Helgol Ma Res (2008) 62:189–199
DOI 10.1007/s10152-008-0107-x
123
ORIGINAL ARTICLE
Communi y s uc u e o cap ellids (C us acea: Amphipoda:
Cap ellidae) on seag asses om sou he n Spain
A. R. González · J. M. Gue a-Ga cía · M. J. Maes e ·
A. Ruiz-Taba es · F. Espinosa · I. Go dillo ·
J. E. Sánchez-Moyano · J. C. Ga cía-Gómez
Recei ed: 10 Ap il 2007 / Re ised: 17 Decembe 2007 / Accep ed: 8 Janua y 2008 / Published online: 5 Feb ua y 2008
© Sp inge -Ve lag and AWI 2008
Abs ac The communi y s uc u e o cap ellids inhabi ing
wo species o seag ass (Cymodocea nodosa and Zos e a
ma ina) was in es iga ed on he Andalusian coas , sou he n
Spain, using uni and mul i a ia e analyses. Th ee meadows
we e selec ed (Alme ía, AL; Málaga, MA; Cádiz, CA), and
changes in seag ass co e and biomass we e measu ed om
2004 o 2005. Fou cap ellid species we e ound; he den-
si y o Cap ella acan hi e a, Ph isica ma ina and Pseudo-
p o ella phasma was co ela ed o seag ass biomass. No
such co ela ion was ound o Pa iambus ypicus, p obably
because his species inhabi s sedimen s and does no cling
o he seag ass lea es. We eco ded a signiWcan dec ease in
seag ass co e and biomass in MA due o illegal bo om
awling Wshe ies. Ph isica ma ina and P. ypicus we e
a ou ed by his pe u ba ion and inc eased hei densi ies
a e he awling ac i i ies. A su ey o epo s on cap el-
lids in seag ass meadows a ound he wo ld showed no clea
la i udinal pa e ns in cap ellid densi ies ( anging om 6 o
1,000 ind/m2 pe meadow) and species di e si y. While
cap ellid abundances in seag ass meadows a e o en e y
high, he numbe o species pe meadow is low ( ange 1–5).
Keywo ds Cap ellidae · Seag asses · Sou he n Spain ·
T awling
In oduc ion
Cap ellids a e small ma ine pe aca idean c us aceans,
which inhabi algae, hyd oids, ascidians, an hozoans, b y-
ozoans, sponges and seag asses (McCain 1968; Gue a-
Ga cía 2001). They eed on suspended ma e ials, p ey on
o he o ganisms, o g aze on epibio ic auna and Xo a
(Caine 1974; Gue a-Ga cía e al. 2002; Thiel e al. 2003),
and hey a e impo an p ey o many coas al Wsh species
(Caine 1987, 1989, 1991). Cap ellids a e mo phologically
well adap ed o cling o he subs a a; wi h hei pe eopods
hey can W mly hold on o b anches o algae, seag ass, b y-
ozoans and hyd ozoans. The pleopods, which a e used o
swimming in o he amphipod c us aceans, a e e y educed
in cap ellids; he e o e, al hough cap ellids can swim
(Caine 1979a) he e a e no e y eYcien swimme s. How-
e e , cap ellids can be dis ibu ed passi ely by clinging o
a iWcial (buoys, opes, li e ) and na u al (mac oalgae)
Xoa ing ma e ials, so he cosmopoli an dis ibu ion o many
li o al cap ellid species migh be acili a ed by he ac ha
hey a e o en associa ed wi h ouling communi ies on
Xoa ing objec s (Thiel e al. 2003). Recen ly, cap ellids
ha e also been ound o be use ul bioindica o s o ma ine
pollu ion and en i onmen al s ess (Gue a-Ga cía and Ga -
cía-Gómez 2001; Ohji e al. 2002; Takeuchi e al. 2004;
Gue a-Ga cía and Koonjul 2005). Al hough amphipods
(gamma ids and cap ellids) a e egula inhabi an s o sea-
g ass meadows, he e is a lack o ecological and beha -
iou al s udies on he cap ellid communi ies associa ed o
seag asses.
Seag asses a e dis ibu ed wo ldwide (600,000 km2 o
he ma ine bo oms a e co e ed by hese spe ma ophy es)
and play an impo an ole in he gene al coas al dynamics
and biology (La kum e al. 1989; Templado 2004). When
compa ed wi h neighbou ing a eas, he meadows e eal
Communica ed by H.-D. F anke.
