so
subs a e assemblages along a shallow subma ine ca e g adien has no been
in es iga ed.
Pe aca id c us aceans a e among he mos di e se and abundan o ganisms in
so -bo om ben hic auna (Fincham 1974; Dau in e al. 1994; P a o and Biandiolino
2005; Lou ido e al. 2008). They also play an impo an ole in s uc u ing ben hic
assemblages (Du y and Hay 2000) and hei abundance and species di e si y may se e
as indica o s o en i onmen al condi ions (Co be a and Ca dell 1995, Gómez-Ges ei a
and Dau in 2000, Gue a-Ga cía and Ga cía-Gomez 2004). Fo anchialine ca es, Ili e
(2005) indica es ha his g oup makes up 90% o species. Fu he mo e, om he
axonomical poin o iew, many new axa a e being desc ibed om ma ine ca es, and
sedimen s o hese ca es a e s ill conside ably unexplo ed. I is di icul o ind ca es
wi h a de ined g adien o sedimen s om he ou e o he inne zones. Consequen ly,
he ca e o Ce o Go do, mo e han 100 m long, is an adequa e si e o s udy, o he i s
ime, he so bo om c us acean communi ies in a Medi e anean ca e g adien .
The aims o his s udy we e: 1) Iden i y he c us acean species which inhabi he
so bo om o Ce o Go do ca e, and 2) de e mine hei dis ibu ion and abundance
along he en i onmen al g adien o he ca e o assess i so bo om communi ies
p esen a simila pa e n o hose ound o he ha d bo om animal communi ies
inhabi ing he walls.
Ma e ial and me hods
S udy a ea
The s udy was conduc ed a he Ce o Go do ca e, a ka s ic subma ine ca e
loca ed in he coas o G anada (Sou h Spain, Albo an Sea, 33º43’46’’ N, 3º45’56’’O)
(Figu e 1). Because o upwelling e en s, which b ing deep and nu ien - ich wa e , his
4
a ea is cha ac e ized o i s e y di e se ben hic communi ies. The ca e is mo e han
100 m long and p esen s a la ge subme ged single en ance. A ma ked na owing o he
ca e 25 m om he en ance sepa a es wo opog aphic a eas. The i s a ea, ex ending
om he en ance o he na owing, is a wide hall, wi h 15 m wide and 15 m high, wi h
an ai chambe on he op. Ligh , al hough educed, is s ill p esen a his hall. The
second a ea is a ec ilinea , blind-ending unnel mo e han 75 m long ha is he da k
a ea. The dep h dec eases along he unnel un il i eaches an ai chambe loca ed a he
end, whe e he e a e eshwa e sp ings. Sampling s a ions we e e e ed as A, B, C and
D. S a ion A was loca ed ou side he ca e, in a sandy bo om a ea nea he en ance.
S a ion B was loca ed in he semi-da k a ea, be ween 15 and 20 m om he en ance.
The o he wo sampling s a ions we e in he da k a ea, one a abou 60 m om he
en ance and he o he a he end o he ca e, in an a ea wi h eshwa e in luence
(Figu e 1).
Sample collec ion and analyses
The sample collec ion was ca ied ou du ing Feb ua y 2001, using a hand-held
co e ec angula o 0,025m2 o a dep h o 10 cm by SCUBA di ing. Six eplica e co e
samples we e aken a each s a ion o he c us acean s udy. Samples we e washed
h ough a 0.5 mm mesh sie e wi h seawa e and ixed wi h e hanol s ained wi h ose
bengal. In he labo a o y, each sample was examined using binocula mic oscopes. All
he c us acean specimens we e coun ed and iden i ied o species whe e possible. Species
di e si y o each sample and he associa ed e enness componen J´ (Pielou, 1966) we e
calcula ed applying he (log2) Shannon Wie e di e si y index (H´) (Shannon &
Wea e , 1963).
Toge he wi h he mac o aunal samples, h ee mo e sedimen co es we e
collec ed a each s a ion o physico-chemical analysis o he sedimen s. All samples
we e immedia ely s o ed ozen un il he labo a o y analysis. G anulome y was
de e mined ollowing he me hod p oposed by Gui ián and Ca ballas (1976). Fo he
chemical analysis, he sedimen s we e ai -d ied, c ushed and sie ed (2mm) i s .
