Vertical distribution of benthic marine organisms on rocky coasts of the Fernando de Noronha Archipelago (Brazil)

Eston, Verena Rapp de; Migotto, Alvaro Esteves; Oliveira Filho, Eurico Cabral de; Rodrigues, Sergio de Almeida; Freitas, Jose Carlos de

doi:10.1590/S0373-55241986000100004

Abstract

Qualitative samples of flora and fauna were collected in several points, from the supralittoral fringe down to 30 m, using mainly aqualungs. Species distribution and abundance were determined along four transects, up to 200 m long, in selected places. Samples were taken from 25 x 25 cm quadrats positioned at fixed intervals along the transects down to a variable depth, depending on algal percentage cover and their substrate. The intertidal zone was populated essentially by molluscs. The subtidal was covered predominantly by macroalgae. Montastrea cavernosa, Siderastrea stellata and Mussismillia hispida were the only animal species with a significant percentage cover within the quadrats. Dictyopteris justii, D. plagiograma, Dictyota cervicornis, D. linearis, D. mertensii, Sargassum platycarpum, Sargassum sp, and Stypopodium zonale were the plants with the highest biomass and/or percentage cover, along with Amphiroa fragilissima, Caulerpa verticillata, and crustose corallines. The scarcity of sea urchins, along with the dominance of algae known to produce chemical feeding deterrents or to have a calcareous nature, points towards a strong pressure by predators, with fishes probably playing a key role in the maintenance of the structure of this subtidal community.

Benthic environment; Community composition; Check lists; Quantitative distribution; Vertical distribution; Oceanic archipelago; Nekton; Fernando de Noronha Archipelago

ARTIGOS

Vertical distribution of benthic marine organisms on rocky coasts of the Fernando de Noronha Archipelago (Brazil)

Verena Rapp de Eston^I,^* (* ) Pós-graduanda da área de oceanografia biológica. ; Alvaro Esteves Migotto^II; Eurico Cabral de Oliveira Filho^II,III; Sergio de Almeida Rodrigues^II,III; Jose Carlos de Freitas^II,III

^IInstituto Oceanográfico da Universidade de São Paulo (Caixa Postal 9075, 01051 São Paulo, SP)

^IICentro de Biologia Marinha da Universidade de São Paulo (Caixa Postal 11461, 05421 São Paulo, SP)

^IIIInstituto de Biociências da Universidade de São Paulo (Caixa Postal 11461, 05421 São Paulo, SP)

SYNOPSIS

Qualitative samples of flora and fauna were collected in several points, from the supralittoral fringe down to 30 m, using mainly aqualungs. Species distribution and abundance were determined along four transects, up to 200 m long, in selected places. Samples were taken from 25 x 25 cm quadrats positioned at fixed intervals along the transects down to a variable depth, depending on algal percentage cover and their substrate. The intertidal zone was populated essentially by molluscs. The subtidal was covered predominantly by macroalgae. Montastrea cavernosa, Siderastrea stellata and Mussismillia hispida were the only animal species with a significant percentage cover within the quadrats. Dictyopteris justii, D. plagiograma, Dictyota cervicornis, D. linearis, D. mertensii, Sargassum platycarpum, Sargassum sp, and Stypopodium zonale were the plants with the highest biomass and/or percentage cover, along with Amphiroa fragilissima, Caulerpa verticillata, and crustose corallines. The scarcity of sea urchins, along with the dominance of algae known to produce chemical feeding deterrents or to have a calcareous nature, points towards a strong pressure by predators, with fishes probably playing a key role in the maintenance of the structure of this subtidal community.

Descriptors: Benthic environment, Community composition; Check lists, Quantitative distribution, Vertical distribution, Oceanic archipelago, Nekton, Fernando de Noronha Archipelago: Brazil.

Descritores: Meio ambiente bentônico, Composição da comunidade, Lista das espécies, Distribuição quantitativa, Distribuição vertical, Arquipélago oceânico, Necton, Arquipélago de Fernando de Noronha: Brasil.

Full text available only in PDF format.

Texto completo disponível apenas em PDF.