A. R. González · J. M. Gue a-Ga cía (&) · M. J. Maes e ·
A. Ruiz-Taba es · F. Espinosa · I. Go dillo ·
J. E. Sánchez-Moyano · J. C. Ga cía-Gómez
Depa amen o de Fisiología y Zoología,
Labo a o io de Biología Ma ina, Facul ad de Biología,
Uni e sidad de Se illa, A da Reina Me cedes 6,
41012 Se illa, Spain
e-mail: jmgue [email p o ec ed]
190 Helgol Ma Res (2008) 62:189–199
123
highe abundances and species ichness (Edga e al. 1994).
The main ac o s con ibu ing o his imp o emen in biodi-
e si y a e a ailabili y o mic ohabi a , p o ec ion om p e-
da o s, ophic esou ces, sedimen se ling, hyd odynamic
o ce educ ion (see P ano i e al. 2000). Seag ass beds o
he empe a e zone suppo la ge numbe s o in e eb a e
species and indi iduals, he eby p o iding abundan ood
o Wshes, compa ed o adjacen un ege a ed a eas (Nakam-
u a and Sano 2005). A Eu opean coas s, ou na i e sea-
g ass species a e known, which a e all dis ibu ed along he
li o al o Andalusia, sou he n Spain: Zos e a ma ina Lin-
naeus, Cymodocea nodosa (Uc ia) Asche son, Posidonia
oceanica (Linnaeus) Delile and Zos e a nol ii Ho nemann.
In spi e o he abundance o hese seag ass meadows in
sou he n Spain, cap ellid communi ies associa ed o hese
plan s ha e been sca cely s udied, and he only eco ds o
cap ellids om hese habi a s come om gene al aunis ic
o ecological s udies (Edga 1990; Sánchez-Je ez and
Ramos Esplá 1996; Rod íguez-Ruiz e al. 2001; Luque
e al. 2004; Balles e os e al. 2004). This lack o in o ma-
ion is also applicable o o he a eas a ound he wo ld
(Takeuchi and Hino 1997).
Because o he ecology and economic impo ance o sea-
g ass meadows, hei p o ec ion has been p oposed in
ecen legisla ion a local, na ional and in e na ional le el.
Seag ass meadows a he Andalusian coas ha e been
dec easing s eadily. In addi ion o na u al p ocesses,
an h opogenic ac o s a e likely o ha e inXuenced his
decline (Sánchez-Je ez and Ramos-Esplá 1996). Besides
he inc easing o u ban and indus ial a eas in he li o al
zone, he e is an impo an eVec o he bo om awling
Wshe ies. La ge numbe s o awle s usually wo k illegally
o e seag ass meadows, causing physical deg ada ion and
c i ical eg ession o he meadows. In ac , al hough a ia-
ion in s uc u al complexi y in seag ass may well be p o-
duced by o he en i onmen al ac o s, human ac i i ies
such as awling play a e y impo an ole in he SE o he
Ibe ian Peninsula (Sánchez-Lizaso e al. 1990). Fo all
hese easons, a esea ch p og amme on seag ass meadows
o he coas o Andalusia, sou he n Spain was ini ia ed in
2004. The p ojec , suppo ed by he En i onmen al Agency
o he Andalusian Go e nmen , is in ended o con ol and
de ec empo al changes in seag ass biomass, densi y, co e
and associa ed auna. To p ope ly assess changes in he
ex en o seag ass meadows h oughou ime, sampling
eVo was mainly ocused on meadows edges. As a pa o
his gene al p ojec , we s udied o he communi y s uc u e
o he cap ellid amphipods associa ed o h ee seag ass
meadows o he Andalusian coas du ing he yea s 2004
and 2005, using uni and mul i a ia e app oaches. Fu he -
mo e, as one o he s udied seag ass meadow was se e ely
aVec ed du ing he s udy by awling Wshe ies, we also
es ed he esponses o cap ellids o seag ass eg ession
p oduced by awle s.
