Biodiversity and distribution of macroinfauna assemblages on sandy beaches along the Gulf of Cadiz (SW Spain)

Biodi e si y and dis ibu ion o mac oin auna assemblages on sandy beaches along he Gul o Cadiz (SW Spain) M. José Reyes-Ma ínez 1, M. Ca men Ruiz-Delgado 1, Juan Emilio Sánchez-Moyano 2, F ancisco José Ga cía-Ga cía 1 1 Depa amen o de Sis emas Físicos, Químicos y Na u ales, Uni e sidad Pablo de Ola ide, C a. U e a km 1, 41013 Se illa, Spain. E-mail: [email p o ec ed] 2 Depa amen o de Zoología, Uni e sidad de Se illa, A . Reina Me cedes 6, 41012 Se illa, Spain. Summa y: In he cu en s udy, he mac oin auna communi ies inhabi ing 12 sandy beaches along he Spanish coas o he Gul o Cadiz we e desc ibe in o de o cha ac e ize hei unexplo ed biodi e si y, hei zona ion pa e ns and hei en i on- men al ea u es. A o al o 66 mac oin auna axa we e eco ded on 12 sandy beaches, anging om 4 o 33 species. Densi y eached 848 indi idual pe m2. The indi idual zona ion pa e n anged om wo o h ee zones, ega dless o he mo phody- namic s a e. A common zona ion pa e n o he whole se o beaches was es ablished, comp ising h ee ac oss-sho e biologi- cal zones. Gene ally, he sup ali o al zone was ypi ied by he ai -b ea hing amphipod Tali us sal a o and coleop e ans; he middle zone was domina ed by ue in e idal species, such as Haus o iidae amphipods (Haus o ius a ena ius), Ci olanidae isopods (Eu ydice a inis), Spionidae polychae es (Scolelepis spp.) and neme eans; and he lowe o subli o al zone was ypi ied by Pon opo eiidae amphipods, mysids and spionid polychae es. Sedimen mois u e, a e age g ain size, o ganic ma - e con en and ele a ion we e he main p edic o a iables o zona ion pa e ns. We used he conse a ion and ec ea ion po en ial indices o show how his in o ma ion can be use ul o coas al managemen . Mos beaches s udied show a sco e abo e 5 on he conse a ion index, so he beaches ha e a easonable po en ial o conse a ion managemen . Keywo ds: sandy beaches; ben hic mac oin auna; zona ion pa e n; en i onmen al a iables; Gul o Cadiz, coas al manage- men . Biodi e sidad y dis ibución de las comunidades de mac oin auna en playas a enosas del Gol o de Cádiz (SO España) Resumen: En el p esen e es udio, se desc iben las comunidades de mac oin auna de doce playas a enosas de la cos a es- pañola del Gol o de Cádiz con el obje i o de ca ac e iza su biodi e sidad, sus pa ones de zonación, y sus ca ac e ís icas ambien ales. Un o al de 66 axones de mac oin auna ue on egis ados en las playas de es udio, oscilando en e 4 y 33 espe- cies, mien as que la densidad o al ue de 848 indi iduos po m2. El pa ón de zonación indi idual a ió de dos a es zonas, independien emen e del es ado mo odinámico de la playa. Se es ableció un pa ón de zonación común de odo el conjun o de playas, o mado po es zonas biológicas. En gene al, la zona sup ali o al ue ipi icada po el an ípodo Tali us sal a o y di e sas amilias de coleóp e os. La zona media ue dominada po e dade as especies in e ma eales, como an ípodos de la amila Haus o iidae (Haus o ius a ena ius), isópodos ci olánidos (Eu ydice a inis), polique os (Scolelepis spp.) y neme - inos. La zona subli o al se ca ac e izó po an ípodos de la amilia Pon opo eiidae, misidáceos y polique os espiónidos. La humedad del sedimen o, el amaño medio de g ano, el con enido en ma e ia o gánica y la ele ación ue on las p incipales a iables in luyen es en los pa ones de zonación. Po úl imo, oda es a in o mación ue in eg ada pa a la aplicación de los índices de Conse ación y Rec eación, pa a mos a cómo es a in o mación puede se ú il pa a la ges ión cos e a. La mayo ía de las playas mos a on una pun uación supe io a 5 en el Índice de Conse ación, indicando un azonable po encial pa a la conse ación. Palab as cla e: playas a enosas; mac oin auna bén ica; pa ones de zonación; a iables ambien ales; Gol o de Cádiz; ges ión cos e a. Ci a ion/Como ci a es e a ículo: Reyes-Ma ínez M.J., Ruíz-Delgado M.C., Sánchez-Moyano J.E., Ga cía-Ga cía F.J. 2015. Biodi e si y and dis ibu ion o mac oin auna assemblages on sandy beaches along he Gul o Cadiz (SW Spain). Sci. Ma . 