Aliquo s o sedimen samples we e also g ound o < 60 µm p io o de e mina ion o
majo , mino and ace elemen concen a ion. O ganic Ma e (OM) was analysed by
dich oma e oxida ion and i a ion wi h e ous ammonium sulpha e (Walkley and Black
1934). Kjeldahl-N was de e mined by he me hod desc ibed by Hesse (1971). To al
majo , mino and ace elemen concen a ion in sedimen s (<60 µm) we e de e mined

5
by Induc i ely Coupled Plasma-Op ical Emission Spec ome e (ICP-OES Va ian ICP
720-ES axially iewed) ollowing aqua- egia diges ion in a mic owa e o en
(Mic owa e Labo a o y S a ion Miles on ETHNOS 900). The e m “ o al” accoun s o
he aqua- egia diges ion, bu i does no comple ely des oy silica es. The accu acy o
he analy ical me hods was assessed by ca ying ou analyses o he BCR (Communi y
Bu eau o Re e ence) e e ence samples: BCR 320R (Channel sedimen ) and soil
sample e e ence ISE 872 om he Wageningen E alua ing P og ams o Analy ical
Labo a o ies, In e na ional Soil-analy ical Exchange (WEPAL; ISE).
S a is ical analyses
Biological, chemical and g anulome y da a we e no no mally dis ibu ed
(Kolmog o -Smi no es ) and did no ea u e homogenei y o a iance (Le ene es ),
no allowing pa ame ic es o be pe o med. Thus, we op ed o use K uskall-Wallis
analysis o de ec di e ences among s a ions. When hose di e ences exis , Tamhane
pos hoc es s we e used o pos hoc compa isons. The uni a ia e s a is ical analyses
we e ca ied ou using he SPSS 17.0 S a is ic p og am.
Clus e analyses we e conduc ed on abundance pe axonomic g oup o assess
he ela ionship be ween s a ions. Hie a chical clus e ing wi h g oup-a e age linking,
based on simila i y ma ices (B ay-Cu is coe icien ) was used. Da a we e p e iously
squa ed oo ans o med o educe he impo ance o ex eme alues. Wi h he same
aim, o he wo clus e analyses we e conduc ed wi h he g anulome y and chemical
da a. In such cases, he clus e ing wi h g oup-a e age linking was based on euclidean
dis ances ma ices. These analyses we e done using he PRIMER package (Cla ke and
Go ley 2001).
Resul s
A o al o 36 c us acean species was eco ded om he ou s a ions o he s udy.
These comp ised 24 amphipod species, ou cumaceans, ou isopods, wo anaids and
wo decapods (Table 1). The ex e io zone was domina ed by he amphipods
Siphonoece es saba ie i and Me aphoxus ul oni, he anaid Apseudes la eilli and he
decapod Diogenes pugila o . These ou species comp ised 74% o he specimens. The
mos abundan species inside he ca e we e h ee species o genus Ha pinia; Ha pinia
ala, Ha pinia c enula a and Ha pinia pec ina a. Ha pinia pec ina a, which has a
shallowe dis ibu ion, was he only one ound in all s a ions, including ou side he ca e.
6
Ha pinia c enula a was p esen in he h ee s a ions inside he ca e, and Ha pinia ala
only li e in he da k pa o he ca e. O he in e es ing deep species ound inside he
ca e we e he amphipod Monoculodes packa di and he isopod Kupellonu a
medi e anea. Amphipods we e he dominan g oup in abundance and numbe o
species o all s a ions (Figu e 2). Howe e , i s numbe o species and indi iduals was
lowe inside he ca e. The subo de Gamma idea was able o colonize he in e io o he
ca e, while he subo de Cap ellidea was only p esen in he ex e io a ea. The o he
c us acean g oup ha inhabi s he ca e sedimen was he o de Isopoda. Al hough i s
dominance was highe in he inne pa o he ca e, his g oup also p esen ed a dec ease
in ichness and abundance inside he ca e. Tanaidacea, Cumacea and Decapoda we e
only p esen in he ex e nal s a ion. Richness, abundance and di e si y alues showed a
clea educ ion pa e n owa ds he inne pa o he ca e. By con as , e enness
inc eased sligh ly in s a ion D (Figu e 3). The K uskal-Wallis analysis e ealed ha
hese di e ences among s a ions we e signi ican o numbe o species, numbe o
indi iduals and di e si y alues, bu we e no o he e enness. The Tamhane pos hoc
es esul s we e di e en in each case (Table 2).