Me hods
S udy si es
The h ee seag ass meadows selec ed o he p esen s udy
a e dis ibu ed along he o ien al coas o Andalusia
(Fig. 1). A he occiden al coas , wa e anspa ency is sig-
niWcan ly lowe and no seag ass can be ound. The main
cha ac e is ics o he h ee selec ed meadows a e gi en in
Table 1.
Fig. 1 S udy a ea showing he loca ion o he h ee selec ed seag ass
meadows. CA Cádiz, MA Málaga, AL Alme ía
Ibe ian
Peninsula
Mde i
e aneanSae
A lan ic Ocean
Andalusia
CA
MA AL
AFRICA
200 km
Table 1 Cha ac e is ics o he h ee seag ass meadows sampled
Code Locali y Coo dina es Seag ass species Su ace (m2) Dep h (m) Biomass (g/m2)
AL Los Geno eses (Alme ía) 36°44.9⬘N–02°06.6⬘WCymodocea nodosa 10,000 10–11 49
MA Ma o (Málaga) 36°44.5⬘N–03°47.6⬘WZos e a ma ina 55,000 11–16 55
CA Ta i a (Cádiz) 36°01.1⬘N–05°37.2⬘WCymodocea nodosa 30,000 11–17 44
Helgol Ma Res (2008) 62:189–199 191
123
Sampling p ocedu e
Es ima ion o changes in seag ass co e
Once he exac loca ion and dimensions o he h ee sea-
g ass meadows had been checked using SCUBA, he ou
edges (no h, sou h, eas and wes ) we e loca ed. On each
edge, 6 Wxed quad a s o 1 £1m
2 (Fig. 2) we e ma ked
wi h shea es o moni o ing he seag ass co e h ough
ime. Each quad a (24 o each meadow, 72 in o al) was
pho og aphed and, using image analysis, di ided in o 64
subsqua es. In each subsqua e he seag ass p esence-
absence was checked and he co e o he quad a was
exp ess as he pe cen age o p esence numbe s om he
o al (64 subsqua es). Co e was measu ed in summe 2004
and 2005.
Es ima ion o seag ass biomass and cap ellid composi ion
On each edge W e andom samples o 15 £15 cm2
(Fig. 2) we e collec ed in summe 2004 and 2005. Sam-
ples we e sie ed (mesh size o 0.5 mm), Wxed in e hanol
85% and s ained wi h bengal ose. In he labo a o y,
cap ellid amphipods we e so ed and iden iWed o spe-
cies le el unde a binocula mic oscope. Seag ass o
each sample was sepa a ed and he biomass (d y weigh )
measu ed.
Sedimen cha ac e is ics
Th ee samples o sedimen we e collec ed om each sea-
g ass meadow in summe 2004 and 2005. Sedimen sam-
ples we e s o ed a ¡20°C in p e-cleaned glass ja s un il
analysis. G anulome y was de e mined by Buchanan and
Kain’s me hod (Buchanan and Kain 1984). O ganic con-
en s we e analysed by ashing samples o sedimen o
500°C o 6 h and e-weighing (Es acio e al. 1997).
S a is ical analysis
Va ia ions in sedimen g anulome y and o ganic ma e
be ween 2004 and 2005 we e es ed using one-way
ANOVA a e e i ying he no mali y o he da a (Kol-
mogo o –Smi no es ) and he homogenei y o a iances
(Le ene es ).
The inXuence o loca ion and ime (yea 2004 s. 2005)
on co e and biomass o he seag ass was analyzed using
wo-way ANOVA. Va ia ions o he seag ass biomass and
co e be ween 2004 and 2005 and changes o cap ellid den-
si ies o each seag ass meadow we e es ed using K uskal–
Wallis es .
To explo e he ela ionship be ween cap ellid densi y
and seag ass biomass, co ela ion analyses we e conduc ed.
The aYni ies among seag ass based on cap ellid species
we e es ablished by clus e analysis using UPGMA
Fig. 2 Sampling design
Quad a s o 15x15 cm
2
o es ima ing he
seag ass biomass and cap ellid densi y
Fixed squa es o 1x1 m
2
o measu ing
changes in seag ass co e om 2004 o 2005
N
W
S
E
SEAGRASS MEADOW
192 Helgol Ma Res (2008) 62:189–199
123
me hod. Mul i a ia e analyses we e ca ied ou using he
PRIMER (Plymou h Rou ines in Mul i a ia e Ecological
Resea ch) package (Cla ke and Go ley 2001). Fo uni a i-
a e analyses, he BMDP (BioMedical Da a P og ams) was
used (Dixon 1983).
