Temporal variation of sandy beach macrofauna at two sites with distinct environmental conditions on Cassino Beach, extreme southern Brazil

Silva, Pedro de Sá Rodrigues da; Neves, Luciano Peyrer das; Bemvenuti, Carlos Emílio

Abstracts

Temporal variations of the macrofauna of sandy beaches have been related to variations in the beach morphodynamics and also to the population dynamics of dominant species. The aim of this article is to describe the temporal variation of the intertidal macrofauna at two sites with distinct environmental condition on Cassino Beach, extreme southern Brazil. At each site three transect lines 50 m apart were defined perpendicular to the shore line, from which samples were collected monthly in triplicate at 4 intertidal levels (10 m apart) from June 2004 to May 2005. During winter a generally low density was observed, due to the absence of recruitments and to the mud deposition, which occurred just before sampling (in April 2004), and to low intensity stranding events. Spring witnessed a population explosion of Scolelepis gaucha, a migration of Mesodesma mactroides adults from the subtidal zone, and a strong stranding event. In the summer, recruitment of M. mactroides, Donax hanleyanus and Emerita brasiliensis was observed. Fall was characterized by low densities, except for D. hanleyanus recruitment. The macrofauna at both sites showed a striking seasonal variation in density and diversity, perhaps attributable to the recruitment of numerically dominant species and physical disturbances (stranding and mud deposition).

Sandy beaches; Benthic macrofauna; Temporal variation; Cassino Beach

Variações sazonais da macrofauna bentônica de praias arenosas têm sido relacionadas com variações da morfodinâmica da praia e também aos recrutamentos das espécies dominantes. Este trabalho objetiva avaliar a variabilidade temporal da macrofauna da zona entremarés de dois locais com distintas características ambientais na praia do Cassino, extremo sul do Brasil. Em cada local foram demarcadas três transversais (separadas por 50m) perpendiculares à linha de água, nas quais amostras foram coletadas em triplicata em 4 níveis entremarés (separados por 10 m), entre junho/2004 e maio/2005. Durante o inverno ocorreram baixas densidades da macrofauna, que foram relacionadas à ausência de recrutamentos, à deposição de lama no mês de abril e à embancamentos de baixa intensidade. Na primavera foi registrada a explosão populacional de Scolelepis gaucha, migração de adultos de Mesodesma mactroides do infralitoral e um evento de embancamento. Durante o verão destacaram-se os recrutamentos dos migradores mareais M. mactroides, Donax hanleyanus e Emerita brasiliensis. O outono foi caracterizado por baixas densidades, exceto pelo recrutamento de D. hanleyanus. A macrofauna bentônica nos dois locais mostrou variações sazonais na densidade e diversidade, que podem ser atribuídas ao recrutamento das espécies dominantes e a ocorrência de perturbações físicas (embancamento e depósito de lama).

