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Evects of Ocean Acidiwcation on Invertebrate Settlement at Volcanic CO Vents

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Evects of Ocean Acidiwcation on Invertebrate Settlement at Volcanic CO Vents View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Plymouth Electronic Archive and Research Library Mar Biol (2010) 157:2489–2502 DOI 10.1007/s00227-010-1513-6 ORIGINAL PAPER EVects of ocean acidiWcation on invertebrate settlement at volcanic CO2 vents M. Cigliano · M. C. Gambi · R. Rodolfo-Metalpa · F. P. Patti · J. M. Hall-Spencer Received: 8 April 2010 / Accepted: 30 June 2010 / Published online: 16 July 2010 © Springer-Verlag 2010 W V V Abstract We present the rst study of the e ects of ocean levels of CO2 can profoundly a ect the settlement of a wide acidiWcation on settlement of benthic invertebrates and range of benthic organisms. microfauna. ArtiWcial collectors were placed for 1 month V along pH gradients at CO2 vents o Ischia (Tyrrhenian Sea, Italy). Seventy-nine taxa were identiWed from six main Introduction taxonomic groups (foraminiferans, nematodes, polychaetes, molluscs, crustaceans and chaetognaths). Calcareous forami- Increasing atmospheric CO2 concentrations are causing a niferans, serpulid polychaetes, gastropods and bivalves rise in pCO2 concentrations at the ocean surface (Houghton showed highly signiWcant reductions in recruitment to the et al. 1992; Keeling and Whorf 1994) due to atmospheric collectors as pCO2 rose from normal (336–341 ppm, pH CO2 ocean exchange on time scales of several months 8.09–8.15) to high levels (886–5,148 ppm) causing acidi- (Zeebe and Wolf-Gladrow 2001). By 2100, the concentra- W ed conditions near the vents (pH 7.08–7.79). Only the syl- tion of CO2 in the ocean is expected to rise to 750 ppm, lid polychaete Syllis prolifera had higher abundances at the which is about twice the present 385–390 ppm (Feely et al. W most acidi ed station, although a wide range of polychaetes 2004; Raven et al. 2005). As CO2 dissolves in the surface and small crustaceans was able to settle and survive under ocean, it reacts with water to form carbonic acid (H2CO3), ¡ these conditions. A few taxa (Amphiglena mediterranea, which dissociates to bicarbonate (HCO3 ), carbonate ions 2¡ + Leptochelia dubia, Caprella acanthifera) were particularly (CO3 ) and protons (H ). With increasing atmospheric W abundant at stations acidi ed by intermediate amounts of pCO2, the equilibrium of the carbonate system will shift to ¡ 2¡ CO2 (pH 7.41–7.99). These results show that increased higher CO2 and HCO3 levels, while CO3 concentration and pH will decrease. These changes in carbonate chemistry, often referred to Communicated by F. Bulleri. as ‘ocean acidiWcation’, are already occurring and are expected to intensify in the future. Models predict that the M. Cigliano and M. C. Gambi contributed equally. pH of surface seawater will drop by 0.4 units by the year 2100 (Caldeira and Wickett 2003). Consequently, the rise M. Cigliano · M. C. Gambi (&) · F. P. Patti Stazione Zoologica Anton Dohrn, of CO2 in ocean waters leads to more corrosive conditions Laboratory of Functional and Evolutionary Ecology, for calcifying organisms, making it more diYcult for them Villa Comunale, 80121 Naples, Italy to build and maintain their carbonate skeletons (Raven e-mail: [email protected] et al. 2005). CalciWcation rates of several species, including R. Rodolfo-Metalpa · J. M. Hall-Spencer coralline algae, coccolithophores, corals, bivalves and echi- Marine Institute, Marine Biology and Ecology Research Centre, noderms, decreases with increasing pCO2 (e.g. Kleypas University of Plymouth, Plymouth, UK et al. 2006; Fabry et al. 2008), although the response is spe- cies speciWc with the up-regulation of calciWcation in some R. Rodolfo-Metalpa IAEA, Marine Environment Laboratories, species (Wood et al. 2008; Ries et al. 2009; Jury et al. Monaco, MC, Monaco 2009; Rodolfo-Metalpa et al. 2010a). The recruitment rate 123 2490 Mar Biol (2010) 157:2489–2502 and the growth of crustose coralline algae is severely inhib- 1970), where leaves Xoat on the water surface at low tide ited under elevated pCO2, suggesting that changes in ben- with the highest mean shoot density recorded around Ischia thic community structure may occur owing to the impact (up to 900 shoots/m2, Buia et al. 2003). Previous studies at of ocean acidiWcation on recruitment and competition the Castello report rich algal (Bourdouresque and Cinelli for space (KuVner et al. 2007; Hall-Spencer et al. 2008; 1971, 1976) and sponge communities (Sarà 1959; Pulitzer Jokiel et al. 2008). Initial results by Porzio et al. (2008) Finali 1970; Pulitzer Finali and Pronzato 1976), as well as demonstrate signiWcant loss of algal diversity and changes diverse invertebrate and Wsh faunas associated with the in macroalgal community structure in a naturally acidiWed Posidonia oceanica meadows (Russo et al. 1984a; Guidetti