The g anulome y da a indica ed a p og essi e inc ease o he ine ac ions in
he inne a eas in he ca e (Figu e 4). The K uskal-Wallis analysis showed he exis ence
o signi ican di e ences in he sil and clay composi ion among s a ions (W=10,38;
p=0,016). Tamhane pos hoc es suppo ed he exis ence o h ee g oups; an ex e io
zone, a semida k zone, and a hi d g oup wi h he samples om he da k zone o he
ca e. Conce ning chemical analyses, uni a ia e es s showed signi ican di e ences
be ween s a ions o all a iables, wi h he excep ion o o ganic ma e , ni ogen and
phospho us. Al hough he e we e some di e ences be ween elemen s in he esul s
ob ained in he Tamhane pos hoc es s, mos o hem ag eed o showed signi ican
di e ences be ween all he s a ions excep o he las wo (s a ion C and s a ion D)
(Table 3). The ends obse ed along he ho izon al ansec also a ied. Fo Al, As, Ba,
Co, C , Cu, Fe, K, Li, Mn, Ni, Pb, V and Zn, he e was an inc ease o hei
concen a ions owa d he inne pa o he ca e. In con as , we ob ained an opposi e
end o B, Ca, Cd, Mg, Na, S and S (Figu e 5).
The clus e analyses pe o med o he biological, g anulome y and chemical
da a ag eed o show he exis ence o he h ee dis inc g oups seen be o e (Figu e 6).
The i s o hem included he samples om s a ion A, wi h sedimen s domina ed by
medium sands and a aunis ic composi ion clea ly di e en om he in e nal
7
composi ion, wi h less han 20% o simila i y. The second g oup was om he semida k
a ea (s a ion B). The samples om his zone, whe e he mos abundan sedimen
ac ion was he e y ine sands, appea ed e y close in he h ee clus e analyses (mo e
han 60% o simila i y in hei axonomic composi ion). Finally, he e was a hi d g oup
including he samples om s a ion C and D ( he da k zone) mixed. This a ea had he
lowes alues o ichness, abundance and di e si y, and sedimen s composed mainly o
sil and clay.
Discussion
All he analyses showed a zona ion in he Ce o-Go do ca e. Th ee zones we e
clea ly di e en in axonomic composi ion as well as physicochemical cha ac e is ics: a
pho ic o ex e io a ea, a semida k a ea and a da k a ea. These h ee biocenoses a e
gene ally ecognized in Medi e anean ma ine ca es and hey ha e been iden i ied o
bo h mobile and ha d ben hic ca e communi ies (Riedl 1966, Pé ès 1967, Bibiloni and
Gili 1982, Balduzzi e al. 1989, Gili e al. 1986, Gili and Mcphe son 1987, Bibiloni e
al. 1989, Fichez 1990, Ma i e al. 2004a, Busso i e al. 2006, Deni o e al. 2007,
Mosca ello and Belmon e 2007, Busso i and Guide i 2009). Ne e heless, i was
unknown whe he he dis ibu ion o so subs a e communi ies showed he same
zona ion so a .
The da a ob ained o he ex e io s a ion we e as expec ed acco ding o p e ious
s udies o his ex e nal a ea (Sanchez-Moyano e al. 2005). On he o he hand, he
esul s ound inside he ca e we e mo e in e es ing, wi h many a e species.
Kupellonu a medi e anea was desc ibed by Ba na d (1925) and comple ed by Wägele
in 1981 and Kensley in 1987. This species was ound be ween 70 and 880 m deep and
only on he coas s o Naples, Sicily and Ligu ian sea. Thus, ou inding shows a huge
inc ease in he dep h ange and he geog aphic dis ibu ion o Kupellonu a
medi e anea. Ano he deep species ound in he Ce o-Go do ca e was Monoculodes
packa di, which had a known ba hyme ic dis ibu ion be ween 90 and 2616 m dep h
(Ru o 1993). So his is, so a , he shallowes disco e y o he species.
The dominance shown o he o de amphipoda o e he es o he c us acean
o de s is a common ea u e (Sanchez-Moyano e al. 2005). The p esence o he o de s
isopoda and amphipoda inside he ca e was no unusual because bo h o hem ha e a
g ea capaci y o li ing in mos ma ine habi a s. Kensley (1998) indica ed ha deep sea
(a ma ked oligo ophic en i omen wi h p edominan ly muddy sedimen , like ca e
8
en i onmen s) was he habi a whe e isopods we e mo e di e se. In e es ingly,
cap ellids, cumaceans, decapods and anaids we e absen in he inne a ea o he ca e.
