Scientia Marina, Vol 76, No 4 (2012)

Benthic assemblages of rock pools in northern Portugal: seasonal and between-pool variability

Iacopo Bertocci
CIIMAR/CIMAR, Centro Interdisciplinar de Investigação Marinha e Ambiental , Portugal

Rita Araújo
CIIMAR/CIMAR, Centro Interdisciplinar de Investigação Marinha e Ambiental , Portugal

Mónica Incera
Centro Tecnológico del Mar – Fundación CETMAR , Spain

Francisco Arenas
CIIMAR/CIMAR, Centro Interdisciplinar de Investigação Marinha e Ambiental , Portugal

Rui Pereira
CIIMAR/CIMAR, Centro Interdisciplinar de Investigação Marinha e Ambiental , Portugal

Helena Abreu
CIIMAR/CIMAR, Centro Interdisciplinar de Investigação Marinha e Ambiental - Daithi O’Murchu Marine Research Station , Portugal

Kim Larsen
CIIMAR/CIMAR, Centro Interdisciplinar de Investigação Marinha e Ambiental , Portugal

Isabel Sousa-Pinto
CIIMAR/CIMAR, Centro Interdisciplinar de Investigação Marinha e Ambiental - Department of Biology, Faculty of Sciences, University of Porto , Portugal


We investigated the seasonal (winter vs summer and within season) and spatial (between-pool) variability of benthic assemblages of rock pools at mid-intertidal level along the shore of Viana do Castelo (North Portugal). Physical traits of rock pools, including size, depth and position along the shore, were also compared between pools. While pools did not differ for any of the examined physical traits, results indicated a clear seasonal difference in the structure of assemblages, including a total of 49 macroalgal and 13 animal taxa. This finding was driven by six taxa that are more abundant in winter (the reef-forming polychaete Sabellaria alveolata, the articulated coralline algae Corallina spp., the brown alga Bifurcaria bifurcata, the encrusting coralline alga Lithophyllum incrustans, the red alga Chondracanthus acicularis and the grazing snails Gibbula spp.) and four algal taxa that are more abundant in summer (the invasive brown Sargassum muticum, the green Ulva spp., the kelp Laminaria ochroleuca and the filamentous red Ceramium spp.). These data provide a new contribution to the knowledge of rock pool systems and have potential implications for monitoring programmes aimed at assessing ecological modifications related to natural and anthropogenic disturbances and for identifying processes responsible for the variability of rock pool assemblages.


rock pools; rocky intertidal; seasonal variability; benthos; Sabellaria alveolata; Portugal

Full Text:



Airoldi L. 2000. Responses of algae with different life histories to temporal and spatial variability of disturbance in subtidal reefs. Mar. Ecol. Prog. Ser. 195: 81-92.

Anderson M.J. 2001. A new method for non-parametric multivariate analysis of variance. Austral. Ecol. 26: 32-46.

Anderson M.J. 2006. Distance-based tests for homogeneity of multivariate dispersions. Biometrics 62: 245-253. PMid :16542252

Anderson M.J., Robinson J. 2003. Generalized discriminant analysis based on distances. Aust. N.Z. J. Stat. 45: 301-318.

Andrew N.L., Mapstone B.D. 1987. Sampling and the description of spatial pattern in marine ecology. Oceanogr. Mar. Biol. Annu. Rev. 52: 39-90.

Araújo R., Sousa-Pinto I., Bárbara I., Quintino V. 2006. Macroalgal communities of intertidal rock pools in the northwest coast of Portugal. Acta Oecol. 30: 192-202.

Araújo R., Vaselli S., Almeida M., Serrão E., Sousa-Pinto I. 2009. Effects of disturbance on marginal populations: human trampling on Ascophyllum nodosum assemblages at its southern distribution limit. Mar. Ecol. Prog. Ser. 378: 81-92.

Arenas F., Fernández C. 2000. Size structure and dynamics in a population of Sargassum muticum (Phaeophyceae). J. Phycol. 36: 1012-1020.

Arenas F., Fernández C., Rico J.M., Fernández E., Haya D. 1995. Growth and reproductive strategies of Sargassum muticum Yendo (Fensholt) and Cystoseira nodicaulis (Whit.) Roberts. Sci. Mar. 59: 1-8.

