Changes in seagrass polychaete assemblages after invasion by Caulerpa racemosa var. cylindracea (Chlorophyta: Caulerpales): community structure, trophic guilds and taxonomic distinctness


  • Antonio Box Laboratorio de Biología Marina, Universidad de las Islas Baleares (UIB)
  • Daniel Martin Centre d’Estudis Avançats de Blanes (CEAB), Consejo Superior de Investigaciones Científicas (CSIC)
  • Salud Deudero Laboratorio de Biología Marina, Universidad de las Islas Baleares (UIB)



Posidonia oceanica, Caulerpa racemosa, Polychaeta, species diversity, trophic guilds, taxonomic distinctness, introduced species, Mediterranean


A two-year study focusing on the associated polychaete assemblages revealed that the degradation of the Posidonia oceanica seagrass meadows, together with the colonisation of rhizomes by invasive Caulerpa racemosa in the Balearic Islands, have produced important changes in the ecosystem functioning, and have therefore affected the benthic faunal communities. The highest abundance and number of species occurred in C. racemosa from August to December. The species composition of the polychaete assemblage in C. racemosa is similar to that of P. oceanica but abundances of shared species differ between Caulerpa and Posidonia. Abundance, number of species and diversity were positively correlated with C. racemosa biomass. The spatial complexity provided by the network of C. racemosa fronds and stolons seemed to combine with that of the remaining seagrass mat to support a well developed polychaete assemblage. Although invaded meadows harboured significantly modified polychaete assemblages when compared with living P. oceanica meadows, the main impacts seem to be non-harmful (i.e. increased abundance and number of species) and mainly affect their seasonal patterns, which is a direct consequence of the corresponding biomass seasonality of C. racemosa.


Download data is not yet available.


Amade, P. and R. Lemee. – 1998. Chemical defence of the Mediterranean alga Caulerpa taxifolia: variations in caulerpenyne production. Aquat. Toxicol. 43: 287-300. doi:10.1016/S0166-445X(98)00054-X

Antoniadou, C. and C. Chintiroglou. – 2006. Trophic relationships of polychaetes associated with different algal growth forms. Helgoland Mar. Res., 60: 39-49. doi:10.1007/s10152-005-0015-2

Antoniadou, C. and C. Chintiroglou. – 2007. Zoobenthos associated with the invasive red alga Womersleyella setacea (Rhodomelacea) in the northern Aegean Sea. J. Mar. Biol. Assoc. the U. K., 87: 629-641. doi:10.1017/S0025315407048151

Argyrou, M., A. Demetropoulos and M. Hadjichristophorou. – 1999. Expansion of the macroalga Caulerpa racemosa and changes in softbottom macrofaunal assemblages in Moni Bay, Cyprus. Oceanol. Acta, 22: 517-528. doi:10.1016/S0399-1784(00)87684-7

Balata, D., L. Piazzi and F. Cinelli. – 2004. A comparison among assemblages in areas invaded by Caulerpa taxifolia and C. racemosa on a subtidal Mediterranean rocky bottom. Mar. Ecol. P.S.Z.N.I, 25: 1-13. doi:10.1111/j.1439-0485.2004.00013.x

Ballesteros, M., J. Castelló, M. Gallés and R. Sarda. – 1987. Invertebrados alguícolas marinos de las Islas Pitiusas. Consell Insular d’Eivissa i Formentera, Conselleria d’Ecologia i Medi Ambient. Ciutat d’Eivissa.

Bax, N., A. Williamson, M. Aguero, E. Gonzalez and W. Geeves. – 2003. Marine invasive alien species: a threat to global biodiversity. Mar. Policy, 27: 313-323. doi:10.1016/S0308-597X(03)00041-1

Bellan-Santini, D., P.M. Arnaud, G. Bellan and M. Verlaque. – 1996. The influence of the introduced tropical alga Caulerpa taxifolia, on the biodiversity of the Mediterranean marine biota. J. Mar. Biol. Assoc. the U. K., 76: 235-237. doi:10.1017/S0025315400029180

Borg, J.A., A.A. Rowden, M.J. Attrill, P.J. Schembri and M.B. Jones. – 2006. Wanted dead or alive: high diversity of macroinvertebrates associated with living and ‘dead’ Posidonia oceanica matte. Mar. Biol., 149: 667-677. doi:10.1007/s00227-006-0250-3

Boudouresque, C.F., R. Lemee, X. Mari and A. Meinesz. – 1996. The invasive alga Caulerpa taxifolia is not a suitable diet for the sea urchin Paracentrotus lividus. Aquat. Bot., 53: 245-250. doi:10.1016/0304-3770(96)01021-2

Boudouresque, C.F. and M. Verlaque. – 2002. Biological pollution in the Mediterranean Sea: invasive versus introduced macrophytes. Mar. Pollut. Bull., 44: 32-38. doi:10.1016/S0025-326X(01)00150-3

Box, A. – 2008. Ecología de caulerpales: fauna y biomarcadores. PhD. thesis. Balearic Islands Univ., Spain.