79(3): 000-000. doi: h p://dx.doi.o g/10.3989/scima .04133.30A Edi o : R. Sa dà. Recei ed: July 31, 2014. Accep ed: May 22, 2015. Published: July 3, 2015. Copy igh : © 2015 CSIC. This is an open-access a icle dis ibu ed unde he C ea i e Commons A ibu ion-Non Com- me cial Lisence (by-nc) Spain 3.0. Scien ia Ma ina 79(3) Sep embe 2015, 000-000, Ba celona (Spain) ISSN-L: 0214-8358 doi: h p://dx.doi.o g/10.3989/scima .04133.30A 2 • M.J. Reyes-Ma ínez e al. SCI. MAR., 79(3), Sep embe 2015, 000-000. ISSN-L 0214-8358 doi: h p://dx.doi.o g/10.3989/scima .04133.30A INTRODUCTION The Gul o Cadiz is loca ed in he sou hwes e n Ibe ian Peninsula, be ween Cape S Vincen (Po ugal) and he S ai o Gib al a (Spain), which connec s he A lan ic Ocean and he Medi e anean Sea. The Span- ish coas al a ea o his gul s e ches some 300 km, be- ween Ayamon e (Huel a p o ince) and Ta i a (Cadiz p o ince). The a ea is in luenced mainly by he mou hs o he i e s Guadiana, Pied as, Tin o, Odiel, Guadale e and Guadalqui i and is domina ed by es ua ine zones and ex ensi e sandy beaches, many o which a e aced by discon inuous ocky-sho e pla o ms (Bena en e e al. 2002), especially on he Cadiz coas . The gene al ai ci cula ion in he Gul o Cadiz is p edominan ly an icyclonic, wi h sho - e m a ia ion in luenced by winds. This egion is cha ac e ized by a mean wa e -su ace empe a u e anging om 18°C o 22°C, a salini y ange o 36.3‰ o 36.5‰, an a e age ni a e, phospha e and silica e concen a ion o abou 0.33, 0.08 and 1.37 μM, espec i ely (An uso e al. 2010), and a chlo ophyll a concen a ion o a ound 10- 40 mg m–2 (P ie o e al. 1999). These ea u es p o ide a sui able habi a o he de elopmen o se e al species, which makes his sys em a e y di e se and p oduc i e a ea (Sob ino e al. 1994). Many species inhabi ing he Gul o Cadiz ha e economic alue, so i is conside ed an a ea wi h g ea socio-economic impo ance o ishe ies and shell ish ga he ing (To es e al. 2013). F equen ly, hese species use sandy sho es as nu se y a eas o ju eniles (Baldó and D ake 2002), eeding on in e eb a es (Speyb oeck e al. 2007), and hey can use biogenic s uc u es (e.g. ubes, mounds and bu - ows) cons uc ed by he in e eb a es as a e uge om p eda ion (Allen-B ooks e al. 2006). The sandy sho es p o ide a la ge ange o se ices, such as sedimen and wa e s o age, decomposi ion o o ganic ma e and pollu an s, wa e dissipa ion, wa e il a ion and pu i ica ion, nu ien ecycling, main e- nance o biodi e si y and a unc ional link be ween ma ine and e es ial en i onmen s, whe e mac oin- auna plays a key ole (De eo e al. 2009). Fu he mo e, he a ou able clima ic condi ions in Spain make he coas al en i onmen s a ac i e o ou ism o se e al mon hs a yea and beaches a e a majo economic e- sou ce (An uso e al. 2003). Despi e he impo ance o sandy beaches and he scale o he coas line a eas occupied in he s udy, da a on bio ic and abio ic cha ac e is ics a e scan . Along he Spanish Gul o Cadiz, pas s udies ha e ocused on s udying he mo phodynamics o sandy beaches in es ic ed a eas (An uso e al. 2003) and he mo - phological changes associa ed wi h me eo ological e en s (Bui ago and An uso 2011). The ew s udies ha ha e desc ibed he auna inhabi ing he beaches ha e ocused on mac oin auna om es ua ine beaches (Mayo al e al. 1994) o on he sup ali o al a h opods associa ed wi h w ack deposi s (Ruiz-Delgado e al. 2014). The e is he e o e a conside able lack o in o - ma ion on mac oin auna communi ies in his egion. Inc easing human in e es in sandy beaches has ex- ace ba ed unsus ainable p ac ices ha in ol e des uc- ion o hese na u al en i onmen s (De eo e al. 2009). Iden i ying and mapping he mac oin auna inhabi ing sandy beaches is an essen ial s ep o es ablishing e - ec i e managemen and conse a ion ools o a be - e use o hese ma ine en i onmen s (McLachlan e al. 2013, Role e al. 2015, Schlache e al. 2014) and o es ima ing he po en ial esponse o bio a o u u e habi a changes. The aim o his s udy is o p o ide he i s desc ip- ion o mac oin auna communi ies inhabi ing sandy beaches and hei en i onmen al cha ac e is ics. To his end, (1) he physical and mo phodynamic cha ac- e is ics o 12 sandy beaches along he Gul o Cadiz coas line we e de ined; (2) he mac oin auna commu- ni ies inhabi ing sandy beaches we e cha ac e ized; (3) he zona ion pa e n o mac oin auna was de e mined; and (4) he in luence o en i onmen al ac o s on he zona ion pa e ns was explo ed. Finally, all his in o ma ion was employed o cal- cula e wo quan i a i e indica o s ollowing McLachan e al. (2013). These indices a e used o assess beach condi ion and sui abili y o di e en uses, om which guidelines o planning, managemen and moni o ing can be de i ed. MATERIALS AND METHODS S udy a ea The s udy a ea comp ised 12 sandy beaches along he Spanish coas o he Gul o Cadiz, om Hoyo beach (37°11’55”N, 07°17’45”W) nea he Po uguese bo - de o Lances beach (36°02’31”N, 05°38’08.031”W) in he a ea nea he S ai o Gib al a (Fig. 1). Sampling p ocedu es Fo all 12 beaches, mac oin auna sampling was ca - ied ou in six eplica ed ansec s haphaza dly sepa a - ed by a 100-m-long s e ch in he cen al a ea o each beach du ing low ide be ween Ma ch and May 2011. In each ansec , mac oin auna samples we e col- lec ed a 10 equally spaced sho e le els ex ending om abo e he d i line (le el 10) o he low swash zone (le el 