Arenas F., Viejo R.M., Fernández C. 2002. Density-dependent regulation in an invasive seaweed: responses at plant and modular levels. J. Ecol. 90: 820-829.

Astles K.L. 1993. Patterns of abundance and distribution of species in intertidal rock pools. J. Mar. Biol. Assoc. U.K. 73: 555-569.

Ayata S.D., Ellien C., Dumas F., Dubois S., Thiébaut E. 2009. Modelling larval dispersal and settlement of the reef-building polychaete Sabellaria alveolata: role of hydroclimatic processes on the sustainability of biogenic reefs. Contin. Shelf Res. 29: 1605-1623.

Baer J., Stengel D.B. 2010. Variability in growth, development and reproduction of the non-native seaweed Sargassum muticum (Phaeophyceae) on the Irish west coast. Estuar. Coast. Shelf Sci. 90: 185-194.

Balestri E., Benedetti-Cecchi L., Lardicci C. 2004. Variability in patterns of growth and morphology of Posidonia oceanica exposed to urban and industrial wastes: contrasts with two reference locations. J. Exp. Mar. Biol. Ecol. 308: 1-21.

Benedetti-Cecchi L. 2000. Priority effects, taxonomic resolution, and the prediction of variable patterns of colonisation of algae in littoral rock pools. Oecologia 123: 265-274.

Benedetti-Cecchi L. 2001. Variability in abundance of algae and invertebrates at different spatial scales on rocky sea shores. Mar. Ecol. Prog. Ser. 215: 79-92.

Benedetti-Cecchi L., Cinelli F. 1992. Effects of canopy cover, herbivores and substratum type on patterns of Cystoseira spp. settlement and recruitment in littoral rockpools. Mar. Ecol. Prog. Ser. 90: 183-191.

Benedetti-Cecchi L., Cinelli F. 1994. Recovery of patches in an assemblage of geniculate coralline algae: variability at different successional stages. Mar. Ecol. Prog. Ser. 110: 9-18.

Benedetti-Cecchi L., Cinelli F. 1995. Habitat heterogeneity, sea urchin grazing and the distribution of algae in littoral rock pools on the west coast of Italy (western Mediterranean). Mar. Ecol. Prog. Ser. 126: 203-212.

Benedetti-Cecchi L., Cinelli F. 1996. Patterns of disturbance and recovery in littoral rock pools: nonhierarchical competition and spatial variability in secondary succession. Mar. Ecol. Prog. Ser. 135: 145-161.

Benedetti-Cecchi L., Nuti S., Cinelli F. 1996. Analysis of spatial and temporal variability in interactions among algae, limpets and mussels in low-shore habitats on the west coast of Italy. Mar. Ecol. Prog. Ser. 144: 87-96.

Benedetti-Cecchi L., Bertocci I., Micheli F., Maggi E., Fosella T., Vaselli S. 2003. Implications of spatial heterogeneity for management of marine protected areas (MPAs): examples from assemblages of rocky coasts in the northwest Mediterranean. Mar. Env. Res. 55: 429-458.

Bertocci I., Maggi E., Vaselli S., Benedetti-Cecchi L. 2005. Contrasting effects of mean intensity and temporal variation of disturbance on assemblages of rocky shores. Ecology 86: 2061-2067.

Breitburg D.L. 1984. Residual effects of grazing: inhibition of competitor recruitment by encrusting coralline algae. Ecology 65: 1136-1143.

Britton-Simmons K.H. 2004. Direct and indirect effects of the introduced alga Sargassum muticum on benthic subtidal communities of Washington State, USA. Mar. Ecol. Prog. Ser. 277: 61-78.

Bulleri F., Bertocci I., Micheli F. 2002. Interplay of encrusting coralline algae and sea urchins in maintaining alternative habitats. Mar. Ecol. Prog. Ser. 243: 101-109.

Caley M., Carr M., Hixon M.A., Hughes T.P., Jones G.P, Menge B.A. 1996. Recruitment and the local dynamics of open marine populations. Annu. Rev. Ecol. Syst. 27: 477-500.