Brito, M.C., D. Martin and J. Nunez. – 2005. Polychaetes associated to a Cymodocea nodosa meadow in the Canary Islands: assemblage structure, temporal variability and vertical distribution compared to other Mediterranean seagrass meadows. Mar. Biol., 146: 467-481. doi:10.1007/s00227-004-1460-1

Buia, M.C., M.C. Gambi, A. Terlizzi and L. Mazzella. – 2001. Colonization of Caulerpa racemosa along the southern italian coast: I.Distribution, phenological variability and ecological role. In: V. Gravez, S. Ruitton, C.F. Boudouresque, L. Le Direac’h, A. Meinesz and S. G. (eds.). Fourth International Workshop on Caulerpa taxifolia, pp 352-360. GIS Posidonie, France.

Buschbaum, C., A.S. Chapman and B. Saier. – 2006. How an introduced seaweed can affect epibiota diversity in different coastal systems. Mar. Biol., 148: 743-754. doi:10.1007/s00227-005-0128-9

Ceccherelli, G. and N. Sechi. – 2002. Nutrient availability in the sediment and the reciprocal effects between the native seagrass Cymodocea nodosa and the introduced rhizophytic alga Caulerpa taxifolia. Hydrobiologia, 474: 57-66. doi:10.1023/A:1016514621586

Ceccherelli, G., L. Piazzi and F. Cinelli. – 2000. Response of the non-indigenous Caulerpa racemosa (Forsskal) J-Agardh to the native seagrass Posidonia oceanica (L.) Delile: effect of density of shoots and orientation of edges of meadows. J. Exp. Mar. Biol. Ecol., 243: 227-240. doi:10.1016/S0022-0981(99)00122-7

Ceccherelli, G., L. Piazzi and D. Balata. – 2002. Spread of introduced Caulerpa species in macroalgal habitats. J. Exp. Mar. Biol. Ecol., 280: 1-11. doi:10.1016/S0022-0981(02)00336-2

Clarke, K.R. and R.M. Warwick. – 2001. A further biodiversity index applicable to species lists: variation in taxonomic distinctness. Mar. Ecol.-Prog. Ser., 216: 265-278. doi:10.3354/meps216265

Crooks, J.A. – 2002. Characterizing ecosystem-level consequences of biological invasions: the role of ecosystem engineers. Oikos, 97: 153-166. doi:10.1034/j.1600-0706.2002.970201.x

Der Hartog, C. – 1994. Suddocation of a littoral Zostera bed by Enteromorpha radiata. Aquat. Bot., 47: 21-28. doi:10.1016/0304-3770(94)90045-0

Fauchald, K. and P. Jumars. – 1979. The diet of worms: a study of Polychaeta feeding guilds. Oceanogr. Mar. Biol., 17: 193-284.

Galil, B.S. and A. Zenetos. – 2002. A sea change -exotics in the Eastern Mediterranean Sea. In: E. Leppäkoski, S. Gollasch and S. Olenin (eds.), Invasive Aquatic Species in Europe, Distribution, Impacts and Management, pp 325-340. Kluwer Academic Publishers, Dordrecht, Boston, London.

Gambi, M.C., A. Giangrande, M. Martinelli and L.A. Chessa. – 1995. Polychaetes of a Posidonia oceanica bed off Sardinia (Italy) - Spatiotemporal distribution and feeding guild analysis. Sci. Mar., 59: 129-141.

Gambi, M.C., G. Conti and C.S. Bremec. - 1998. Polychaete distribution, diversity and seasonality related to seagrass cover in shallow soft bottoms of the Tyrrhenian Sea (Italy). Sci. Mar., 62: 1-17.

Gambi, M.C., B.I. Van Tussenbroek and A. Brearley. – 2003. Mesofaunal borers in seagrasses: world-wide occurrence and a new record of boring polychaetes in the Mexican Caribbean. Aquat. Bot., 76: 65-77. doi:10.1016/S0304-3770(03)00031-7

Gambi, M.C., B. Trani and M.C. Buia. – 2005. Taxonomic diversity and distribution of polychaete and isopod borers on the sheaths of the seagrass Posidonia oceanica: analysis at regional scale along the coast off Sardinia (Italy). Ital. J. Zool., 72: 141-151. doi:10.1080/11250000509356664

Givernaud, T.J., J. Cosson and A. Givernaud-Mouradi. – 1991. Etudes des populations de Sargassum muticum sur les cores de Basse, Normandies (France). In: M. Elliott and J.P. Ducrotoy (eds.).19th Symposium Estuarines and Coasts: Spatial and Temporal Intercomparison, pp 129-132. International Symposium Series.