All hese g oups ha e been epo ed om muddy sedimen s (see e.g. Gue a-Ga cía and
Ga cía-Gomez 2004; Lou ido e al. 2008) so o he ac o s apa om he g anulome y
could also a ec .
The esul s showed a clea decline in ichness, abundance and di e si y o he
c us acean so -bo om communi y om he ex e io o he inne da k pa s o he ca e.
This end has been obse ed o many g oups like suspension and il e eede s
(sponges, cnida ians, b yozoans and unica es) (Bibiloni and Gili 1982, Gili e al. 1986,
Balduzzi e al 1989, Bibiloni e al. 1989, Ha melin 1997, Benede i-Cechi e al 1998,
Co ie o 2000, Bell 2002, Ma í e al 2004a, Busso i e al 2006), big decapods (Gili and
Macphe son 1987), meio auna (Toda o e al 2006), plank onic o ganisms (Ga abou and
Flos 1995, Mosca ello and Belmon e 2007), polichae s (Deni o and Licciano 2006),
ishes (Busso i e al. 2002, Busso i e al. 2003), e c. Howe e , his end has no been
epo ed o he mac oin auna so a .
The e a e e y ew in es iga ions dealing wi h mac oin auna o subma ine
ca es. Mo eo e , i we wan o unde s and how he en i onmen al g adien s in ma ine
ca es a ec he c us acean communi y, i is di icul o ex ac conclusions om he
compa ison o ou esul s wi h hose ob ained in hese o he s udies. Mon ei o-Ma ques
(1981), made a axonomical desc ip ion o mac oin auna assemblages om some
subma ine ca es om sou he n F ance. Ne e heless, his samples we e washed h ough
a 1 mm mesh sie e, so mos o c us aceans we e no aken in o accoun . The wo k o
Akoumianaki and Hughes (2004) ook place in he G o a Azzu a, on he coas o I aly.
They s udied he dis ibu ion o mac oin auna inside he ca e and no inwa d dec ease o
mac oin auna abundance o di e si y was obse ed. The explana ion o his was he
exis ence o sulphu sp ings a he end o he ca e, which p o ided an addi ional sou ce
o ood which has a posi i e e ec on he coloniza ion o mac oin auna inside he
esou ce limi ed ca e en i onmen . Thus, o ou knowledge, he p esen s udy is he i s
e idence o a dec ease in so bo om mac oin auna di e si y in Medi e anean ma ine
ca es, when compa ed o communi ies ou side he ca e. Fo ha d ben hic communi ies,
he usual explica ion o his g adien is a educed wa e u no e owa ds he inne pa
o he ca e, which gene ally de e mines oligo ophic condi ions (Ha melin 1985, Fichez
1990). In Ce o-Go do ca e, he g adien in he sil and clay pe cen age in he sedimen
would suppo his heo y, since i indica es a low wa e u no e inside he ca e.
15
Gue a-Ga cía JM, Ga cía-Gómez JC (2004) C us acean assemblages and sedimen
pollu ion in an excep ional case s udy: a ha bou wi h wo opposing en ances.
C us aceana 77:353-370
Gui ián F, Ca ballas T (1976). Técnicas de análisis de suelos. 2º ed. Pico Sac p,
San iago de Compos ela, Spain.
Ha melin JG, Vacele J, Vasseu P (1985) Les g o es sous-ma ines obscu es: un milieu
ex eme e un emà quable bio ope e uge. Té hys 11(3-4);214-229
Ha melin JG (1997) Di e si y o b yozoans in a Medi e anean subli o al ca e wi h
ba hyal-like condi ions: ole o dispe sal p ocesses and local ac o s. Ma Ecol P og Se
153:139-152
Hesse PR (1971) Tex book o soil chemical analysis. J.Mu ay, London.