Acknowledgements

We are very grateful to Naercio A. Menezes, Gustavo S. de Mello, Beatriz M. de Moraes, Paulo S. Young, Paulo Cesar de Paiva and Edmundo F. Nonato, Maria Helena Baeta Neves, Edison J. de Paula, Y. Yoneshigue, and Luiz R. Tommasi for the identification of the fishes, crabs, sponges, corals and cirriped, polychaetes, bluegreen algae, Sargassum spp, P. boundouresqueri, and Tripneustes ventricosus, respectively. We also thank José Luiz M. Leme for the confirmation of the identification of some species of molluscs, Rosana M. Rocha for help in the identification of the tunicates, and Patricia R. Blauth for the revision of the english text. This work was possible due to a FAPESP grant given to Eurico C. de Oliveira Filho (Proc nº 85/1454-3).

References

ALMEIDA, F. F. M. de 1955. Geologia e petrologia do arquipélago de Fernando de Noronha. Monogr. Dep. nac. Prod, miner. Div. Geol. Miner., Rio de Janeiro, (13):1-181.

BANDEL, K. & KALDOSKY, D. 1982. Western Atlantic species of Nodilittorina (Gastropoda; Prosobranchia): comparative morphology and its functional, ecological, phylogenetic and taxonomic implications. Veliger, 25:1-42.

DAYTON, P. K. 1985. The structure and regulation of some South American kelp communities. Ecol. Monogr., 55(4):447-468.

DICKIE, G. 1874. Enumeration of algae collected at Fernando de Noronha by H. M. Moseley, N.A., naturalist to H.M.S. "Challenger". J. Linn. Soc. (Bot.), 14:363-365.

DUGGINS, D. O. 1983. Starfish predation and the creation of mosaic patterns in a kelp-dominated community. Ecology, 64(6):1610-1619.

DÜING, W.; OSTAPOFF, F. & MERLE, J., eds 1980. Physical oceanography of the tropical Atlantic during GATE. Miami, Fla., University of Miami, 117p.

FAUSTO FILHO, J. 1974. Stomatopod and decapod crustaceans of the archipelago of Fernando de Noronha, northeast Brazil. Arq. Ciênc. Mar, Univ. fed. Ceará, 14(1):1-35,

HAY, M. E. 1981. The functional morphology of turf-forming seaweeds: persistence in stressful marine habitats. Ecology, 62:739-750.

______ 1984. Predictable spatial escapes from herbivory: how do these affect the evolution of herbivore resistance in tropical communities? Oecologia, 64:396-407.

KEMPF, M. & LABOREL, J. L. 1968. Formations de vermets et d'algues calcaires sur les côtes du Brésil. Recl Trav. Stn mar. Endoume, 43(59): 9-23.

LABOREL, J. L. 1969. Les peuplements de madreporaires des côtes tropicales du Brésil. Annls Univ. Abidjan, sér. E, 2(3):1-261.

______ & KEMPF, M. 1965/1966. Formações de vermetos e algas calcárias nas costas do Brasil. Trabhs Inst. oceanogr. Univ. fed. Pernamb., Recife, 7/8:33-50.

LEWIS, J. R. 1964. The ecology of rocky shores. London, The English Universities Press, 323p.

LITTLER, M. M. & LITTLER, D. S. 1980. The evolution of thallus form and survival strategies in benthic marine macroalgae: field and laboratory tests of a functional form model. Am. Nat., 116:25-44.

LOBEL, P. S. 1981. Trophic biology of herbivorous reef fishes: alimentary pH and digestive capabilities. J. Fish Biol., 19:365-397.

______ & OGDEN, J. C. 1981. Foraging by the herbivorous parrotfish Sparisoma radians. Mar. Biol., 64:173-183.

LOPES, H. S. & ALVARENGA, M. 1955. Contribuição ao conhecimento dos moluscos da ilha de Fernando de Noronha, Brasil. Bolm Inst. paul. Oceanogr., S Paulo, 6(1/2):157-190.

LUBBOCK, R. & EDWARDS, A. 1981. The fishes of Saint Paul's Rocks. J. Fish Biol., 18(2):135-157.

MATTHEWS, H. R. & KEMPF, M. 1970. Moluscos marinhos do Norte e Nordeste do Brasil. II. Moluscos do arquipélago de Fernando de Noronha (com algumas reterências ao atol das Rocas). Arq. Ciênc. Mar., Univ. fed. Ceará, 10(1):1-53.

MILLER, R. J. 1985. Succession in sea urchin and sea weed abundance in Nova Scotia, Canada. Mar. Biol., 84 (3):275-286.