Resul s
Sedimen cha ac e is ics
The seag ass meadows o Alme ía (AL) and Cádiz (CA)
we e domina ed by Wne sands while in he seag ass o
Málaga (MA) e y Wne sands p edomina ed (Fig. 3). The
seag ass o MA was aVec ed by awling ac i i ies om 2004
o 2005. As a esul , a signiWcan change in g anulome y was
measu ed (an inc ease in Wne sand, F=21.7, P< 0.05, and a
dec ease in e y Wne sand, F= 39.3, P< 0.01). As o o ganic
ma e (Fig. 4), he e we e no signiWcan diVe ences be ween
2004 and 2005 o he h ee s udied meadows.
Seag ass co e and biomass
Using he Wxed squa es o 1 £1m
2 o es ima e changes in
co e , we measu ed a signiWcan dec ease o seag ass co e
( om 57 o 17%, K uskal–Wallis (K) = 32.4, P< 0.001) in
Málaga (MA), p obably due o he eVec o Wshing awle s
in he a ea. The eVec was also signiWcan o seag ass bio-
mass in MA ( om 55 o 23g/m
2; K=18.2, P< 0.001)
(Fig. 5). When pooled da a o all seag ass meadows we e
analysed using wo-way ANOVA, signiWcan diVe ences in
co e and biomass we e measu ed o bo h loca ion (AL,
MA o CA) and yea (2004 o 2005) (Table 2). Howe e , a
signiWcan in e ac ion was also measu ed be ween he wo
ac o s, due o he di e gen beha iou o he Málaga
meadow.
Cap ellid communi y
Fou cap ellid species we e ound du ing he p esen s udy:
Cap ella acan hi e a (Leach, 1814), Pa iambus ypicus
(K öye , 1844), Ph isica ma ina Slabbe , 1769, and
Pseudop o ella phasma (Mon agu, 1804). Species abun-
dances o he h ee s udied seag ass meadows a e gi en in
Fig. 6. Ph isica ma ina and P. ypicus we e he dominan
species, while P. phasma was la gely es ic ed o Alme ía
and Cap ella acan hi e a was only ound in Cádiz.
In Alme ía (AL) and Cádiz (CA) he e we e no signiW-
can diVe ences in cap ellid densi ies be ween 2004 and
2005. Howe e , a signiWcan inc ease in numbe o speci-
mens we e measu ed o P. ma ina and P. ypicus in Mál-
aga (MA), p obably as a esul o he pe u ba ion
associa ed wi h awling ac i i ies (P.ma ina, om 5.9 o
15.7 ind/225 cm2, K= 9.8, P<0.05; P. ypicus, om 0.6 o
3.1 ind/225 cm2, K= 4.4, P<0.05).
Fig. 3 Pa icle size dis ibu ion in he sedimen s o he h ee s udied
seag ass meadows o 2004 and 2005. Values a e means o h ee epli-
ca es each. AL Alme ía, MA Málaga, CA Cádiz
0%
50%
100%
2004 2005 2004 2005 2004 2005
Sil -clay (<0.063 mm)
Ve y ine sand (0.125-0.063 mm)
Fine sand (0.25-0.125mm)
Medium sand (0.5-0.25 mm)
G oss sand (1-0.5 mm)
Ve y g oss sand (2-1 mm)
G a el (>2 mm)
AL MA CA
Fig. 4 O ganic ma e con en o he sedimen s in he h ee s udied
seag ass meadows o 2004 and 2005 (means and SD); AL Alme ía,
M
A Málaga, CA Cádiz
0
AL MA CA
1
2
3
4
5
2004
2005
%
Helgol Ma Res (2008) 62:189–199 193
123
Fo bo h 2004 and 2005 he cap ellid communi y,
al hough consis ing only o ou species, allowed o dis-
c imina ing among he h ee meadows, acco ding o he
clus e analysis (Fig. 7). Each meadow was cha ac e ised
by a speciWc composi ion and/o abundance o he cap ellid
communi y.