Ili e TM (2005) Biodi e si y in anchialine ca es. In: Cul e DC, Whi e WB (eds)
Encyclopedia o Ca es. Else ie Academic P ess, London, pp 24-30
Kensley B (1987) A e-e alua ion o he sys ema ics o K. H. Ba na d's e iew o
an hu idean isopods. S eens upia 13(3):101-139
Kensley B (1998) Es ima es o species di e si y o ee-li ing ma ine isopod
c us aceans on co al ee s. Co al ee s 17:83-88
Labo el J, Vacele J (1959) Les g o es sous-ma ines obscu es en Médi e anée. C R
Acad Sci Pa is 248:2619-2621
Lou ido A, Mo ei a J, T oncoso JS (2008) Assemblages o pe aca id c us aceans in
sub idal sedimen s om he Ría de Aldán (Galicia, NW Spain). Helgol Ma Res 62:289-
301

16
Ma í R, U iz JM, Balles e os E, Tu on X (2004a) Ben hic assemblages in wo
Medi e anean ca es: species di e si y and co e age as a unc ion o abio ic pa ame e s
and geog aphic dis ance. J. Ma Biol Ass UK 84:557-572
Ma í R, U iz JM, Balles e os E, Tu on X (2004b) Tempo al a ia ion o se e al
s uc u e desc ip o s in animal-domina ed ben hic communi ies in wo Medi e anean
ca es. J. Ma Biol Ass UK 84:573-580
Ma í R, U iz JM, Tu on X (2005) Spa ial and empo al a ia ion o na u al oxici y in
cnida ians, b yozoans and unica es in Medi e anean ca es. Sci Ma 69(4):485-492
Mon ei o-Ma ques V (1981) Peuplemen s des planche s en asés de ois g o es sous-
ma ines de la égion de Ma seille. E ude p éliminai e. Té hys 10:89-96
Mosca ello S, Belmon e G (2007) The plank on o a shallow subma ine ca e ('G o a di
Ciolo' Salen o Peninsula, SE I aly). Ma Ecol 28(1): 27-59
O JA, S oboda A (1976) Sea ca es as model sys ems o ene gy low s udies in
p ima y ha d bo om communi ies. Pubbl S az Zool Napoli 40:477-485.
Pé ès JM (1967) The Medi e anean ben hos. Ocenog Ma Biol Ann Re 5:449-533
Pielou EC (1966) Species di e si y and pa e n di e si y in he s udy o ecological
sucession. J Theo Biol 10:370-383
P a o E, Biandolino F (2005) Amphipod biodi e si y o shallow wa e in he Ta an o
seas (no h-wes e n Ionian Sea) J Ma Biol Assoc UK 85:333-338
Riedl R (1966) Biologie de Mee eshöhlen. Ve lag Paul Pa ey. Hambu g and Be lin
Robe son MR, Hall SJ, Ele he iou A (1989) En i onmen al co ela es wi h amphipod
dis ibu ion in a Sco ish sea loch. Cah Biol Ma 30:243-258
17
Ros J, Rome o J, Balles e os E, Gili JM (1989) Buceando en las aguas azules. El
ben os. In: Ma gale R (ed) El Medi e aneo Occiden al. Omega, Ba celona, pp 235-297
Ru o S (1993) Pa 3. Oedice o idae. In: Ru o S (ed) The amphipoda o he
Medi e anean. Mémoi es de l'ins i u océanog aphique, Monaco, 13, pp 579-615
Sanchez-Moyano E, Ga cía-Asencio I, Ga cía-Adiego E, Ga cía-Gómez JC, Leal-
Galla do A, Olle o C, F aidías-Ama illo J (2005) Vigilancia ecológica del li o al
andaluz. Conseje ía de Medio Ambien e, Jun a de Andalucía, Se illa.
Sa à M (1961) Zonacione dei Po i e i nella g o a della “Gaiola”. Ann. Is . Mus. Zool.
Uni . Napoli 12:1-10
Sa à M (1976) Il popolamen o delle g o e ma ine: in e esse di una sal agua dia. Pub
S az Zool Napoli 40:50-505.
Shannon CE, Wea e N (1963) The ma hema ical heo y o communica ion. Uni e si y
o Illinois P ess: U bana.