MOLINARI, R. L.; VOITURIEZ, B. & DUNCAN, P. 1981. Observations in the subthermocline undercurrent of the Equatorial South Atlantic Ocean:1978-1980. Oceanol. Acta, 4(4):451-456.

NORRIS, J. N. & FENICAL, W. 1982. Chemical defense in tropical marine algae. In: Rützler,& MacIntyre, I. G., eds - The Altantic barrier reef ecosystem at Carrie Bow Cay, Belize. I. Structure and communities. Smithson. Contr. mar. Sci., (12):417-432.

NORTH, W. J. & PEARSE, J. S. 1970. Sea urchin population explosion in southern California coastal waters. Science, N.Y., 167:203.

OLIVEIRA FILHO, E. C. 1974. An annotated list of the Brazilian seaweeds in Dickie's herbarium. J. Linn. Soc, (Bot.), 69(3):229-238.

______ 1977. Algas marinhas bentônicas do Brasil. Tese de livre-docência. Universidade de São Paulo, Instituto de Biociencias. 407p.

PEREIRA, S. M. B.; UGADIM, Y.; PEDRINI, A. G. & BRAGA, M. R. A. Algas marinhas bentônicas do Território de Fernando de Noronha (Brasil). Phycol. lat.-am., 4. (in press).

PHILANDER, S. G. K. 1973. Equatorial Undercurrent: measurements and theories. Rev. Geophys. Space Phys., 11:513-570.

PRICE, J. H. & JOHN, D. M. 1980. Ascencion Island, South Atlantic: a survey of inshore benthic macroorganisms, communities and interactions. Aquat. Bot., 9(3): 251-278.

RANDALL, J. E, 1967. Food habits of reef fishes of the West Indies. In: International Conference on Tropical Oceanography, 1965, Miami Beach, Florida. Proceedings, p.665-847.

RIDLEY, H. N. 1890. Notes on the zoology of Fernando de Noronha. J. Linn. Soc. (Zool.), 20:473-592.

SAZIMA, I. & SAZIMA, M. 1983. Aspectos de comportamento alimentar e dieta da tartaruga marinha, Chelonia mydas, no litoral norte paulista. Bolm Inst. oceanogr., S Paulo, 32(2): 199-203.

SCHELTEMA, R. S. 1968. Dispersal of larvae by equatorial ocean currents and its importance to zoogeography of shoal-water tropical species. Nature, Lond., 217:1159-1162.

STENECK, R. S. & WATLING, L. 1982. Feeding capabilities and limitations of herbivorous molluscs: a functional group approach. Mar. Biol., 68:299-319.

VERMEIJ, G. J. 1972. Endemism and environment: some shore molluscs of the tropical Atlantic. Am. Nat., 106(947):89-101.

______ 1973. Morphological patterns in high intertidal gastropods: adaptive strategies and their limitations. Mar. Biol., 20:319-346.

______ & PORTER, J. W. 1971. Some characteristics of the dominant intertidal molluscs from rocky shores in Pernambuco, Brazil. Bull, mar. Sci., 21(2):440-454.

WALTER, H. & LEITH, H. 1960. Klimadiagramm weltatlas. Jena, Gustav Fisher Verlag.

(Received 08-Oct-86; accepted 19-Dec-86)