Praias arenosas; Macrofauna bentônica; Variabilidade temporal; Praia do Cassino

BARROS, F.; CALLIARI, L. J.; GIANUCA, N. M. Estudo preliminar da macrofauna bentonica da praia dos Concheiros. Notas Técnicas CECO/UFRGS, v. 7, p. 33-37, 1994.
BARROS, F.; BORZONE, C. A.; ROSSO, S. Macroinfauna of six beaches near Guaratuba Bay, Southern Brazil. Braz. Arch. Biol. Technol, v. 44, n. 4, p. 351-364, 2001.
BORZONE C. A.; SOUZA, J. R. B. Estrutura da macrofauna bentônica no supra, meso e infralitoral de uma praia arenosa do sul do Brasil. Oecol. Brasiliensis, v.3, p. 197-212, 1997.
BORZONE, C. A.; SOUZA, J. R. B.; SOARES, A. G. Morphodynamic influence on the structure of inter and subtidal macrofaunal communities of subtropical sandy beaches. Rev. Chil. Hist. Nat,v. 69, p. 565-577, 1996.
BRAZEIRO, A.; DEFEO, O. Effects of harvesting and density dependence on the demography of sandy beach populations: The yellow clam Mesodesma mactroides in Uruguay. Mar. Ecol. Prog. Ser, v. 182, p. 127-135, 1999.
BROWN, A. C. ; MCLACHLAN, A. Ecology of sandy shores Amsterdam: Elsevier, 1990. 327 p.
CALLIARI, L. J. ; FACHIN, S. Laguna dos Patos. Influência nos depósitos lamíticos costeiros. Pesquisas, v. 20, n. 1, p. 57-69, 1993.
CALLIARI, L. J; KLEIN, A. H. F. Características morfodinâmicas e sedimentológicas das praias oceânicas entre Rio Grande e Chuí, RS. Pesquisas, v. 20, n. 1, p. 48-56, 1993.
CARDOSO, R. S.; VELOSO, V. G. Population dynamics and secondary production of the wedge clam Donax hanleyanus (Bivalvia: Donacidae) on a high energy subtropical beach of Brazil. Mar. Biol, v. 142, n. 1, p. 153-162, 2003.
CLARKE, K. R.; WARWICK, R. M. Change in marine communities: an approach to statistical analysis and interpretation. Plymouth: Natural Environment Research Council, 1994. 144 p.
COSCARÓN, S. La Almeja Amarilla (Mesodesma (T.) mactroides Deshayes) de la costa de la Província de Buenos Aires. Agro. Publ. Tec, v. 1, p. 1-66, 1959.
DEFEO, O. Recruitment variability in sandy beach macroinfauna: much to learn yet. Revta Chil. Hist. Nat., v. 69, p. 615-630, 1996.
DEFEO, O.; SCARABINO, V. Ecological significance of a possible deposit-feeding strategy in Mesodesma mactroides (Deshayes, 1854) (Mollusca: Pelecypoda). Atlântica, v. 12, p. 55-65, 1990.
DEFEO, O.; DE ALAVA, A. Effects of human activities on long-term trends in sandy beach populations: the wedge clam Donax hanleyanus in Uruguay. Mar. Ecol. Prog. Ser, v. 123, p. 73-82, 1995.
DEFEO, O.; RUEDA, M. Spatial structure, sampling design and abundance estimates in sandy beach macroinfauna: some warnings and new perspectives. Mar. Biol, v. 140, p. 1215-1225, 2002.
DEFEO, O.; CARDOSO, R. S. Latitudinal patterns in abundance and life-history traits of the mole crab Emerita brasiliensis on South American sandy beaches. Diversity Distrib, v. 10, n. 2, p.89-98, 2004.
DEFEO, O.; ORTIZ, E.; CASTILLA, J. C. Growth, mortality and recruitment of the yellow clam Mesodesma mactroides on Uruguayan beaches. Mar. Biol, v. 114, p. 429-437, 1992.
