Community structure of caprellids (Crustacea: Amphipoda: Caprellidae) on seagrasses from southern Spain

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