Gollasch, S. - 2006. Overview on introduced aquatic species in European navigational and adjacent waters. Hegoland Mar. Res., 60: 84-89. doi:10.1007/s10152-006-0022-y

Jung, V., T. Thibaut, A. Meinesz and G. Pohnert. – 2002. Comparison of the wound-activated transformation of caulerpenyne by invasive and noninvasive Caulerpa species of the Mediterranean. J. Chem. Ecol., 28: 2091-2105. doi:10.1023/A:1020710230532 PMid:12474902

Klein, J. and M. Verlaque. -2008. The Caulerpa racemosa invasion: A critical review. Mar. Pollut. Bull., 56: 205-225. doi:10.1016/j.marpolbul.2007.09.043 PMid:18061625

Levi, F. and P. Francour. – 2004. Behavioural response of Mullus surmuletus to habitat modification by the invasive macroalga Caulerpa taxifolia. J. Fish Biol., 64: 55-64. doi:10.1111/j.1095-8649.2004.00280.x

Longepierre, S., A. Robert, F. Levi and P. Francour. – 2005. How an invasive alga species (Caulerpa taxifolia) induces changes in foraging strategies of the benthivorous fish Mullus surmuletus in coastal Mediterranean ecosystems. Biodivers. Conserv., 14: 365-376. doi:10.1007/s10531-004-5356-7

Mack, R.N., D. Simberloff, W.M. Lonsdale, H. Evans, M. Clout and F.A. Bazzaz. – 2000. Biotic invasions: causes, epidemiology, global consequences, and control. Ecol. Applications, 10: 689-710. doi:10.1890/1051-0761(2000)010[0689:BICEGC]2.0.CO;2

Martin, D., S. Pinedo and R. Sarda. – 2000. Distribution patterns and trophic structure of soft-bottom Polychaeta assemblages in north-western Mediterranean shallow-water bay. Ophelia, 53: 1-17.

Meinesz, A. – 2004. Caulerpa taxifolia continues its invasion. Biofutur, 244: 41-46.

Meinesz, A., T. Belsher, T. Thibaut, B. Antolic, K.B. Mustapha, C.F. Boudouresque, D. Chiaverini, F. Cinelli, J.M. Cottalorda, A.S. Djellouli, A. El Abed, C. Orestano, A.M. Grau, L. Ivesa, I. Vaugelas, N. Zavodnik and A. Zuljevic. – 2001. The introduced green alga Caulerpa taxifolia continues to spread in Mediterranean. Biol. Inv., 3: 201-210. doi:10.1023/A:1014549500678

Molenaar, H., A. Meinesz, and T. Thiebaut. – 2009. Alterations of the structure of Posidonia oceanica beds due to the introduced alga Caulerpa taxifolia. Sci. Mar., 73(2): 329-335. doi:10.3989/scimar.2009.73n2329

Montefalcone, M., C. Morri, A. Peirano, G. Albertelli and C.N. Bianchi.– 2007. Substitution and phase shift within the Posidonia oceanica seagrass meadows of NW Mediterranean Sea. Estuar.Coast. Shelf Sci., 75: 63-71. doi:10.1016/j.ecss.2007.03.034

Pandolfo, A. and R. Chemello. – 1995. Prime nota sulla malacofauna associata a Caulerpa racemosa nella Baia di Santa Panagia (Sicilia orientale). Biol. Mar. Mediterr., 2: 649-651.

Parent-Massin, D., V. Fournier, P. Amade, R. Lemee, M. DurandClement, C. Delescluse and D. Pesando. – 1996. Evaluation of the toxicological risk to humans of caulerpenyne using human hematopoietic progenitors, melanocytes, and keratinocytes in culture. J. Toxicol. Environ. Health, 47: 47-59. doi:10.1080/009841096161924

Pesando, D., P. Huitorel, V. Dolcini, P. Amade and J.P. Girard. – 1998. Caulerpenyne interferes with microtubule-dependent events during the first mitotic cycle of sea urchin eggs. Eur. J. Cell Biol., 77: 19-26. PMid:9808285

Piazzi, L. and F. Cinelli. – 1999. Development and seasonal dynamics of a population of the tropical alga Caulerpa racemosa (Forsskal) J. Agardh in the Mediterranean. Cryptogamie Algol., 20: 295-300. doi:10.1016/S0181-1568(00)88146-5