DEFEO, O.; BRAZEIRO, A.; DE ALAVA, A.; RIESTRA, G. Is sandy beach macrofauna only physically controlled? Role of substrate and competition in isopods. Estuar. coast. Shelf Sci, v. 45, p. 453-462, 1997.
DEXTER, D. M. Community structure and seasonal variation in intertidal Panamanian sandy beaches. Estuar. coast. mar. Sci, v. 9, p. 543-558, 1979.
DEXTER, D. M. Temporal and spatial variability in the community structure of the fauna of four sandy beaches in south-eastern New South Wales. Aust. J. mar. freshwat. Res,v. 35, n. 6, p. 663-672, 1984.
DEXTER, D. M. Distribution and life histories of abundant crustaceans of four sandy beaches of south-eastern New South Wales. Aust. J. mar. freshw. Res., v. 36, n. 2, p:281-289, 1985.
DUNGAN, J. E.; MCLACHLAN, A. An assessment of longshore movement in Donax serra Roding (Bivalvia: Donacidae) on an exposed sandy beach. J. exp. mar. Biol. Ecol, v. 234 , n. 1, p. 111-124, 1999.
EFFORD, I. E. 1976. Distribution of the sand crab in the genus Emerita (Decapoda: Hippidae). Crustaceana, v. 30, p. 169-183, 1976.
ELEFTHERIOU, A.; NICHOLSON, M. D. 1975. The effect of exposure on beaches fauna. Cah. Biol. mar., v. 16, p. 685-710, 1975.
GIANUCA, N. M. A preliminary account of the ecology of sandy beaches in southern Brazil. In: MCLACHLAN, A.; ERASMUS, T . (Ed.) Sandy beaches as ecosystems The Hauge: Dr. W. Junk Publishers, 1983. p. 413-420.
HALL, S. J. Physical disturbance and marine benthic communities: Life in unconsolidated sediments. Oceanogr. mar. Biol. a. Rev, v. 32, p. 179-239, 1994.
HAYNES, D.; QUINN, G. P. Temporal and spatial variability in community structure of a sandy intertidal beach, Cape Paterson, Victoria, Australia. Mar. freshw. Res, v. 46, n. 6, p. 931-942, 1995.
HOLLAND, A. F.; POLGAR, T. T. Seasonal changes in the structure of an intertidal community. Mar. Biol, v. 37, n. 4, p. 341-348, 1976.
JARAMILLO, E.; MCLACHLAN, A.; COETZEE, P. Intertidal zonation patterns of macroinfauna over a range of exposed sandy beaches in south-central Chile. Mar. Ecol. Prog. Ser. , v.101, p. 105-118, 1993.
JARAMILLO, E.; CONTRERAS, H.; QUINJON, P. Macroinfauna and human disturbance in Chile. Revta Chil. Hist. Nat., v. 69, p. 655-663, 1996.
LEBER, K. M. 1982. Seasonality of macroinvertebrates on a temperate, high wave, energy sandy beach. Bull. mar. Sci., v. 32, p. 86-98, 1982.
MCLACHLAN, A. Dissipative beaches and macrofaunal communities on exposed intertidal sands. J. coast. Res, v. 6, p. 57-71, 1990.
MCLACHLAN, A. Physical factors in benthic ecology: effects of changing sand particle size on beach fauna. Mar. Ecol. Prog. Ser, v.131, p. 205-211, 1996.
MCLACHLAN, A.; JARAMILLO, E. Zonation on sandy beaches. Oceanogr. mar. Biol. a. Rev, v. 33, p. 305-335, 1995.
MCLACHLAN, A.; WOODRIDGE, T.; DYE, A. H. The ecology of sandy beaches in southern Africa. S. Afr. J. Zool., v.16, p. 219-231, 1981.
MCLACHLAN, A.; JARAMILLO, E.; DONN, T. E.; WESSELS, E. Sandy beach macrofauna communities and their control by the physical environment: a geographical comparison. J. coast. Res, v.15, p. 27-38, 1993.
MÉNDES, S. Bivalve mortality in southwest Atlantic shores. Harmful Algae News, v. 10, p. 11-12, 1995.