The densi y o P. ma ina, P. phasma and C. acan hi e a
was signiWcan ly co ela ed o he seag ass biomass
( = 0.51, 0.40 and 0.50, espec i ely; P< 0.05) while he e
was no co ela ion o P. ypicus.
Discussion
Cap ellid communi y on Andalusian seag asses
The ou cap ellid species ound in he s udied seag ass
meadows, C. acan hi e a, P. ma ina, P. phasma and P. yp-
icus, a e common species on many diVe en subs a a (see
Gue a-Ga cía 2001). Ph isica ma ina is a cosmopoli an
species ha li es on algae, hyd oids, ascidians, an hozoans,
sponges, b yozoans and sedimen s (Gue a-Ga cía 2001). I
can bea s essed a eas o low hyd odynamics and high
a es o sedimen a ion and o ganic ma e (Gue a-Ga cía
and Ga cía-Gómez 2001). Cap ella acan hi e a is also
widely dis ibu ed in he Medi e anean, being especially
abundan on algae, bu also associa ed wi h a a ie y o
subs a a. Pseudop o ella phasma is usually associa ed wi h
hyd oids, bu can be ound also on algae, an hozoans and
sedimen . Pa iambus ypicus has been ound mainly on
sedimen s and also associa ed wi h echinode ms (Gue a-
Ga cía 2001). Consequen ly, no speciWc associa ions could
be es ablished be ween hese cap ellids and seag ass spe-
cies; howe e , speciWc associa ions be ween cap ellids and
seag ass species ha e been epo ed in o he a eas (e.g.
Cap ella japonica li ing on he seag ass Phyllospadix iwa -
ensis in Japan, Takeuchi and Hino 1997). Al hough he e is
a lack o s udies dealing wi h cap ellids on seag ass a ound
he Ibe ian Peninsula, he cap ellid composi ion is e y
simila in he ou seag ass species (Table 3).
Cap ellids may play an impo an ole in seag ass mead-
ows. Cap ellids li ing on seag ass a e impo an p ey o
many Wshes (Caine 1991; Ho inouchi e al. 1998; Rod í-
guez-Ruiz e al. 2001; Sánchez-Je ez e al. 2000). Some
species o Wshes associa ed wi h seag ass meadows con-
sume p ima ily cap ellids, especially du ing ju enile s ages
(see Kwak e al. 2005). In ac , in shallow wa e ecosys-
ems, cap ellidean and gamma idean amphipods a e consid-
e ed o be one o he mos impo an p ey i ems o Wshes,
especially o hose less han 10 cm in body leng h (Takeu-
chi and Hino 1997). Fu he mo e, he amphipod and gas o-
pod g aze s a e e y impo an in con olling pe iphy on
and ephiphy es o seag ass (Je nakoV and Nielsen, 1997).
Fo example, Caine (1980) epo ed ha in he absence o
Cap ella lae iuscula, pe iphy on biomass inc eased by
411% in Z. ma ina beds.
Fig. 5 Seag ass co e and biomass o 2004 and 2005 in he h ee
s udied meadows (means and SD); AL Alme ía, MA Málaga, CA
Cádiz; * P<0.001
0
AL MA CA
AL MA CA
20
40
60
80
%
2004
2005
0
20
40
60
80
g/m2
Seag ass co e
Seag ass biomass
*
*

194 Helgol Ma Res (2008) 62:189–199
123
Cap ellids and seag ass biomass. Ve ical dis ibu ion
o cap ellids in he meadow
In opical seag ass meadows, seag ass biomass has usually
been ound co ela ed wi h bo h numbe and abundance o
in e eb a e species. A hick ege a ion p o ides be e p o-
ec ion om p eda o s and a la ge plan su ace o cling on
(Heck and We s one 1977). In he p esen s udy, he abun-
dance o P. ma ina, P. phasma and C. acan hi e a co e-
la ed wi h seag ass biomass while he abundance o P.
ypicus did no . This may be explained by he e ical dis-
ibu ion o he cap ellid species in he seag ass meadow.
Ph isica ma ina is dis ibu ed on bo h blades and sedimen ;
P. phasma and C. acan hi e a li e mainly on lea es while
P. ypicus can be ound wi hin sedimen s, among sand
g ains (pe sonal obse a ion; Fig. 8). This species seems o
li e in sedimen s ega dless o he local seag ass biomass,
i.e. e en in plain sedimen s. In con as , he abundances o
he o he species we e highly co ela ed wi h seag ass bio-
mass, since hey depend on seag ass blades o cling on.