1) wi h a 25-cm-diame e plas ic co e o a dep h o 20 cm. T ansec leng h depended on each beach’s wid h. A o al o 60 samples we e collec ed wi hin a o al sam- pled a ea o 3.75 m2 pe beach. On empe a e beaches his a ea is conside ed su icien o collec 90% o all he mac oin auna (Ja amillo e al. 1995). Samples we e sie ed on si e h ough a 1-mm mesh sie e, collec ed in labelled plas ic bags and p ese ed in 70% e hanol s ained wi h Rose Bengal. Addi ionally, one sedimen sample was aken a each sampling le el wi h a plas ic ube (3.5 cm diame e ) bu ied 15 cm deep o analyse he mean g ain size, so ing coe icien (T ask 1950), sand mois u e and o ganic ma e o he sedimen . In he labo a o y, he mac oin auna was quan i ied and iden i ied o he lowes axonomic le el possible. The mean g ain size was de e mined ollowing he me hod p oposed by Gui ián and Ca ballas (1976). This me hod disc imina es di e en g anulome ic ac ions Mac oin auna assemblages on sandy beaches • 3 SCI. MAR., 79(3), Sep embe 2015, 000-000. ISSN-L 0214-8358 doi: h p://dx.doi.o g/10.3989/scima .04133.30A when he sedimen composi ion is mainly sand and he peli ic ac ion is low (less han 5%). Sand mois u e was de e mined by measu ing he weigh loss a e d y- ing he samples a 90°C. The o ganic ma e con en o he sedimen ac ion <0.5 mm was es ima ed as he di e ence be ween d y sedimen weigh and sedimen weigh a e calcina ion a 500°C. To cha ac e ize he mo phodynamic s a e, he ela- i e idal ange (Masselink and Sho 1993), he beach index (McLachlan and Do lo 2005), he beach s a e index (McLachlan e al. 1993) and he dimensionless all- eloci y pa ame e (Dean’s pa ame e ; Dean 1973) we e used. The beach ace slope was es ima ed by he heigh di e ence acco ding o Eme y (1961). The heigh and wa e pe iod, empe a u e and idal ange we e aken om a e age annual da a om he SIMAR oceanog aphic da abase o Pue os del Es ado (Spanish Minis y o Public Wo ks) (h p://www.pue os.es/es- es/oceanog a ia/Paginas/po us.aspx). Conse a ion and ec ea ion po en ial indices To cha ac e ize he main use o beaches, he conse - a ion index (CI) and he ec ea ion po en ial index (RI) we e applied ollowing McLachlan e al. (2013). Fo each indica o a simple sco ing sys em was de eloped based on h ee key d i e s in o de o e alua e he po- en ial/sui abili y o a beach o conse a ion and/o ec- ea ion. CI akes in o accoun 1) he ex en , na u e and condi ion o he dunes, hei well-de eloped ege a ion and hei connec ion wi h he beach; 2) he p esence o iconic and endange ed species; and 3) he mac oben hic communi y abundance and species ichness. In con as , RI is based on 1) a ailable in as uc u es o suppo ec- ea ional ac i i ies (beach access, oile s, e c.); 2) beach sa e y and heal h s a us; and 3) physical ca ying capac- i y. CI and RI alues ange om 0 o 10, in inc easing o de o conse a ion alue o ec ea ion po en ial. Da a analysis Uni a ia e analyses we e used o cha ac e ize he auna communi ies p esen on each beach s udied, cal- cula ing he Ma gale ichness index (d), he Shannon- Wiene di e si y index (H’) and he Pielou e enness index (J’), using he PRIMER so wa e package. The zona ion pa e n on each beach s udied was iden i ied using clus e analysis, based on he B ay-Cu - is simila i y ma ix, ollowed by a simila i y p o ile es (SIMPROF) (Cla ke and Go ley 2006), o e alua e he signi icance o he classi ica ion (p<0.05). P e iously, abundance da a we e ou h- oo ans o med o down- weigh he con ibu ion o he majo abundan species. Once he zona ion pa e ns had been de ined on indi idual beaches, a modal pa e n o zona ion was es ablished o he en i e se o beaches. To his end, species om each sampling le el we e pooled based on zones iden i ied by clus e analysis. Then, a single ma ix o “species × zone” o each beach was gene a ed and all o hem we e combined in o a global ma ix. This global biological ma ix was ou h- oo ans o med and subjec ed o non-me ic mul i-dimensional scaling o dina ion (n-MDS). Fu - he mo e, he simila i y pe cen ages analysis (SIM- PER) was pe o med in o de o ind he ypi ying species in each iden i ied zone o he en i e se o beaches along he Gul o Cadiz. Beaches ha did no ha e a clea zona ion pa e n we e excluded om hese analyses. All mul i a ia e analyses we e pe o med wi h PRIMER-E 6.1 (PRIMER-E L d) (Cla ke and Wa wick 2001). To de e mine associa ions o mac oin auna com- muni ies wi h en i onmen al a iables, a canonical co espondence analysis (CCA) was applied (Te B aak 1986). Fi s , a global biological ma ix was submi ed o de ended co espondence analysis (DCA) in o de o measu e he g adien leng hs and o ensu e an uni- modal species esponse. G adien leng h o he i s axis was g ea e han 3.0 SD, and a CCA o dina ion me hod was used. Fo his