NEVES, F. M.; BEMVENUTI, C. E. Spatial distribution of macrobenthic fauna on three sandy beaches from northern Rio Grande do Sul, southern Brazil. Braz. J. Oceanogr,v. 54, n. 2, p. 135-145, 2006.
ODEBRECHT, C.; RORIG, L.; GARCIA, V. T.; ABREU, P. C. Shellfish mortality and red tide event in southern Brazil. In: LASSUS, P. (Ed.). Harmful algae Blooms Berlin: Springer-Verlag, 1995. p. 213-218.
OLIVIER, S.; CAPEZZANI, D.; CARRETO, J.; CHISTIANSEN, H.; MORENO, V.; DE MORENO, J. A.; PENCHASZADEH & OLIVIER, S. 1975. Ecología de una población de "Berberecho" (Donax hanleyanus) en Villa Gesell, Argentina. Malacologia, v.15, p. 133-146, 1975.
RAMÍREZ, F. C.; MIANZAN, H.; CHIAVERANO, L. 2004. Varamientos y arribazones. In: BOSCHI, E. E.; COUSSEAU, M.B. (Ed.). La vida entre mareas: vegetales e animales de las costas de Mar del Plata, Argentina. Mar del Plata: INIDEP, 2004. p. 59-64. (Publnes especial.
SANTOS, P. J. P. 1994. Population dynamics and production of Scolelepis gaucha (Polychaeta: Spionidae) on the sandy beaches of Southern Brazil. Mar. Ecol. Prog. Ser, v. 110, p. 159-165, 1994.
SCHOEMAN, D. S.; WHELLER, M.; WAIT, M. The relative accuracy of the standard estimators of macrofaunal abundance and species richness from selected intertidal transect designs used to sample exposed sand beaches. Estuar. coast. Shelf Sci, v.58, p. 5-16, 2003.
SHORT, A. D.; WRIGHT, L. D. Physical variability of sandy beaches. In: MCLACHLAN, A.; ERASMUS, T. (Ed.). Sandy beaches as ecosystems The Hague: Dr. W. Junk Publishers, 1983. p. 133-144
SOUZA, J. R. B.; GIANUCA, N. M. Zonation and seasonal variation of the intertidal macrofauna on a sandy beach of Paraná State, Brazil. Sci. Mar, v.59, n.2, p. 103-111, 1995.
YANNICELLI, B.; PALÁCIOS, R.; GIMÉNEZ, L. Swimming ability and burrowing time of two cirolanid isopods from different levels of exposed sandy beaches. J. expl mar. Biol. Ecol, v. 273, p. 73-88, 2002.
UNDERWOOD, A. J. Experiments in ecology: their logical design and interpretation using analysis of variance. Cambridge: Cambridge University Press, 1997. 504 p.
VELOSO, V. G.; CARDOSO, R. S. 2001. Effects of morphodynamics on the spatial and temporal variation of the macrofuana on three sandy beaches, Rio de Janeiro State, Brazil. J. mar. Biol. Assoc. U.K., v. 81, p. 369-375, 2001.
VELOSO, V. G.; CARDOSO, R. S.; FONSECA, D. B. 1997. Adaptações e biologia da macrofauna de praias arenosas expostas com ênfase nas espécies da região entre-marés do litoral Fluminense. Oecol. Brasiliensis, v. 3, p. 135-154, 1997.
FIGUEIREDO, S. A. Sedimentologia e morfodinâmica das praias oceânicas adjacentes às desembocaduras lagunares e fluviais do RS. In: Congresso da Associação Brasileira de Estudos do Quartenário, 9, 2003, Recife. Anais ..., v. 1. Recife, 2003. p. 136.
PEREIRA, P. S. 2004. Variabilidade da orla oceânica da costa do Rio Grande do Sul e suas implicações na elaboração de planos de contingência: aspectos morfodinâmicos, sedimentológicos e geomorfológicos. Master's thesis, Fundação Universidade Federal do Rio Grande, 160 p.