environment. and Bussotti 1998; Scipione 1999; Gambi and CaWero However, eVects of ocean acidiWcation on larval pelagic 2001) in the areas nearby and partially inXuenced by the stages of invertebrates are still poorly understood (Vézina vents. Hall-Spencer et al. (2008) reported a total of 64 and Hoegh-Guldberg 2008). Many laboratory studies have megabenthic taxa along the gradient at the vents areas, shown that the early life history stages of several organisms where reductions in the diversity of adult populations are are negatively impacted by acidiWed seawater, including caused partly by the dissolution of calciWed species due to work on echinoderms, crustaceans and molluscs (Kurihara lowered pH (Martin et al. 2008; Rodolfo-Metalpa et al. and Shirayama 2004; Kurihara et al. 2004, 2007; Dupont 2010b). Loss of macroalgal diversity was also shown by et al. 2008; Kurihara and Ishimatsu 2008; Ellis et al. 2009; Porzio et al. (2008). Although CO2 vents are localised and Findlay et al. 2009). However, potential shifts in benthic highly variable in pH, they provide information about the V V recruitment that may result from these e ects on early life ecological e ects of long-term exposures to high CO2 lev- history changes are unknown due to the diYculties of main- els encompassing the life cycles of interacting macroben- taining mixed populations of delicate larval stages in labo- thic organisms as well as the feedbacks and indirect eVects ratory conditions. that occur within natural marine systems (Hall-Spencer Hall-Spencer et al. (2008) have shown that natural CO2 et al. 2008; Riebesell 2008). The aim of this study was to venting sites may be useful for assessing the long-term use artiWcial collectors to determine whether invertebrates eVects of ocean acidiWcation on benthic biota and sea-Xoor and microfauna varied along gradients in pH where acidiW- V ecosystems. They indicate that, although natural CO2 vent- cation of water by CO2 a ects natural marine communities. ing sites are not precise analogues of global-scale ocean We focus on early-settled stages of invertebrates and acidiWcation, they can provide essential information about foraminiferans, a fauna component which was not consid- V high-CO2 e ects on spatial and temporal scales which are ered at the vents initial survey by Hall-Spencer et al. otherwise diYcult to address. Here, we provide Wrst data on (2008). Our null hypothesis was that there would be no sig- the eVects of acidiWcation on invertebrates and microfauna niWcant diVerences in species composition and community settled on artiWcial collectors placed at various distances structure in the collectors placed along the pH gradients V from CO2 vents, creating a gradient of di erent pH condi- studied. tions. Materials and methods Study site ArtiWcial collectors (scouring pads) were placed in situ Castello Aragonese, located at the north-eastern side of along 300 m transects on the north and south sides of Cas- Ischia island (Fig. 1, 40° 43.84Ј N, 13° 57.08Ј E) is part of a tello Aragonese at six stations (N1, N2, N3, S1, S2, S3) 132,000 years old volcano (Rittmann and Gottini 1981) where Hall-Spencer et al. (2008) had recorded signiWcant V where gas vents occur in shallow water (Tedesco 1996). di erences in pH due to CO2 vents (Fig. 1). N1 and S1 are The gas comprises 90–95% CO2, 3–6% N2, 0.6–0.8% O2, monitoring stations located under normal pH conditions, 0.2–0.8% CH4 and 0.08–0.1% air and bubbles at about N2 and S2 are intermediate stations, characterized by high 1.4 £ 106 l d¡1 at ambient temperature. The site is micro- pH Xuctuations and mean intermediate values, while N3 W tidal (0.30–0.50 m range) and the CO2 vents lack sulphur and S3 are characterized by low pH, acidi ed conditions. (Tedesco 1996); they acidify normal salinity and alkalinity Water samples (n = 3–5) were taken at buoyed stations by of seawater along a pH gradient from 8.17 down to 6.57 for SCUBA divers using glass bottles (250 cc volume) during 300 m running parallel to the rocky shore on the north and the month of collector deployment, and the pHT (total south sides of the Castello (Hall-Spencer et al. 2008). The scale) was measured immediately using a meter accurate to south side is more sheltered from wave action and has a 0.01 pH units (Metrohm 826 pH mobile) calibrated using shallow (0.5 m depth) Posidonia oceanica meadow form- TRIS/HCl and 2-aminopyridine/HCl buVer solutions (DOE ing a reef-like structure (sensu Augier and Boudouresque 1994). Seawater samples were then passed through 0.45-m 123 Mar Biol (2010) 157:2489–2502 2491 Class and Order) and counted, the foraminiferans, poly- chaetes, molluscs, isopods and amphipods were classiWed to genus and, where possible, to species. Data analyses Number of species, abundance, Shannon diversity (H’) and Pielou’s evenness (J) were calculated for each collector and plotted as means § SD (n = 3 at each station).
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