Chapman A.R.O. 1990. Effects of grazing, canopy cover and substratum type on the abundances of common species of seaweeds inhabiting littoral fringe tide pools. Bot. Mar. 33: 319-326.

Cheshire A.C., Westphalen G., Wenden A., Scriven L.J., Rowland B.C. 1996. Photosynthesis and respiration of phaeophycean-dominated macroalgal communities in summer and winter. Aquat. Bot. 55: 159-170.

Clarke K.R. 1993. Nonparametric multivariate analyses of changes in community structure. Austral. J. Ecol. 18: 117-143.

Coelho H.S., Neves R.J.J., White M., Leitão P.C., Santos A.J. 2002. A model for ocean circulation on the Iberian coast. J. Mar. Syst. 32: 153-179.

Davison I.R., Pearson G.A. 1996. Stress tolerance in intertidal seaweeds. J. Phycol. 32: 197-211.

Desroy N., Dubois S.F., Fournier J., Ricquiers L., Le Mao P., Guerin L., Gerla D., Rougerie M., Legendre A. 2011. The conservation status of Sabellaria alveolata (L.) (Polychaeta: Sabellariidae) reefs in the Bay of Mont-Saint-Michel. Aquat. Conserv.: Mar. Freshwat. Ecosyst. 21: 462-471.

Dethier M.N. 1982. Pattern and process in tidepool algae: factors influencing seasonality and distribution. Bot. Mar. 25: 55-66.

Dethier M.N. 1984. Disturbance and recovery in intertidal pools: maintenance of mosaic patterns. Ecol. Monogr. 54: 99-118.

Dethier M.N., Graham E.S., Cohen S., Tear L.M. 1993. Visual versus random-point percent cover estimations: "objective" is not always better. Mar. Ecol. Prog. Ser. 96: 93-100.

Dias J.M.A., Gonzalez R., Garcia C., Diaz-del Rio V. 2002. Sediment distribution patterns on the Galicia-Minho continental shelf. Prog. Oceanogr. 52: 215-231.

Díez I., Santolaria A., Secilla A., Gorostiaga J.M. 2009. Recovery stages over long- term monitoring of the intertidal vegetation in the Abra de Bilbao area and on the adjacent coast (N. Spain). Eur. J. Phycol. 4: 1-14.

Dubois S., Retière C., Olivier F. 2002. Biodiversity associated with Sabellaria alveolata (Polychaeta: Sabellariidae) reefs: effects of human disturbances. J. Mar. Biol. Assoc. U.K. 82: 817-826.

Dubois S., Commito J.A., Olivier F., Retière C. 2006. Effects of epibionts on Sabellaria alveolata (L.) biogenic reefs and their associated fauna in the Bay of Mont Saint-Michel. Estuar. Coast. Shelf Sci. 68: 635-646.

Dubois S., Comtet T., Retière C., Thiébaut E. 2007. Distribution and retention of Sabellaria alveolata larvae (Polychaeta: Sabellariidae) in the Bay of Mont-Saint-Michel, France. Mar. Ecol. Prog. Ser. 344: 15-28.

Femino R., Mathieson A. 1980. Investigations of New England marine algae IV. The ecology and seasonal succession of tide pool algae at Bald Head Cliff, York, Maine, USA. Bot. Mar. 23: 319-332.

Fournier J., Etienne S., Le Cam J.B. 2010. Inter- and intraspecific variability in the chemical composition of the mineral phase of cements from several tube-building polychaetes. Geobios 43: 191-200.

Fraschetti S., Terlizzi A., Benedetti-Cecchi L. 2005. Patterns of distribution of marine assemblages from rocky shores: evidence of relevant scales of variation. Mar. Ecol. Prog. Ser. 296: 13-29.

Godet L., Fournier J., Jaffré M., Desroy N. 2011. Influence of stability and fragmentation of a worm-reef on benthic macrofauna. Estuar. Coast. Shelf Sci. 92: 472-479.

Golléty C., Migné A., Davoult D. 2008. Benthic metabolism on a sheltered rocky shore: role of the canopy in the carbon budget. J. Phycol. 44: 1146-1153.