ALMEIDA, F. F. M. de 1955. Geologia e petrologia do arquipélago de Fernando de Noronha. Monogr. Dep. nac. Prod, miner. Div. Geol. Miner., Rio de Janeiro, (13):1-181.
BANDEL, K. & KALDOSKY, D. 1982. Western Atlantic species of Nodilittorina (Gastropoda; Prosobranchia): comparative morphology and its functional, ecological, phylogenetic and taxonomic implications. Veliger, 25:1-42.
DAYTON, P. K. 1985. The structure and regulation of some South American kelp communities. Ecol. Monogr., 55(4):447-468.
DICKIE, G. 1874. Enumeration of algae collected at Fernando de Noronha by H. M. Moseley, N.A., naturalist to H.M.S. "Challenger". J. Linn. Soc. (Bot.), 14:363-365.
DUGGINS, D. O. 1983. Starfish predation and the creation of mosaic patterns in a kelp-dominated community. Ecology, 64(6):1610-1619.
DÜING, W.; OSTAPOFF, F. & MERLE, J., eds 1980. Physical oceanography of the tropical Atlantic during GATE. Miami, Fla., University of Miami, 117p.
FAUSTO FILHO, J. 1974. Stomatopod and decapod crustaceans of the archipelago of Fernando de Noronha, northeast Brazil. Arq. Ciênc. Mar, Univ. fed. Ceará, 14(1):1-35,
HAY, M. E. 1981. The functional morphology of turf-forming seaweeds: persistence in stressful marine habitats. Ecology, 62:739-750.
______ 1984. Predictable spatial escapes from herbivory: how do these affect the evolution of herbivore resistance in tropical communities? Oecologia, 64:396-407.
KEMPF, M. & LABOREL, J. L. 1968. Formations de vermets et d'algues calcaires sur les côtes du Brésil. Recl Trav. Stn mar. Endoume, 43(59): 9-23.
LABOREL, J. L. 1969. Les peuplements de madreporaires des côtes tropicales du Brésil. Annls Univ. Abidjan, sér. E, 2(3):1-261.
______ & KEMPF, M. 1965/1966. Formações de vermetos e algas calcárias nas costas do Brasil. Trabhs Inst. oceanogr. Univ. fed. Pernamb., Recife, 7/8:33-50.
LEWIS, J. R. 1964. The ecology of rocky shores. London, The English Universities Press, 323p.
LITTLER, M. M. & LITTLER, D. S. 1980. The evolution of thallus form and survival strategies in benthic marine macroalgae: field and laboratory tests of a functional form model. Am. Nat., 116:25-44.
LOBEL, P. S. 1981. Trophic biology of herbivorous reef fishes: alimentary pH and digestive capabilities. J. Fish Biol., 19:365-397.
______ & OGDEN, J. C. 1981. Foraging by the herbivorous parrotfish Sparisoma radians Mar. Biol., 64:173-183.
LOPES, H. S. & ALVARENGA, M. 1955. Contribuição ao conhecimento dos moluscos da ilha de Fernando de Noronha, Brasil. Bolm Inst. paul. Oceanogr., S Paulo, 6(1/2):157-190.
LUBBOCK, R. & EDWARDS, A. 1981. The fishes of Saint Paul's Rocks. J. Fish Biol., 18(2):135-157.
MATTHEWS, H. R. & KEMPF, M. 1970. Moluscos marinhos do Norte e Nordeste do Brasil. II. Moluscos do arquipélago de Fernando de Noronha (com algumas reterências ao atol das Rocas). Arq. Ciênc. Mar., Univ. fed. Ceará, 10(1):1-53.
MILLER, R. J. 1985. Succession in sea urchin and sea weed abundance in Nova Scotia, Canada. Mar. Biol., 84 (3):275-286.
MOLINARI, R. L.; VOITURIEZ, B. & DUNCAN, P. 1981. Observations in the subthermocline undercurrent of the Equatorial South Atlantic Ocean:1978-1980. Oceanol. Acta, 4(4):451-456.
NORRIS, J. N. & FENICAL, W. 1982. Chemical defense in tropical marine algae. In: Rützler,& MacIntyre, I. G., eds - The Altantic barrier reef ecosystem at Carrie Bow Cay, Belize. I. Structure and communities. Smithson.
Contr. mar. Sci., (12):417-432.
NORTH, W. J. & PEARSE, J. S. 1970. Sea urchin population explosion in southern California coastal waters. Science, N.Y., 167:203.
OLIVEIRA FILHO, E. C. 1974. An annotated list of the Brazilian seaweeds in Dickie's herbarium. J. Linn. Soc, (Bot.), 69(3):229-238.
______ 1977. Algas marinhas bentônicas do Brasil. Tese de livre-docência. Universidade de São Paulo, Instituto de Biociencias. 407p.
PEREIRA, S. M. B.; UGADIM, Y.; PEDRINI, A. G. & BRAGA, M. R. A. Algas marinhas bentônicas do Território de Fernando de Noronha (Brasil). Phycol. lat.-am., 4 (in press).
PHILANDER, S. G. K. 1973. Equatorial Undercurrent: measurements and theories. Rev. Geophys. Space Phys., 11:513-570.
PRICE, J. H. & JOHN, D. M. 1980. Ascencion Island, South Atlantic: a survey of inshore benthic macroorganisms, communities and interactions. Aquat. Bot., 9(3): 251-278.
RANDALL, J. E, 1967. Food habits of reef fishes of the West Indies. In: International Conference on Tropical Oceanography, 1965, Miami Beach, Florida. Proceedings, p.665-847.
RIDLEY, H. N. 1890. Notes on the zoology of Fernando de Noronha. J. Linn. Soc. (Zool.), 20:473-592.
SAZIMA, I. & SAZIMA, M. 1983. Aspectos de comportamento alimentar e dieta da tartaruga marinha, Chelonia mydas, no litoral norte paulista. Bolm Inst. oceanogr., S Paulo, 32(2): 199-203.
SCHELTEMA, R. S. 1968. Dispersal of larvae by equatorial ocean currents and its importance to zoogeography of shoal-water tropical species. Nature, Lond., 217:1159-1162.
STENECK, R. S. & WATLING, L. 1982. Feeding capabilities and limitations of herbivorous molluscs: a functional group approach. Mar. Biol., 68:299-319.
VERMEIJ, G. J. 1972. Endemism and environment: some shore molluscs of the tropical Atlantic. Am. Nat., 106(947):89-101.
______ 1973. Morphological patterns in high intertidal gastropods: adaptive strategies and their limitations. Mar. Biol., 20:319-346.
______ & PORTER, J. W. 1971. Some characteristics of the dominant intertidal molluscs from rocky shores in Pernambuco, Brazil. Bull, mar. Sci., 21(2):440-454.
WALTER, H. & LEITH, H. 1960. Klimadiagramm weltatlas. Jena, Gustav Fisher Verlag.