Cu iously, in a s udy conduc ed in a seag ass meadow o Z.
ma ina in he Salcombe Es ua y, UK, P. ypicus was mo e
abundan in agmen ed han in con inuous a eas (F os
e al. 1999).
Acco ding o Vi ns ein e al. (1984), mos ee-li ing
gamma id amphipods end o hide be ween seag ass blades
Table 2 Two-way ANOVA esul s o he inXuence o he loca ion (AL, MA, CA) and yea (2004, 2005) on he seag ass co e and biomass
(*p < 0.05, **p < 0.01)
Sou ce o a ia ion Co e (%) Biomass (g/225 cm2)
Mean § SE DF SS F Mean § SE DF SS F
Loca ion 2 6531.3 23.2** 2 2.7 4.2*
AL 57.7 §12.4 1.2 §0.7
MA 37.5 §23.1 0.9 §0.5
CA 43.2 §13.2 1.2 §0.6
Yea 1 11166.4 79.4** 1 0.01 0.02*
2004 54.9 §12.3 1.2 §0.6
2005 32.4 §18.5 1.0 §0.6
Loca ion £yea 2 7244.9 25.7** 2 7.4 11.3**
Fi
g.
6C
ap e
llid
a
b
un
d
ances
(ind/225 cm2) in he h ee sea-
g ass meadows o 2004 and
2005 (means and SD); AL
Alme ía, MA Málaga, CA Cádiz;
*P< 0.05; scale ba s 1mm
0
5
10
15
20
CAMA
AL CAMA
AL
CA
MA
AL CA
MA
AL
0
5
10
15
20
0
5
10
15
20
0
5
10
15
20
acisi hP
ani am
a e ih nacaalle paC
*
*
2004
2005
submai aP
sucipy
alle o poduesP
a
m
s
a
h
p
Helgol Ma Res (2008) 62:189–199 195
123
while cap ellids li e mo e exposed on he su ace and ips
o he blades. Pe haps coun e ac ing hei ulne able posi-
ion on he lea su aces, cap ellids ha e a skele on-like
mo phology, which may make hem less conspicuous o
isual p eda o s (Vi ns ein e al. 1984). In ac , al hough
mo e beha iou al s udies a e necessa y o mos species,
Cap ella japonica and C. suga ensis ha e been obse ed
o hold seag asses in a “pa allel” pos u e (Takeuchi and
Hino 1997). Al hough cap ellids a e epo ed o be mo e
“camouXaged” in seag ass meadows han in o he habi a s,
he e a e o he subs a es, such as hyd oids, whe e cap ellids
can emain unno iced by p eda o s. Caine (1979b) com-
pa ed popula ions o Cap ella lae iuscula li ing on Zos e a
and on he hyd oid Obelia, and ound ha p eda ion was
highe on Zos e a han on Obelia, especially o ju eniles
and emales. Ju enile and emale selec i i y by p eda o s
a e ela ed o isual disce nibili y; ju eniles lack a p o ec i e
colo a ion on Zos e a, and he b ood pouch o o ige ous
emales is whi e. On Obelia, he ligh e colou blends wi h
he backg ound and ju eniles esemble polyps.
T awling eVec on cap ellid communi y
The p esen s udy showed ha P. ma ina and P. ypicus
we e clea ly a ou ed by he pe u ba ion associa ed wi h
awling ac i i ies. Sánchez-Je ez and Ramos-Esplá (1996)
also ound highe densi ies o some cap ellid species in
a eas impac ed by awle s in compa ison wi h con ol
a eas along C. nodosa meadows. F os e al. (1999) ound
highe densi ies o P. ypicus in agmen ed a eas o Z.
ma ina beds han in con inuous zones. Ph isica ma ina and
P. ypicus ha e been epo ed o be e y good colonize s o
sedimen s a e d edging in ha bou s (Gue a-Ga cía e al.