analysis, only he mos abundan species (>6% o o al con ibu ion in each biological zone iden i ied) we e aken in o accoun a e ou h- oo ans o ma ion. The en i onmen al pa ame e s ma ix was ans- o med (Log(x+1)) and s anda dized p io o educing ex eme alues and p o iding be e canonical coe i- cien compa isons. Only a iables signi ican ly ela ed o he auna a ia ion we e included (p<0.05); o his end, each a iable was analysed sepa a ely and i s sig- ni icance was es ed using a Mon e Ca lo pe mu a ion es (999 pe mu a ions) (Te B aak 1995). In CCA analysis, he s a is ical signi icance o ca- nonical eigen alues and he signi icance o he i s wo axes we e es ed by a Mon e Ca lo es (999 pe - mu a ions). DCA and CCA we e ca ied ou wi h he s a is ical so wa e package PC-ORD (McCune and Med o d 1997). RESULTS Beach cha ac e is ics The physical and mo phodynamic cha ac e is ics o he 12 beaches s udied a e shown in Table 1. Acco ding Fig. 1. – S udy a ea showing he 12 sandy beaches sampled. 4 • M.J. Reyes-Ma ínez e al. SCI. MAR., 79(3), Sep embe 2015, 000-000. ISSN-L 0214-8358 doi: h p://dx.doi.o g/10.3989/scima .04133.30A o he idal ange and ela i e idal ange, he beaches we e ca ego ized as meso idal, domina ed by wa es. The beaches showed a wide ange o mo phodynamic ypes, classi ied by Dean’s pa ame e as in e media e (Te ón, Mazagón, Ma alascañas, Ba osa and Zaha a), dissipa i e (Bo a, Cos a Ballena, Valdelag ana, Le- an e, Co adu a, and Lances) and e lec i e (Hoyo). Beach s a e index alues classi ied mos beaches as in e media e o dissipa i e, wi h high ene gy le els, excep o Hoyo and Zaha a, which we e in e media e beaches wi h low o medium ene gy le els. Mac oin auna A o al o 63 mac oin auna axa we e eco ded om he beaches o he Gul o Cadiz (Appendix 1). C us a- ceans we e he mos di e se axa, wi h 23 species, ol- lowed by polychae es (22 species), insec s and molluscs (9 and 8 species, espec i ely). Appendix 1 shows he o- al abundance, o al species, Ma gale species ichness, Shannon-Wiene di e si y index and Pielou e enness index. Bo a and Le an e had he highes ichness, wi h 33 and 24 species, espec i ely, while he lowes alue Table 1. – Physical cha ac e iza ion o sampled beaches. L, beach leng h (m); Mgs, median g ain size (mm); OM, o ganic ma e con en (%); RTR, Rela i e ide ange; BI, beach index; BSI, beach s a e index. Beaches L Slope(1/m) Mgs Sand ype So ing Dean RTR BI BSI OM Hoyo 2800 10.99 0.65 coa se sand 1.54 1.6 2.27 1.36 0.92 0.62 Te ón 3500 29.52 0.42 medium sand 1.45 2.53 2.27 2.09 1.09 0.48 Bo a 3800 46.59 0.22 ine sand 1.33 5.23 2.7 2.51 1.36 0.89 Mazagón 5500 15.84 0.49 medium sand 1.57 2.1 2.41 1.75 1.05 0.62 Ma alascañas 4200 13.97 0.41 medium sand 1.34 2.59 2.34 1.77 1.1 0.31 Cos a Ballena 4500 29.99 0.23 ine sand 1.35 5.91 2.27 2.31 1.43 0.68 Valdelag ana 1880 17.69 0.21 ine sand 1.6 6.8 2.28 2.11 1.48 1.19 Le an e 4600 26.46 0.22 ine sand 1.43 6.32 2.49 2.25 1.42 0.75 Co adu a 2500 84.31 0.20 ine sand 1.25 7.73 2.02 2.81 1.55 0.81 Ba osa 4000 17.6 0.47 medium sand 1.55 2.42 2.05 1.76 1.03 2.92 Zaha a 2900 11.5 0.51 coa se sand 1.75 2.26 1.58 1.43 0.93 0.83 Lances 4300 24.76 0.23 ine sand 1.35 6.41 1.07 1.94 1.19 0.57 Fig. 2. – Zona ion pa e n in each s udied beach de ined by simila i y p o ile (SIMPROF). G ey lines ep esen signi ican e idence o com- muni y s uc u e (P<0.05). Black lines indica e no signi ican e idence. Mac oin auna assemblages on sandy beaches • 5 SCI. MAR., 79(3), Sep embe 2015, 000-000. ISSN-L 0214-8358 doi: h p://dx.doi.o g/10.3989/scima .04133.30A was ound a Ma alascañas (4 species). Densi y was also highly a iable, anging om 23 o 249 indi iduals pe m2. The lowes di e si y alue (H’) was obse ed a Ma alascañas beach (0.40), while he highes was ound a Le an e (2.68). The e enness index anged om 0.29 a Ma alascañas o 0.86 a Lances beach. Scolelepis spp., Pon oc a es a ena ius and Haus- o ius a ena ius we e he mos equen species. The polychae e was p esen on 100% o he o al beaches sampled, while he amphipods we e p esen on 90%. In e ms o abundance, polychae es o he genus Scolelepis we e dominan , accoun ing o 28% o o al abundance, ollowed by he amphipod Haus o ius a e- na ius, which accoun ed o 15% o he o al. Zona ion Ac oss-sho e species dis ibu ion on each beach s udied is shown in Figu e 2. Clus e o dina ion and he SIMPROF es iden i ied wo biological zones a Valdelag ana and Co adu a and Lances, and h ee bio- logical zones a Hoyo, Te ón, Bo a, Mazagón, Cos a Ballena, Le an e, Ba osa and Zaha a. Excep ionally, Ma alascañas did no show a clea zona ion pa e n. Fo his analysis, he sampling le els whe e no species we e p esen we e emo ed. A global zona ion pa e n o he en i e se o beaches om he Spanish Gul o Cadiz coas line was de i ed om he indi idual ac oss-sho e species dis ibu ion; Fig. 3. – n-MDS o dina ion o he global zona ion pa e n. Black iangles ep esen he lowe zones, g ey in e ed iangles he middle zones and black quad a es he uppe zones o he s udied beaches. Fig. 4. – Mean o al densi y (±SE) o he axa ound in he lowe , middle and uppe zones. Fig. 5. – T iplo esul ing om CCA analysis. C osses show he mos abundan species in each zone. The lowe zones a e ep esen ed by i- angles, he middle zones by in e ed iangles and he uppe zones by ci cles. A ows ep esen s explana o y a iables (Mois , sand mois u e; Mgs, median g ain size; Ele , ele a ion; OM, o ganic ma e con en ). 