(*

) Pós-graduanda da área de oceanografia biológica.

Publication Dates

Publication in this collection
01 June 2012
Date of issue
1986

History

Accepted
19 Dec 1986
Received
08 Oct 1986

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

[1] ALMEIDA, F. F. M. de 1955. Geologia e petrologia do arquipélago de Fernando de Noronha. Monogr. Dep. nac. Prod, miner. Div. Geol. Miner., Rio de Janeiro, (13):1-181.

[2] BANDEL, K. & KALDOSKY, D. 1982. Western Atlantic species of Nodilittorina (Gastropoda; Prosobranchia): comparative morphology and its functional, ecological, phylogenetic and taxonomic implications. Veliger, 25:1-42.

[3] DAYTON, P. K. 1985. The structure and regulation of some South American kelp communities. Ecol. Monogr., 55(4):447-468.

[4] DICKIE, G. 1874. Enumeration of algae collected at Fernando de Noronha by H. M. Moseley, N.A., naturalist to H.M.S. "Challenger". J. Linn. Soc. (Bot.), 14:363-365.

[5] DUGGINS, D. O. 1983. Starfish predation and the creation of mosaic patterns in a kelp-dominated community. Ecology, 64(6):1610-1619.

[6] DÜING, W.; OSTAPOFF, F. & MERLE, J., eds 1980. Physical oceanography of the tropical Atlantic during GATE. Miami, Fla., University of Miami, 117p.

[7] FAUSTO FILHO, J. 1974. Stomatopod and decapod crustaceans of the archipelago of Fernando de Noronha, northeast Brazil. Arq. Ciênc. Mar, Univ. fed. Ceará, 14(1):1-35,

[8] HAY, M. E. 1981. The functional morphology of turf-forming seaweeds: persistence in stressful marine habitats. Ecology, 62:739-750.

[9] ______ 1984. Predictable spatial escapes from herbivory: how do these affect the evolution of herbivore resistance in tropical communities? Oecologia, 64:396-407.

[10] KEMPF, M. & LABOREL, J. L. 1968. Formations de vermets et d'algues calcaires sur les côtes du Brésil. Recl Trav. Stn mar. Endoume, 43(59): 9-23.

[11] LABOREL, J. L. 1969. Les peuplements de madreporaires des côtes tropicales du Brésil. Annls Univ. Abidjan, sér. E, 2(3):1-261.

[12] ______ & KEMPF, M. 1965/1966. Formações de vermetos e algas calcárias nas costas do Brasil. Trabhs Inst. oceanogr. Univ. fed. Pernamb., Recife, 7/8:33-50.

[13] LEWIS, J. R. 1964. The ecology of rocky shores. London, The English Universities Press, 323p.

[14] LITTLER, M. M. & LITTLER, D. S. 1980. The evolution of thallus form and survival strategies in benthic marine macroalgae: field and laboratory tests of a functional form model. Am. Nat., 116:25-44.

[15] LOBEL, P. S. 1981. Trophic biology of herbivorous reef fishes: alimentary pH and digestive capabilities. J. Fish Biol., 19:365-397.