Guerra-García J.M., Cabezas M.P., Baeza-Rojano E., García-Gómez J.C. 2011. Spatial patterns and seasonal fluctuations of intertidal macroalgal assemblages from Tarifa Island, southern Spain: relationship with associated Crustacea. J. Mar. Biol. Assoc. U.K. 91: 107-116.

Guimaraens M.A., Coutinho R. 2000. Temporal and spatial variation of Ulva spp. And water properties in the Cabo Frio upwelling region of Brazil. Aquat. Bot. 66: 101-114.

Guiry M.D., Cunningham E.M. 1984. Photoperiodic and temperature responses in the reproduction of north-eastern Atlantic Gigartina acicularis (Rhodophyta: Gigartinales). Phycologia 23: 357-367.

Hewitt J.E., Thrush S.F., Cummings V.J., Turner S.J. 1998. The effect of changing sampling scales on our ability to detect effects of large-scale processes on communities. J. Exp. Mar. Biol. Ecol. 227: 251-264.

Huggett J., Griffiths C.L. 1986. Some relationships between elevation, physico-chemical variables and biota of intertidal rock pools. Mar. Ecol. Prog. Ser. 29: 189-197.

Huthnance J.M., Van Aken H.M., White M., Barton E.D., LeCann B., Coelho E.F., Fanjul E.A., Miller P., Vitorino J. 2002. Ocean margin exchange-water flux estimates. J. Mar. Syst. 32: 107-137.

Izquierdo J.L., Pérez-Ruzafa I.M., Gallardo T. 2002. Effect of temperature and photon fluence rate on gametophytes and young sporophytes of Laminaria ochroleuca Pylaie. Helgoland Mar. Res. 55: 285-292

Konar B., Foster M.S. 1992. Distribution and recruitment of subtidal geniculate coralline algae. J. Phycol. 28: 273-280.

Kooistra W.H.C.F., Joosten A.M.T., van den Hoek C. 1989. Zonation patterns in intertidal pools and their possible causes: a multivariate approach. Bot. Mar. 32: 9-26.

Lawton J.H. 1996. Patterns in ecology. Oikos 75: 145-147.

Lemos R.T., Pires H.O. 2004. The upwelling regime off the west Portuguese coast, 1941–2000. Int. J. Climat. 24: 511-524.

Levin S.A. 1992. The problem of pattern and scale in ecology. Ecology 73: 1943-1967.

Littler M.M., Kauker B.J. 1984. Heterotrichy and survival strategies in the red alga Corallina officinalis L. Bot. Mar. 27: 37-44.

Masterson P., Arenas F.A., Thompson R.C., Jenkins S.R. 2008. Interaction of top down and bottom up factors in intertidal rockpools: Effects on early successional macroalgal community composition, abundance and productivity. J. Exp. Mar. Biol. Ecol. 363: 12-20.

McArdle B.H., Anderson M.J. 2001. Fitting multivariate models to community data: a comment on distance-based redundancy analysis. Ecology 82: 290-297.[0290:FMMTCD]2.0.CO;2

Menge B.A., Ashkenas L.R., Matson A. 1983. Use of artificial holes in studying community development in cryptic marine habitats in a tropical rocky intertidal region. Mar. Biol. 77: 129-142.

Metaxas A., Scheibling R.E. 1993. Community structure and organization of tidepools. Mar. Ecol. Prog. Ser. 98: 187-198.

Metaxas A., Hunt H.L., Scheibling R.E. 1994. Spatial and temporal variability of macrobenthic communities in tidepools on a rocky shore in Nova Scotia, Canada. Mar. Ecol. Prog. Ser. 105: 89-103.

Neto A.I. 2000. Observations on the biology and ecology of selected macroalgae from the littoral of São Miguel (Azores). Bot. Mar. 43: 483-498.

Pedersen M.F., Borum J., Fotel F.L. 2010. Phosphorus dynamics and limitation of fast- and slow-growing temperate seaweeds in Oslofjord, Norway. Mar. Ecol. Prog. Ser. 399: 103-115.

Reed D.C., Laur D.R., Ebeling A.W. 1988. Variation in algal dispersal and recruitment: the importance of episodic events. Ecol. Monogr. 58: 321-335.