6 • M.J. Reyes-Ma ínez e al. SCI. MAR., 79(3), Sep embe 2015, 000-000. ISSN-L 0214-8358 doi: h p://dx.doi.o g/10.3989/scima .04133.30A he e o e, aunal zones iden i ied a each beach we e ga he ed o a global MDS o dina ion (Fig. 3). SIMPER analysis pe o med on his o dina ion showed a deg ee o simila i y o 40% be ween all lowe zones, whe e Pon- oc a es a ena ius, Gas osaccus sanc us and Scolel- epis spp. showed he highes pe cen ages o p esences (17.8%, 17.2% and 11.0%, espec i ely). The middle zones showed a simila i y o abou 30%. Polychae es o he genus Scolelepis (37.70%), he isopod Eu ydice a inis (26.40%), he amphipod Haus o ius a ena ius (11.56%), and neme eans (9.95%) highligh ed he simi- la i y in aunal composi ion be ween all middle zones. Finally, he uppe zones showed a 20% simila i y and he ypi ying species we e he ai -b ea hing amphipod Tali us sal a o (56.7%) and coleop e ans (37%). Biologically, densi y alues dec eased om he lowe o he uppe zone. In he lowe and middle zones he mos abundan axa we e c us aceans and poly- chae es, while in he uppe zones, besides c us aceans, insec s we e dominan (Fig. 4). Rela ionship be ween en i onmen al a iables and mac oin auna En i onmen al a iables signi ican ly ela ed o auna a ia ions, es ed by Mon e Ca lo pe mu a- ion es , we e ele a ion (p=0.002), sand mois u e (p=0.001), o ganic ma e con en (p=0.015) and g ain size (p=0.001). Howe e , hese p edic o a iables we e no s ongly co ela ed ( 2<0.5). The Mon e Ca lo es o he se o en i onmen al a iables was signi ican o bo h axes (p=0.001) and o eigen alues (p=0.001) showing a signi ican ela ionship be ween biological da a and p edic o en i onmen al a iables. Table 2. – Conse a ion index (CI) and ec ea ion po en ial index (RI) sco es o beaches. Beach name Dune s a us Iconic species Mac o-ben hos CI In as uc u e Sa e y heal h Ca ying capaci y RI Hoyo 4 1 1 6 1 3 2 6 Well de eloped, li le dis u bance Low numbe s, no nes ing In e media e No in as uc u e, limi ed access Low haza d, clean and pollu ion ee Ex ensi e beach wi h wide backsho e Te ón 4 2 1 7 1 3 1 5 Well de eloped, li le dis u bance P esen in good numbe s, may be nes ing In e media e No in as uc u e, limi ed access Low haza d, clean and pollu ion ee In e media e Bo a 4 1 2 7 1 3 2 6 Well de eloped, li le dis u bance Low numbe s, no nes ing Species ich and abundan No in as uc u e, limi ed access Low haza d, clean and pollu ion ee Ex ensi e beach wi h wide backsho e Mazagón 4 2 1 7 1 2 1 4 Well de eloped, li le dis u bance P esen in good numbe s, may be nes ing In e media e No in as uc u e, limi ed access Mode a e haza d and clean In e media e Ma alascañas 0 0 1 1 5 2 0 7 Absen Absen In e media e Excellen access and ameni ies Mode a e haza d and clean Limi ed, pocke beach, no backsho e Cos a Ballena 3 1 2 6 3 3 2 8 Dis u bed bu la gely in ac Low numbe s, no nes ing Species ich and abundan Good access, some ameni ies Low haza d, clean and pollu ion ee Ex ensi e beach wi h wide backsho e Valdelag ana 0 1 2 3 5 3 1 9 Absen Low numbe s, no nes ing Species ich and abundan Excellen access and ameni ies Low haza d, clean and pollu ion ee In e media e Le an e 4 3 2 9 1 3 1 5 Well de eloped, li le dis u bance Signi ican nes ing a ea o ma ine bi ds Species ich and abundan No in as uc u e and limi ed access Low haza d, clean and pollu ion ee In e media e Co adu a 1 0 2 3 2 2 1 5 Se e ely dis u bed Absen Species ich and abundan Modes in as uc u e, easonable access Mode a e haza d and clean In e media e Ba osa 2 0 2 4 2 3 2 7 Ex ensi e dis u bance Absen Species ich and abundan Modes in as uc u e, easonable access Low haza d, clean and pollu ion ee Ex ensi e beach wi h wide backsho e Zaha a 4 2 1 7 2 2 2 6 Well de eloped, li le dis u bance P esen in good numbe s, may be nes ing In e media e Modes in as uc u e, easonable access Mode a e haza d and clean Ex ensi e beach wi h wide backsho e Lances 4 2 1 7 2 2 2 6 Well de eloped, li le dis u bance P esen in good numbe s, may be nes ing In e media e Modes in as uc u e, easonable access Mode a e haza d