Publication Dates

Publication in this collection
26 Jan 2009
Date of issue
Dec 2008

History

Accepted
20 June 2008
Reviewed
14 Apr 2008
Received
27 June 2006

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

[1] BARROS, F.; CALLIARI, L. J.; GIANUCA, N. M. Estudo preliminar da macrofauna bentonica da praia dos Concheiros. Notas Técnicas CECO/UFRGS, v. 7, p. 33-37, 1994.

[2] BARROS, F.; BORZONE, C. A.; ROSSO, S. Macroinfauna of six beaches near Guaratuba Bay, Southern Brazil. Braz. Arch. Biol. Technol, v. 44, n. 4, p. 351-364, 2001.

[3] BORZONE C. A.; SOUZA, J. R. B. Estrutura da macrofauna bentônica no supra, meso e infralitoral de uma praia arenosa do sul do Brasil. Oecol. Brasiliensis, v.3, p. 197-212, 1997.

[4] BORZONE, C. A.; SOUZA, J. R. B.; SOARES, A. G. Morphodynamic influence on the structure of inter and subtidal macrofaunal communities of subtropical sandy beaches. Rev. Chil. Hist. Nat,v. 69, p. 565-577, 1996.

[5] BRAZEIRO, A.; DEFEO, O. Effects of harvesting and density dependence on the demography of sandy beach populations: The yellow clam Mesodesma mactroides in Uruguay. Mar. Ecol. Prog. Ser, v. 182, p. 127-135, 1999.

[6] BROWN, A. C. ; MCLACHLAN, A. Ecology of sandy shores Amsterdam: Elsevier, 1990. 327 p.

[7] CALLIARI, L. J. ; FACHIN, S. Laguna dos Patos. Influência nos depósitos lamíticos costeiros. Pesquisas, v. 20, n. 1, p. 57-69, 1993.

[8] CALLIARI, L. J; KLEIN, A. H. F. Características morfodinâmicas e sedimentológicas das praias oceânicas entre Rio Grande e Chuí, RS. Pesquisas, v. 20, n. 1, p. 48-56, 1993.

[9] CARDOSO, R. S.; VELOSO, V. G. Population dynamics and secondary production of the wedge clam Donax hanleyanus (Bivalvia: Donacidae) on a high energy subtropical beach of Brazil. Mar. Biol, v. 142, n. 1, p. 153-162, 2003.

[10] CLARKE, K. R.; WARWICK, R. M. Change in marine communities: an approach to statistical analysis and interpretation. Plymouth: Natural Environment Research Council, 1994. 144 p.

[11] COSCARÓN, S. La Almeja Amarilla (Mesodesma (T.) mactroides Deshayes) de la costa de la Província de Buenos Aires. Agro. Publ. Tec, v. 1, p. 1-66, 1959.

[12] DEFEO, O. Recruitment variability in sandy beach macroinfauna: much to learn yet. Revta Chil. Hist. Nat., v. 69, p. 615-630, 1996.

[13] DEFEO, O.; SCARABINO, V. Ecological significance of a possible deposit-feeding strategy in Mesodesma mactroides (Deshayes, 1854) (Mollusca: Pelecypoda). Atlântica, v. 12, p. 55-65, 1990.

[14] DEFEO, O.; DE ALAVA, A. Effects of human activities on long-term trends in sandy beach populations: the wedge clam Donax hanleyanus in Uruguay. Mar. Ecol. Prog. Ser, v. 123, p. 73-82, 1995.

[15] DEFEO, O.; RUEDA, M. Spatial structure, sampling design and abundance estimates in sandy beach macroinfauna: some warnings and new perspectives. Mar. Biol, v. 140, p. 1215-1225, 2002.

[16] DEFEO, O.; CARDOSO, R. S. Latitudinal patterns in abundance and life-history traits of the mole crab Emerita brasiliensis on South American sandy beaches. Diversity Distrib, v. 10, n. 2, p.89-98, 2004.

[17] DEFEO, O.; ORTIZ, E.; CASTILLA, J. C. Growth, mortality and recruitment of the yellow clam Mesodesma mactroides on Uruguayan beaches. Mar. Biol, v. 114, p. 429-437, 1992.

[18] DEFEO, O.; BRAZEIRO, A.; DE ALAVA, A.; RIESTRA, G. Is sandy beach macrofauna only physically controlled? Role of substrate and competition in isopods. Estuar. coast. Shelf Sci, v. 45, p. 453-462, 1997.

[19] DEXTER, D. M. Community structure and seasonal variation in intertidal Panamanian sandy beaches. Estuar. coast. mar. Sci, v. 9, p. 543-558, 1979.