Piazzi, L. and F. Cinelli. – 2000. Effects of the spread of the introduced Rhodophyceae Acrothamnion preissii and Womersleyella setacea on the macroalgal community of Posidonia oceanica rhizomes in the western Mediterranean sea. Cryptogamie Algol., 21: 291-300. doi:10.1016/S0181-1568(00)00116-1

Raniello, R., E. Mollo, M. Lorenti, M. Gavagnin and M.C. Buia. – 2007. Phytotoxic activity of caulerpenyne from the Mediterranean invasive variety of Caulerpa racemosa: a potential allelochemical. Biol. Inv., 9: 361-368. doi:10.1007/s10530-006-9044-2

Rosenzweig, M.L. – 2001. The four questions: What does the introduction of exotic species do to diversity? Evol. Ecol. Res., 3: 361-367.

Sakai, A.K., F.W. Allendorf, J.S. Holt, D.M. Lodge, J. Molofsky, K.A. With, S. Baugman, R.J. Cabin, J.E. Cohen, N.C. Ellstrand, D.E. McCauley, O. O’Neil, I.M. Parker, J.N. Thompson and S.G. Weller. – 2001. The population biology of invasive species. Ann. Rev. Ecol. Syst., 32: 305-332. doi:10.1146/annurev.ecolsys.32.081501.114037

Sanchez-Moyano, J.E., F.J. Estacio, E.M. Garcia-Adiego and J.C. Garcia-Gomez. – 2001. Effect of the vegetative cycle of Caulerpa prolifera on the spatio-temporal variation of invertebrate macrofauna. Aquat. Bot., 70: 163-174. doi:10.1016/S0304-3770(00)00144-3

Sax, D.F. and S.D. Gaines. – 2003. Species diversity: from global decreases to local increases. Trends Ecol. Evol., 18: 561-566. doi:10.1016/S0169-5347(03)00224-6

Smith, S.D. – 2008. Interpreting molluscan death assemblages on rocky shores: Are they representative of the regional fauna? J. Exp. Mar. Biol. Ecol., 366: 151-159. doi:10.1016/j.jembe.2008.07.019

Streftaris, N. and A. Zenetos. – 2006. Alien marine species in the Mediterranean - the 100 ‘Worst Invasives’ and their impact. Medit. Mar. Sci., 7(1): 87-118.

Sureda, A., A. Box, M. Ensenat, E. Alou, P. Tauler, S. Deudero and A. Pons. – 2006. Enzymatic antioxidant response of a labrid fish (Coris julis) liver to environmental caulerpenyne. Comp. Biochem. Physiol. C, 144: 191-196.

Sureda, A., A. Box, S. Deudero and A. Pons. – 2008. Reciprocal effects of caulerpenyne and intense herbivorism on the antioxidant response of Bittium reticulatum and Caulerpa taxifolia. Ecotoxicol. Environ. Saf.

Vazquez-Luis, M., P. Sanchez-Jerez and J.T. Bayle-Sempere. – 2008. Changes in amphipod (Crustacea) assemblages associated with shallow-water algal habitats invaded by Caulerpa racemosa var. cylindracea in the western Mediterranean Sea. Mar. Environ. Res., 65: 416-426. doi:10.1016/j.marenvres.2008.01.006 PMid:18342365

Viejo, R.M. – 1999. Mobile epifauna inhabiting the invasive Sargassum muticum and two local seaweeds in northern Spain. Aquat. Bot., 64: 131-149. doi:10.1016/S0304-3770(99)00011-X

Warwick, R.M. and J. Light. – 2002. Death assemblages of molluscs on St Martin’s Flats, Isles of Scilly: a surrogate for regional biodiversity ? Biodivers. Conserv. 11: 99-112. doi:10.1023/A:1014094829984

Wentworth, W. - 1972. A scale of grade and class terms for clastic sediments. J. Geol., 30: 377–392. doi:10.1086/622910

Zenetos, A., S. Gofas, G.F. Russo and J. Templado. – 2003. CIESM Atlas of Exotic Species in the Mediterranean, Volume 3: Mollusc. CIESM Publishers, Monaco.

Zuljevic, A., B. Antolic and V. Onofri. – 2003. First record of Caulerpa racemosa (Caulerpales : Chlorophyta) in the Adriatic Sea. J. Mar. Biol. Assoc. U.K., 83: 711-712. doi:10.1017/S0025315403007689h




How to Cite

Box A, Martin D, Deudero S. Changes in seagrass polychaete assemblages after invasion by Caulerpa racemosa var. cylindracea (Chlorophyta: Caulerpales): community structure, trophic guilds and taxonomic distinctness. Sci. mar. [Internet]. 2010Jun.30 [cited 2024Mar.1];74(2):317-29. Available from:




Most read articles by the same author(s)

1 2 > >>