and clean Ex ensi e beach wi h wide backsho e Mac oin auna assemblages on sandy beaches • 7 SCI. MAR., 79(3), Sep embe 2015, 000-000. ISSN-L 0214-8358 doi: h p://dx.doi.o g/10.3989/scima .04133.30A The CCA esul s show ha he o al a ia ion o da a was 2.49 (ine ia), while he o al a ia ion explained was 0.802 ( he sum o all canonical eigen alues). Pea - son species-en i onmen al co ela ions we e ela i ely high: 0.93 o Axis 1 and 0.82 o Axis 2. The i s axis (accoun ing o 66% o he o al explained a ia- ion) co ela ed posi i ely wi h ele a ion (0.745) and nega i ely wi h sand mois u e (–0.887) and o ganic ma e con en (–0.465). The second axis (accoun ing o 0% o he explained a ia ion) co ela ed mainly wi h medium-sized g ain (0.806). The o dina ion diag am o CCA (Fig. 5) showed a g adien o zones (lowe , middle, and uppe ) ma ked mainly by he i s axis and showed ha c us aceans (Ba hypo eia pelagica, Eu ydice a inis, E. pulch a, Gas osaccus sanc us, G. spini e , Haus o ius a e- na ius and Pon oc a es a ena ius) and Polychae a (Scolelepis spp.) esponded posi i ely o sand mois u e and o ganic ma e con en bu nega i ely o ele a- ion, inc easing hei densi y o he le along he i s axis. Coleop e ans and Tali us sal a o exhibi ed he opposi e pa e n. Densi y o neme eans was he leas explained by hese en i onmen al a iables. The neme eans P. a ena ius and G. sanc us also e- sponded posi i ely o medium-coa se g ain size, while he densi y o Ba hypo eia pelagica, Donax unculus and Coleop e a sp1 was mo e in luenced by ine g ain size, due o hei dis ibu ion along he second axis. Conse a ion agains ec ea ional po en ial Table 2 p o ides a summa y o 12 cases o beach usage by means o CI and RI sco es and he ensuing managemen s a egy is illus a ed in Figu e 6. Mos beaches a e in he mul iple-use ca ego y, indica ing ha hey ha e high conse a ion and ec ea ion alues. Only Ma alascañas and Valdelag ana a e in he in en- si e ec ea ion ca ego y, wi h high RI alues (7 and 9, espec i ely). DISCUSSION Mac oin auna Sandy beaches a e ex emely dynamic ecosys ems wi h hos ile condi ions o li e, so he numbe o axa adap ed o li ing unde hese condi ions is expec ed o be low compa ed wi h o he coas al sys ems. Howe e , he mac oin auna biodi e si y ound in sandy beaches o he Spanish coas o he Gul o Cadiz we e ela- i ely high ( om 4 o 33 species). This alue is simila o hose epo ed in nea by la i udes, such as he 9 o 31 species ound in no he n Spain (Rodil e al. 2006), he 27 o 30 species ound on he wes e n coas o Po ugal (Gonçal es e al. 2009), and he 4 o 24 species ound on Mo occan sandy beaches (Bayed 2003). The beaches showed a wide ange o mo phody- namic ypes and a gene al inc easing end in species ichness om e lec i e o dissipa i e beaches was obse ed, in ag eemen wi h McLachlan e al. (1993). Bo a beach showed he highes species ichness and abundance. This beach, loca ed nea he mou h o he Pied as Ri e , whe e he in luence o wa e ac ion is lowe , is one o he mos shel e ed o he en i e se o beaches s udied, as is also e lec ed in he ela i e idal ange, which showed high alues on his sandy beach. The high ichness and abundance alues ound on Bo a beach suppo s he gene al end o bio ic a iables o inc ease wi h exposu e, as epo ed by o he au ho s (Dex e 1983, Ja amillo and MacLachlan 1993, Rodil e al. 2007). Abundance and ichness o mac oin auna a e di ec ly ela ed o ood supply (Rodil e al. 2012), so i is also possible ha Pied as i e mou h inc eases ood a ailabili y, enabling he es ablishmen and de- elopmen o mo e species. C us aceans, polychae es and molluscs we e usual- ly dominan among he mac oin auna o sandy beaches (McLachlan and B own 2006). In ou s udy, amphipod and isopod c us aceans and spionid polychae es we e he mos abundan and included di e se axa. In ac , 74% o all indi iduals collec ed belong o six species o hese g oups: Ba hypo eia pelagica, Haus o ius a ena ius, Pon oc a es a ena ius, Siphonoece es saba- ie i, Eu ydice a inis and Scolelepis spp. The dominan and mos equen species occu ing on e e y beach s udied we e polychae es o he genus Scolelepis. This genus has a wide geog aphical dis ibu ion (Souza and Bo zone 2000) and is also e y abundan on many A - lan ic beaches (Papageo giou e al. 2006, Deg ae e al. 2003, Ba os e al. 2001). Li le impo ance is gi en o neme eans, which a e no mally no conside ed ypical axa on sandy beaches, due o esidual con ibu ions ha hey exhibi , al hough his axon is conside ed a use ul bio-indica o (He e a- Bachille e al. 2008). Fo example, he neme eans Lienus acu i ons and Psammamphipo us elonga us we e a ec ed by he “P es ige” oil spill on Galician Fig. 6. – Loca ion o each beach lis ed in Table 2 ela i e o i s conse a ion index and ec ea ion po en ial index sco es. The plo shows he dema ca ion o beaches o in ensi e ec ea ion, conse - a ion and mul i-pu pose use. H, Hoyo; T, Te ón; Bo, Bo a; M, Mazagón; Ma, Ma alascañas; Cb, Cos a Ballena; V, Valdelag ana; Le, Le an e; C, Co adu a; Ba, Ba osa; Z, Zaha a; La, Lances. 