[16] ______ & OGDEN, J. C. 1981. Foraging by the herbivorous parrotfish Sparisoma radians Mar. Biol., 64:173-183.

[17] LOPES, H. S. & ALVARENGA, M. 1955. Contribuição ao conhecimento dos moluscos da ilha de Fernando de Noronha, Brasil. Bolm Inst. paul. Oceanogr., S Paulo, 6(1/2):157-190.

[18] LUBBOCK, R. & EDWARDS, A. 1981. The fishes of Saint Paul's Rocks. J. Fish Biol., 18(2):135-157.

[19] MATTHEWS, H. R. & KEMPF, M. 1970. Moluscos marinhos do Norte e Nordeste do Brasil. II. Moluscos do arquipélago de Fernando de Noronha (com algumas reterências ao atol das Rocas). Arq. Ciênc. Mar., Univ. fed. Ceará, 10(1):1-53.

[20] MILLER, R. J. 1985. Succession in sea urchin and sea weed abundance in Nova Scotia, Canada. Mar. Biol., 84 (3):275-286.

[21] MOLINARI, R. L.; VOITURIEZ, B. & DUNCAN, P. 1981. Observations in the subthermocline undercurrent of the Equatorial South Atlantic Ocean:1978-1980. Oceanol. Acta, 4(4):451-456.

[22] NORRIS, J. N. & FENICAL, W. 1982. Chemical defense in tropical marine algae. In: Rützler,& MacIntyre, I. G., eds - The Altantic barrier reef ecosystem at Carrie Bow Cay, Belize. I. Structure and communities. Smithson.

[23] Contr. mar. Sci., (12):417-432.

[24] NORTH, W. J. & PEARSE, J. S. 1970. Sea urchin population explosion in southern California coastal waters. Science, N.Y., 167:203.

[25] OLIVEIRA FILHO, E. C. 1974. An annotated list of the Brazilian seaweeds in Dickie's herbarium. J. Linn. Soc, (Bot.), 69(3):229-238.

[26] ______ 1977. Algas marinhas bentônicas do Brasil. Tese de livre-docência. Universidade de São Paulo, Instituto de Biociencias. 407p.

[27] PEREIRA, S. M. B.; UGADIM, Y.; PEDRINI, A. G. & BRAGA, M. R. A. Algas marinhas bentônicas do Território de Fernando de Noronha (Brasil). Phycol. lat.-am., 4 (in press).

[28] PHILANDER, S. G. K. 1973. Equatorial Undercurrent: measurements and theories. Rev. Geophys. Space Phys., 11:513-570.

[29] PRICE, J. H. & JOHN, D. M. 1980. Ascencion Island, South Atlantic: a survey of inshore benthic macroorganisms, communities and interactions. Aquat. Bot., 9(3): 251-278.

[30] RANDALL, J. E, 1967. Food habits of reef fishes of the West Indies. In: International Conference on Tropical Oceanography, 1965, Miami Beach, Florida. Proceedings, p.665-847.

[31] RIDLEY, H. N. 1890. Notes on the zoology of Fernando de Noronha. J. Linn. Soc. (Zool.), 20:473-592.

[32] SAZIMA, I. & SAZIMA, M. 1983. Aspectos de comportamento alimentar e dieta da tartaruga marinha, Chelonia mydas, no litoral norte paulista. Bolm Inst. oceanogr., S Paulo, 32(2): 199-203.

[33] SCHELTEMA, R. S. 1968. Dispersal of larvae by equatorial ocean currents and its importance to zoogeography of shoal-water tropical species. Nature, Lond., 217:1159-1162.

[34] STENECK, R. S. & WATLING, L. 1982. Feeding capabilities and limitations of herbivorous molluscs: a functional group approach. Mar. Biol., 68:299-319.

[35] VERMEIJ, G. J. 1972. Endemism and environment: some shore molluscs of the tropical Atlantic. Am. Nat., 106(947):89-101.

[36] ______ 1973. Morphological patterns in high intertidal gastropods: adaptive strategies and their limitations. Mar. Biol., 20:319-346.

[37] ______ & PORTER, J. W. 1971. Some characteristics of the dominant intertidal molluscs from rocky shores in Pernambuco, Brazil. Bull, mar. Sci., 21(2):440-454.

[38] WALTER, H. & LEITH, H. 1960. Klimadiagramm weltatlas. Jena, Gustav Fisher Verlag.