[20] DEXTER, D. M. Temporal and spatial variability in the community structure of the fauna of four sandy beaches in south-eastern New South Wales. Aust. J. mar. freshwat. Res,v. 35, n. 6, p. 663-672, 1984.

[21] DEXTER, D. M. Distribution and life histories of abundant crustaceans of four sandy beaches of south-eastern New South Wales. Aust. J. mar. freshw. Res., v. 36, n. 2, p:281-289, 1985.

[22] DUNGAN, J. E.; MCLACHLAN, A. An assessment of longshore movement in Donax serra Roding (Bivalvia: Donacidae) on an exposed sandy beach. J. exp. mar. Biol. Ecol, v. 234 , n. 1, p. 111-124, 1999.

[23] EFFORD, I. E. 1976. Distribution of the sand crab in the genus Emerita (Decapoda: Hippidae). Crustaceana, v. 30, p. 169-183, 1976.

[24] ELEFTHERIOU, A.; NICHOLSON, M. D. 1975. The effect of exposure on beaches fauna. Cah. Biol. mar., v. 16, p. 685-710, 1975.

[25] GIANUCA, N. M. A preliminary account of the ecology of sandy beaches in southern Brazil. In: MCLACHLAN, A.; ERASMUS, T . (Ed.) Sandy beaches as ecosystems The Hauge: Dr. W. Junk Publishers, 1983. p. 413-420.

[26] HALL, S. J. Physical disturbance and marine benthic communities: Life in unconsolidated sediments. Oceanogr. mar. Biol. a. Rev, v. 32, p. 179-239, 1994.

[27] HAYNES, D.; QUINN, G. P. Temporal and spatial variability in community structure of a sandy intertidal beach, Cape Paterson, Victoria, Australia. Mar. freshw. Res, v. 46, n. 6, p. 931-942, 1995.

[28] HOLLAND, A. F.; POLGAR, T. T. Seasonal changes in the structure of an intertidal community. Mar. Biol, v. 37, n. 4, p. 341-348, 1976.

[29] JARAMILLO, E.; MCLACHLAN, A.; COETZEE, P. Intertidal zonation patterns of macroinfauna over a range of exposed sandy beaches in south-central Chile. Mar. Ecol. Prog. Ser. , v.101, p. 105-118, 1993.

[30] JARAMILLO, E.; CONTRERAS, H.; QUINJON, P. Macroinfauna and human disturbance in Chile. Revta Chil. Hist. Nat., v. 69, p. 655-663, 1996.

[31] LEBER, K. M. 1982. Seasonality of macroinvertebrates on a temperate, high wave, energy sandy beach. Bull. mar. Sci., v. 32, p. 86-98, 1982.

[32] MCLACHLAN, A. Dissipative beaches and macrofaunal communities on exposed intertidal sands. J. coast. Res, v. 6, p. 57-71, 1990.

[33] MCLACHLAN, A. Physical factors in benthic ecology: effects of changing sand particle size on beach fauna. Mar. Ecol. Prog. Ser, v.131, p. 205-211, 1996.

[34] MCLACHLAN, A.; JARAMILLO, E. Zonation on sandy beaches. Oceanogr. mar. Biol. a. Rev, v. 33, p. 305-335, 1995.

[35] MCLACHLAN, A.; WOODRIDGE, T.; DYE, A. H. The ecology of sandy beaches in southern Africa. S. Afr. J. Zool., v.16, p. 219-231, 1981.

[36] MCLACHLAN, A.; JARAMILLO, E.; DONN, T. E.; WESSELS, E. Sandy beach macrofauna communities and their control by the physical environment: a geographical comparison. J. coast. Res, v.15, p. 27-38, 1993.

[37] MÉNDES, S. Bivalve mortality in southwest Atlantic shores. Harmful Algae News, v. 10, p. 11-12, 1995.

[38] NEVES, F. M.; BEMVENUTI, C. E. Spatial distribution of macrobenthic fauna on three sandy beaches from northern Rio Grande do Sul, southern Brazil. Braz. J. Oceanogr,v. 54, n. 2, p. 135-145, 2006.