8 • M.J. Reyes-Ma ínez e al. SCI. MAR., 79(3), Sep embe 2015, 000-000. ISSN-L 0214-8358 doi: h p://dx.doi.o g/10.3989/scima .04133.30A sandy beaches, so hey could be a good indica o s o beach quali y (He e a-Bachille e al. 2008, Pue a e al. 2010). On sandy beaches o sou hwes e n Spain, Neme - ean abundance was simila o ha o molluscs, showing high occu ences (67% o he sampled beaches), high- ligh ing he impo ance o neme eans a hese la i udes. Simila ly, Tali us sal a o was equen ly ound on he beaches s udied. This ‘sand-hoppe ’ is ecognized as a good biomoni o o ace-me al pollu ion and he e ec o human ampling (Ugolini e al. 2008). Mac oin auna zona ion The zona ion o mac oin auna, de ined as he dis- ibu ion o species h oughou he in e idal zone on sandy beaches, has been unde aken a ound he wo ld (De eo e al. 1992, Ja amillo e al. 2000, Schlache and Thompson 2013). Mac oin auna ac oss-sho e dis- ibu ion is highly a iable, anging om one o i e biological zones, al hough h ee a e mos common (see Schlache and Thompson 2013). In he cu en s udy, 67% o o al beaches showed h ee dis inc zones and 25% showed wo. This pa e n was di e en o ha ound on beaches in no he n Spain, whe e mac oin- auna was di ided in o wo main zones (Rodil e al. 2006), and on Mo occan sandy beaches, whe e ou zones we e iden i ied acco ding o Sal a ’s scheme (Sal a 1964, Bayed 2003). Ja amillo e al. (1993) de e mined ha in e medi- a e and dissipa i e beaches include h ee aunal zones, whe eas he e lec i e beaches had only wo. Along he Spanish coas o he Gul o Cadiz, his pa e n was no ound. In ac , he mo e dissipa i e beaches showed wo biological zones, while beaches closes o he e- lec i e s a e (Hoyo and Mazagón) had h ee zones. In gene al, he numbe o zones al e na ed, indepen- den ly o Dean’s pa ame e . Thus, no clea e idence was ound o suppo he con en ion ha he numbe o zones is closely ela ed o mo phodynamics. These esul s co obo a e he conclusion d awn by Schlache and Thompson (2013), who de ec ed no signi ican co ela ion be ween habi a me ic (habi a dimensions, sedimen p ope ies, and mo phodynamic s a e) and he numbe o auna zones. Al hough he numbe o biological zones a ied among beaches, a common zona ion pa e n was es ab- lished o he en i e se o beaches s udied. This was done in o de o cha ac e ize he mos ypical species inhabi ing each zone. The gene al pa e n showed h ee biological zones acco ding o he zona ion scheme o Dahl (1952). In gene al, he sup ali o al zone was ypi ied by ai -b ea hing amphipods (Tali us sal a o ) and coleop e ans. The middle zone was domina ed by ue in e idal species, such as Haus o iidae amphipods (Haus o ius a ena ius), Ci olanidae isopods (Eu ydice a inis), Spionidae polychae es (Scolelepis spp.) and neme eans. The lowe o subli o al zone was ypi ied by amphipods belonging o he Pon opo eiidae amily: mysids, and spionid polychae es. The dis ibu ion o he species in each zone is in ag eemen wi h indings on o he nea by empe a e sandy beaches, such as hose along he no he n coas s o Spain, Tunisia, and Mo- occo (Rodil e al. 2006, Pé ez-Domingo e al. 2008, Bayed 2003). Di e si y and densi ies o indi iduals inc ease o- wa ds he lowe zones. This is a gene al ea u e ound in nume ous s udies o sandy beaches wo ldwide (Ja amil- lo e al. 1993, Gonçal es e al. 2009, Rodil e al. 2006). Some au ho s ha e de e mined ha his pa e n could be due o a e lec ion o he high sub- idal di e si y and sho pe iods o ai exposu e, hus allowing mo e species o inhabi zones closes o he seawa e (Deg ae e al. 1999, Ae s e al. 2004). The high abundance ound in he lowe a eas o all he beaches s udied e idences how impo an hese en i onmen s a e as po en ial sou ces o ood o o he p eda o y species ( ish and bi ds). Rela ionship be ween en i onmen al a iables and mac oin auna Dis ibu ion o mac oin auna is ela ed o he ole - ance o hese communi ies o en i onmen al a iables (McLachlan and B own 2006). Al hough he ela ion- ship be ween species and he en i onmen could change wi h he scale o s udy (Rodil e al. 2012), abio ic p e- dic o a iables a he local scale we e examined. Beach slope and g ain size ha e been iden i ied as main ac o s con olling mac oin auna dis ibu- ion h oughou he in e idal zone (McLachlan e al. 1993, Ja amillo and MacLachlan 1993). Resul s om CCA analysis showed ha sand mois u e and o ganic ma e con en , in addi ion o ele a ion and g ain size, we e he main en i onmen al a iables con olling he mac oin