2003), and excellen ecolonize s o de auna ed sands in
expe imen al ays (Gue a-Ga cía and Ga cía-Gómez
2006). These wo species can be conside ed as “oppo unis ic”,
Fig. 7 Clus e analysis conduc ed using he mean alues o cap ellid ob ained o each edge (N, S, E and W), yea (2004, 2005) and seag ass meadow
(CA, MA, AL)
S
-
4
002
-LA
W-
5
00
2
-L
A
N-4
0
02-LA
E-
5
002-LA
E-
40
0
2
-L
A
W-4002-
L
A
S
-
5
0
02-LA
N-5
0
02-LA
N
-
4
002-
A
C
S-
4
0
02-
A
C
W
-
5002
-
A
C
E-
5
0
0
2
-
A
C
W
-40
02-
AC
N
-5
0
0
2
-A
C
E
-4
0
0
2
-
A
C
E
-
5
0
0
2-
AM
E
-
4
0
02
-
A
M
S
-50
0
2-
A
M
S-5
0
0
2
-
AC
S
-
4
0
02
-
AM
W-4
0
0
2
-
AM
W
-5
0
0
2
-
A
M
N-4
0
0
2-
A
M
N
-
5
0
02
-
A
M
100
80
60
40
20
0
Simila i y (%)
AL MACA
Table 3 Cap ellids ound in seag ass meadows o he Ibe ian Peninsula, Cana y Islands and Balea es
1Sánchez-Je ez e al. 1999 (Alican e); 2Rie a e al. 2003 (Cana y Islands); 3Balles e os e al. 2004 (Andalusia); 4Luque e al. 2004 (Andalusia);
5Box, unpublished da a (Balea es); 6p esen s udy (Andalusia); 7Cu ás 1990 (Galicia); 8Mo ei a 2003 (Galicia) 9Cacabelos, unpublished da a
(Galicia); 10Gue a-Ga cía, unpublished da a (Andalusia)
C. nodosa1,2,3 P. oceanica4,5,6 Z. ma ina6,7,8 Z. nol ii9,10
Cap ella acan hi e a ++ +
Cap ella equilib a +
Cap ella apax +
Pa iambus ypicus +++
Ph isica ma ina ++ ++
Pseudop o ella phasma ++ ++
196 Helgol Ma Res (2008) 62:189–199
123
Table 4 Cap ellid composi ion and o al abundance (ind/m2) in diVe en seag ass meadows om se e al wo ld a eas
T : Thalassia es udinum; S : Sy ingodium Wli o me; Zm: Zos e a ma ina; Cn: Cymodocea nodosa; Po: Posidonia oceanica; Zn: Zos e a nol ii; Zj: Zos e a japonica; Aa: Amphibolis an a c ica;
Ag: Amphibolis g iY hii; Ps: Posidonia sinuosa; Pi: Phyllospadix iwa ensis; Zc: Zos e a caulescens
1S one (1983), 2Nelson (1979), 3Sánchez-Je ez e al. (1999, 2000), 4S iso e al. (2001), 5Bologna and Heck (1999), 6Caine (1991) and Thom e al. (1995), 7Edga (1990), 8F ed iksen e al.
(2005), 9Je nakoV and Nielsen (1998), 10Takeuchi and Hino (1997), 11p esen s udy; 12Cu ás (1990)
Indian Ri e 1
Flo ida, USA
Beau o 2,
USA
Alican e3,
SE Spain
Venice4,
I aly
S Joseph5,
Flo ida, USA
Padilla Bay6,
Washing on, USA
Se en Mile7,
W Aus alia
Skage ak8,
No way
Pe h coas 9,
W Aus alia
O suchi Bay10,
NE Japan
Andalusia11,
S Spain
Lugo12,
NW Spain
T +S Zm Cn Po Cn Zn T Zm Z,j Aa + Ag Zm Ag Ps Pi Zm Zc Cn Zm Zm
Cap ella acan hi e a – – 0.5 2.7 6 17 – – – – 26 – – – – – 1.8 – –
Cap ella bispinosa – – ––––– – – – – – – –6–– – –
Cap ella danile skii – – ––––– – – – – – – 4.2––– – –
Cap ella equilib a – 35 ––––– – – – – – – –––– – –
Cap ella japonica – – ––––– – – – – – – 411––– – –
Cap ella lae iuscula – – – – – – – 908 1,000 – – – – – – – – – –
Cap ella linea is – – ––––– – – – 80 – – –––– – –
Cap ella penan is – 166 – – – – – – – – – – – – – – – – –
Cap ella polyacan ha – – ––––– – – – – – – –10–– – –
Cap ella scau a – – ––––– – – – – – – –107.1– – –
Cap ella sp aV. b e i os is – – ––––– – – – – – – 21––– – –
Cap ella sp1 – – – – – – – – – 250 – 443 298 – – – – – –
Cap ella sp2 – – ––––– – – – – – – ––1.8– – –