[39] ODEBRECHT, C.; RORIG, L.; GARCIA, V. T.; ABREU, P. C. Shellfish mortality and red tide event in southern Brazil. In: LASSUS, P. (Ed.). Harmful algae Blooms Berlin: Springer-Verlag, 1995. p. 213-218.

[40] OLIVIER, S.; CAPEZZANI, D.; CARRETO, J.; CHISTIANSEN, H.; MORENO, V.; DE MORENO, J. A.; PENCHASZADEH & OLIVIER, S. 1975. Ecología de una población de "Berberecho" (Donax hanleyanus) en Villa Gesell, Argentina. Malacologia, v.15, p. 133-146, 1975.

[41] RAMÍREZ, F. C.; MIANZAN, H.; CHIAVERANO, L. 2004. Varamientos y arribazones. In: BOSCHI, E. E.; COUSSEAU, M.B. (Ed.). La vida entre mareas: vegetales e animales de las costas de Mar del Plata, Argentina. Mar del Plata: INIDEP, 2004. p. 59-64. (Publnes especial.

[42] SANTOS, P. J. P. 1994. Population dynamics and production of Scolelepis gaucha (Polychaeta: Spionidae) on the sandy beaches of Southern Brazil. Mar. Ecol. Prog. Ser, v. 110, p. 159-165, 1994.

[43] SCHOEMAN, D. S.; WHELLER, M.; WAIT, M. The relative accuracy of the standard estimators of macrofaunal abundance and species richness from selected intertidal transect designs used to sample exposed sand beaches. Estuar. coast. Shelf Sci, v.58, p. 5-16, 2003.

[44] SHORT, A. D.; WRIGHT, L. D. Physical variability of sandy beaches. In: MCLACHLAN, A.; ERASMUS, T. (Ed.). Sandy beaches as ecosystems The Hague: Dr. W. Junk Publishers, 1983. p. 133-144

[45] SOUZA, J. R. B.; GIANUCA, N. M. Zonation and seasonal variation of the intertidal macrofauna on a sandy beach of Paraná State, Brazil. Sci. Mar, v.59, n.2, p. 103-111, 1995.

[46] YANNICELLI, B.; PALÁCIOS, R.; GIMÉNEZ, L. Swimming ability and burrowing time of two cirolanid isopods from different levels of exposed sandy beaches. J. expl mar. Biol. Ecol, v. 273, p. 73-88, 2002.

[47] UNDERWOOD, A. J. Experiments in ecology: their logical design and interpretation using analysis of variance. Cambridge: Cambridge University Press, 1997. 504 p.

[48] VELOSO, V. G.; CARDOSO, R. S. 2001. Effects of morphodynamics on the spatial and temporal variation of the macrofuana on three sandy beaches, Rio de Janeiro State, Brazil. J. mar. Biol. Assoc. U.K., v. 81, p. 369-375, 2001.

[49] VELOSO, V. G.; CARDOSO, R. S.; FONSECA, D. B. 1997. Adaptações e biologia da macrofauna de praias arenosas expostas com ênfase nas espécies da região entre-marés do litoral Fluminense. Oecol. Brasiliensis, v. 3, p. 135-154, 1997.

[50] FIGUEIREDO, S. A. Sedimentologia e morfodinâmica das praias oceânicas adjacentes às desembocaduras lagunares e fluviais do RS. In: Congresso da Associação Brasileira de Estudos do Quartenário, 9, 2003, Recife. Anais ..., v. 1. Recife, 2003. p. 136.

[51] PEREIRA, P. S. 2004. Variabilidade da orla oceânica da costa do Rio Grande do Sul e suas implicações na elaboração de planos de contingência: aspectos morfodinâmicos, sedimentológicos e geomorfológicos. Master's thesis, Fundação Universidade Federal do Rio Grande, 160 p.