auna dis ibu ion ac oss he sho e on sandy beaches o he Gul o Cadiz coas . Lowe and middle zones showed an in e nal g adi- en in luenced mainly by a e age g ain size. Species inhabi ing hese zones we e Pon oc a es a ena ius, Siphonoece es saba ie i and neme eans, closely e- la ed o coa se g ain size, while Donax unculus and Ba hypo eia pelagica we e ela ed o ine g ain size. The mos abundan species in uppe zones, he ali- id amphipod Tali us sal a o and coleop e ans, we e co ela ed posi i ely wi h ele a ion bu nega i ely wi h sand mois u e and o ganic ma e con en . G ain size was no a good explana o y a iable o hese species. In ac , Ugolini e al. (2008) ound no ela ionship be ween sand-hoppe abundance and sand-g ain size. Al hough hese species showed signi ican ela ion- ships wi h abio ic a iables, o he ac o s no aken in o accoun could a ec hei dis ibu ion. Fo example, i has been epo ed ha s anded ma e ial (e.g. mac o- phy es and mac oalgae) p o ides a physical s uc u e ha can be used as a shel e o b eeding si e and as a ood sou ce by sup ali o al a h opods (Olaba ia e al. 2007) and he age o hese deposi s plays a signi ican ole in he s uc u e o uppe -sho e assemblages (Ruiz- Delgado e al. 2014). Conse a ion and ec ea ion po en ial indices The conse a ion and ec ea ion po en ial indices de eloped by McLachlan e al. (2013) a e use ul ools o de e mining he mos sui able beach managemen Mac oin auna assemblages on sandy beaches • 9 SCI. MAR., 79(3), Sep embe 2015, 000-000. ISSN-L 0214-8358 doi: h p://dx.doi.o g/10.3989/scima .04133.30A app oach, p o iding a se o managemen p inciples ha can easily be ollowed, e en wi h limi ed e- sou ces. In hese cases, mapping plays a key ole o he collec ion o in o ma ion and has been iden i ied as a p incipal s ep owa ds managemen and conse - a ion policies (Role e al. 2015). In addi ion, he implemen a ion o hese indices p o ides he b oades cha ac e iza ion o sandy beaches, no only based on physical ea u es, since sandy beaches a e also social and ecological en i onmen s. In his wo k, 12 case s udies illus a e he applica ion o he CI and RI indices. Al hough hese indices include i e ca ego ies, ou beaches a e g ouped mainly in o wo: in ensi e ec ea ion and mul iple use. Mos beaches ha e shown sco es abo e 5 on he CI, so hey ha e a ea- sonable po en ial o conse a ion. Fo example, Te ón, Mazagón and Le an e beaches, ca ego ized by hese indices as o mul iple use, ha e a high conse a ion po- en ial. Coas al managemen policies should he e o e be designed o p omo e measu es o he conse a ion o hese en i onmen s. On he o he hand, only Ma alas- cañas and Valdelag ana, ha e a high ec ea ion po en- ial, which can be exploi ed wi h app op ia e manage- men s a egies o he possible e-e alua ion o exis ing measu es ha educe he CI alue, o example. In conclusion, he beaches on he Spanish coas - line along he Gul o Cadiz a e cha ac e ized by high biodi e si y, including majo bio-indica o species, and by a clea zona ion o mac oin auna. The o e all dis ibu ion pa e n in ol es h ee biological zones: he sup ali o al zone, cha ac e ized by ai -b ea hing amphipods and coleop e ans; he middle zone, cha - ac e ized by Haus o iidae amphipods, Ci olanidae iso- pods, Spionidae polychae es and neme eans; and he subli o al zone, cha ac e ized by amphipods belong- ing o he Pon opo eiidae amily, mysids and spionid polychae es. The ac oss-sho e dis ibu ion o he mac- oin auna is in luenced p ima ily by sand mois u e, o ganic ma e con en , ele a ion and g ain size. O he ac o s such as w ack deposi and o ganic inpu s om i e s and es ua ies could in luence he abundance and dis ibu ion o mac oin auna inhabi ing sandy beach- es. The e o e, u u e s udies a e needed o elucida e whe he he p esence o s anded ma e ial could a ec he global zona ion pa e ns on sandy beaches. The ec ea ion and conse a ion indices showed ha he s udied beaches ha e mul iple unc ionali ies, gi en hei social, physical, mo phodynamic and bio ic ea- u es, al hough many o hem ha e a high po en ial o conse a ion ha needs o be managed. ACKNOWLEDGEMENTS We would like o hank he “Los To uños” Na u al Pa k (Cadiz) and i s s a o he acili ies p o ided du - ing sampling. This wo k was suppo ed by he incen i e p og amme o excellen esea ch p ojec s inanced by he go e nmen o Andalucía (P09- HUM-4717). This p ojec also suppo ed a PhD G an awa ded o he i s au ho . M.C. Ruiz-Delgado was suppo ed by he Spanish Minis y o Educa ion ia a p e-doc o al g an (FPU) (AP-2009-3906). REFERENCES Ae s K., Vanag T., Deg ae S., e al. 2004. Mac oin aunal com- muni y s uc u e and zona ion o an Ecuado ian sandy beach (Bay o Valdi ia). Belg. Zool. 134: 15-22. 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