Roberts D.E., Smith A., Ajani P., Davis A.R. 1998. Rapid changes in encrusting marine assemblages exposed to anthropogenic point-source pollution: a beyond BACI approach. Mar. Ecol. Prog. Ser. 163: 213-224.

Rubal M., Veiga P., Vieira R., Sousa-Pinto I. 2011. Seasonal patterns of tidepool macroalgal assemblages in the North of Portugal. Consistence between species and functional group approaches. J. Sea Res. 66: 187-194.

Sánchez I., Fernández C. 2005. Impact of the invasive seaweed Sargassum muticum (Phaeophyta) on an intertidal macroalgal assemblage. J. Phycol. 41: 923-930.

Sánchez I., Fernández C., Arrontes J. 2005. Long-term changes in the structure of intertidal assemblages after invasion by Sargassum muticum (Phaeophyta). J. Phycol. 41: 942-949.

Steen H. 2003. Intraspecific competition in Sargassum muticum (Phaeophyceae) germlings under various density, nutrient and temperature regimes. Bot. Mar. 46: 36-43.

Steneck R.S. 1986. The ecology of coralline algal crusts: convergent patterns and adaptive strategies. Annu. Rev. Ecol. Syst. 17: 273-303.

Terlizzi A., Anderson M.J., Fraschetti S., Benedetti-Cecchi L. 2007. Scales of spatial variation in Mediterranean subtidal sessile assemblages at different depths. Mar. Ecol. Prog. Ser. 332: 25-39.

Thompson R.C., Crowe T.P., Hawkins S.J. 2002. Rocky intertidal communities: past environmental changes, present status and predictions for the next 25 years. Environ. Conserv. 29: 168-191.

tom Dieck I. 1992. North Pacific and North Atlantic digitate Laminaria species (Phaeophyta): hybridization experiments and temperature responses. Phycologia 31: 147-163.

Underwood A.J. 1997. Experiments in Ecology: their logical design and interpretation using analysis of variance. Cambridge University Press, Cambridge, UK.

Underwood A.J., Chapman M.G. 1996. Scales of spatial patterns of distribution of intertidal invertebrates. Oecologia 107: 212-224.

Underwood A.J., Petraitis P.S. 1993. Structure of intertidal assemblages in different locations: how can local processes be compared? In: Ricklefs R., Schutler D. (eds.) Species diversity in ecological communities. Univ. Chicago Press, Chicago, pp. 38-51. PMid :8480505

Underwood A.J., Skilleter G.A. 1996. Effects of patch-size on the structure of assemblages in rock pools. J. Exp. Mar. Biol. Ecol. 197: 63-90.

Underwood A.J., Chapman M.G., Connell S.D. 2000. Observations in ecology: you can't make progress on processes without understanding the patterns. J. Exp. Mar. Biol. Ecol. 250: 97-115.

Valdivia N., Scrosati R.A., Molis M., Knox A.S. 2011. Variation in community structure across vertical intertidal stress gradients: how does it compare with horizontal variation at different scales? PLoS ONE 6(8): e24062. PMid :21887371 PMCid:3161098

Van Tamelen P.G. 1996. Algal zonation in tidepools: experimental evaluation of the roles of physical disturbance, herbivory and competition. J. Exp. Mar. Biol. Ecol. 201: 197-231.

Viejo R.M. 1997. The effects of colonization by Sargassum muticum on tidepool macroalgal assemblages. J. Mar. Biol. Assoc. U.K. 77: 325-40.

Wilson D.P. 1970. Additional observations on larval growth and settlement of Sabellaria alveolata. J. Mar. Biol. Assoc. U.K. 50: 1-31.

Wolfe J., Harlin M. 1988a. Tidepools in Southern Rhode Island, U.S.A. I. Distribution and seasonality of macroalgae. Bot. Mar. 31: 525-536.

Wolfe J., Harlin M. 1988b. Tidepools in Southern Rhode Island, U.S.A. II. Species diversity and similarity analysis of macroalgal communities. Bot. Mar. 31: 537-546.

Copyright (c) 2012 Consejo Superior de Investigaciones Científicas (CSIC)

Creative Commons License
This work is licensed under a Creative Commons Attribution 4.0 International License.

Contact us

Technical support