Toda o MA, Leasi F, Biza i N, Tongio gi P (2006) Meio auna densi ies and gas o ich
communi y composi ion in a Medi e anean sea ca e. Ma Biol 149:1079-1091
Tu on X, Ma í R, U iz JM (2009) Chemical bioac i i y o sponges along an
en i onmen al g adien in a Medi e anean ca e. Sci Ma 73(2):387-397
Wägele JW (1981) S udy o he Hyssu idae (C us acea: Isopoda: An hu idea) om he
Medi e anean and he Red Sea. Is J Zool 30(1-2):47-87
Walkley A, Black IA (1934) An examina ion o Deg ja e me hod o de e mining soil
o ganic ma e and a p oposed modi ica ion o he ch omic acid i a ion me hod. Soil
Sci 37:29-37
18
Zabala M, Rie a T, Gili JM, Ba ange M, Lobo A, Peñuelas J (1989) Wa e low, ophic
deple ion, and ben hic mac o auna impo e ishmen in a subma ine ca e om he
wes e n medi e anean. Ma Ecol 10(3):271-287.
Fig. 1. Loca ion and longi udinal sec ion o Ce o Go do ca e, showing he samples
s a ions.
Fig. 2. Dominance (%) o di e en g oups calcula ed in unc ion o numbe o species
(A) and indi iduals (B)
Fig. 3. Mean alues ± S anda d de ia ion o numbe o species/sample (S),
indi iduals/m2 (N), e eness (J’) and di e si y (H’) in each s a ion.
Fig. 4. Pe cen age o each g anulome ic ac ion in he sedimen pe sample. OM=
O ganic ma e
Fig. 5. Mean concen a ion ± S anda d de ia ion o each chemical a iable pe s a ion.
Fig. 6. Clus e analysis o samples using he abundance o each species (A), he
chemical da a (B) and he g anulome ic da a (C)
19
020 40 60 80 100
D
C
B
A
CUMACEA
TANAIDACEA
DECAPODA
ISOPODA
AMPHIPODA
020 40 60 80 100
D
C
B
A
CUMACEA
TANAIDACEA
DECAPODA
ISOPODA
AMPHIPODA
A)
B)
S a ions
S a ions
Dominance (%)
Dominance (%)
20
0%
20%
40%
60%
80%
100%
A1
A2
A3
B1
B2
B3
C1
C2
C3
D1
D2
D3
Sil and clay
Ve y ine and ine sands
Medium sands
Ve y g oss and g oss sands
Samples
Pe cen age(%)

21
A B C D
0
0,5
1
OM
S a ion
% OM
A B C D
0
0,04
0,08 N
S a ion
% N
A B C D
0
300
600 P
S a ion
P (mg/kg)
A B C D
0
10000
20000
30000 Al
S a ion
Al (mg/kg)
A B C D
0
4
8As
S a ion
As (mg/kg)
A B C D
0
10
20
30 B
S a ion
B (mg/kg)
A B C D
0
50
100 Ba
S a ion
Ba (mg/kg)
A B C D
0
100000
200000
300000 Ca
S a ion
Ca (mg/kg)
A B C D
0
0,06
0,12 Cd
S a ion
Cd (mg/kg)
A B C D
0
10
20 Co
S a ion
Co (mg/kg)
A B C D
0
20
40 C
S a ion
C (mg/kg)
A B C D
0
10
20 Cu
S a ion
Cu (mg/kg)
A B C D
0
20000
40000
Fe
S a ion
Fe (mg/kg)
A B C D
0
3000
K
S a ion
K (mg/kg)
A B C D
0
20
40 Li
S a ion
Li (mg/kg)
A B C D
0
12000
24000 Mg
S a ion
Mg (mg/kg)
A B C D
0
200
400 Mn
S a ion
Mn (mg/kg)
A B C D
0
2000
4000 Na
S a ion
Na (mg/kg)
A B C D
0
20
40 Ni
S a ion
Ni (mg/kg)
A B C D
0
20
40 Pb
S a ion
Pb (mg/kg)
A B C D
0
2000
4000 S
S a ion
S (mg/kg)
A B C D
0
600
1200 S
S a ion
S (mg/kg)
A B C D
0
50
100 V
S a ion
V (mg/kg)
A B C D
0
50
100 Zn
S a ion
Zn (mg/kg)
22
100806040200B ay Cu isSimila i y (%)
A4
A1
A6
A2
A3
A5
D5
D2
D3
D6
C5
C6
C2
C3
C4
D1
C1
B1
B3
B2
B6
B4
B5
050000100000150000200000 Euclidean Dis ance
A1
A2
A3
C1
D3
D1
D2
C2
C3
B2
B1
B3
A2
020406080 Euclidean Dis ance
A1
A3
B1
B2
B3
D1
D2
C2
C1
C3
D3
A)
B)
C)