Cap ella subine mis – – ––––– – – – – – – 8.3––– – –
Cap ella suga ensis – – ––––– – – – – – – –3.841– – –
Cap ella e ucosa – – ––––– – – – – – – 4.2––– – –
Cap ellids uniden i . – – – – – – 144 – – – 26 – – – – – – – –
Pa acap ella enuis 45 27 ––––– – – – – – – –––– – –
Pa iambus ypicus – – – – – – – – – – – – – – – – 156 82.2 –
Ph isica ma ina – – 4.3 1.3 – – – – – – – – – – – – 37 481 90
Pseudop o ella phasma – – 0.1 0.7 – – – – – – – – – – – – 44 3.3 –
To al numbe o species 1 3 3 3 1 1 ? 1 1 1 3 1 1 5 4 3 4 3 1
To al abundance (Ind/m2) 45 228 ? ? 6 17 144 908 1,000 250 132 443 298 449 64 50 239 557 90
Helgol Ma Res (2008) 62:189–199 197
123
p oW ing om pe u ba ions o so bo oms, such as d edging
o awling.
La i udinal pa e ns o cap ellid dis ibu ion on seag ass
meadows
Al hough he biogeog aphical dis ibu ion o cap ellid
species on a global scale is no well known so a , i has
been epo ed ha he highes cap ellid species di e si y is
ound in empe a u e wa e s (a ound 30° la i ude), and ha
species di e si y dec eases bo h owa ds colde wa e s o
highe la i udes and owa ds he equa o (Laubi z 1970;
Abele 1982; Thiel e al. 2003). This pa e n has been also
measu ed o gamma idean amphipods. Howe e , he ypi-
cal pa e n o many o he g oups o c us aceans such as
os acods, copepods, s oma opods o decapods is diVe en :
a dec ease o species ichness om he equa o owa ds
highe la i udes (Abele 1982). I only seag ass meadows
a e conside ed, he amphipod pa e n changes as di e si y
inc eases signiWcan ly wi h dec easing la i ude, being
maximal nea he equa o (Vi ns ein e al. 1984). These
au ho s summa ised he a ailable in o ma ion o de e mine
whe he any la i udinal pa e n exis s o he seag ass-
associa ed epi auna, and o examine hypo heses, which
migh explain he obse ed pa e ns. They ound ha he
di e si y o decapods and amphipods in seag ass meadows
inc eases wi h dec easing la i ude, while he di e si y o
isopods and Wshes showed nonsigniWcan ends wi h
la i ude. In e es ingly, densi y o amphipods showed no
pa e n wi h la i ude, which is appa en ly inconsis en wi h
ei he he opical dec ease o inc ease p edic ed by he
p eda ion and p ima y p oduc ion hypo heses. Consequen ly,
Vi ns ein e al. (1984) poin ed ou ha , con a y o e i-
dence om o he biological sys ems, i appea s ha
la i ude is, in gene al, an inconsis en p edic o o diVe ences
in he epi auna o seag ass communi ies. I we conside
only he cap ellidean amphipods, species ichness and
abundances ob ained in he p esen s udy a e simila o
hose measu ed in o he seag ass meadows a ound he
wo ld, and da a o Table 4 suppo he idea ha he e is no
clea la i udinal pa e n o densi y alues (in ac , he
co ela ion be ween la i ude alues o locali ies in Table 4
and cap ellid densi ies is no signiWcan , = 0.19, NS).
Da a o o al cap ellid abundance ange om 6 o
1,000 indi iduals/m2, bu he me hods diVe ed among
s udies, which makes compa isons diYcul . Fu he mo e,
he e is a lack o da a o a eas nea he equa o and u he
s udies a e necessa y.
Fi
g.
8S
c
h
ema
i
c ep esen a-
ion o he e ical dis ibu ion o
cap ellids in he h ee seag ass
meadows; AL Alme ía,
M
A Málaga, CA Cádiz