Comparative analysis of shell occupation by two southern populations of the hermit crab Loxopagurus loxochelis (Decapoda, Diogenidae)

Ayres-Peres, Luciane; Quadros, Aline Ferreira; Mantelatto, Fernando L.

Abstracts

The present study aimed to comparatively verify the relation between the hermit crabs and the shells they use in two populations of Loxopagurus loxochelis. Samples were collected monthly from July 2002 to June 2003, at Caraguatatuba and Ubatuba Bay, São Paulo, Brazil. The animals sampled had their sex identified, were weighed and measured; their shells were identified, measured and weighed, and their internal volume determined. To relate the hermit crab's characteristics and the shells' variables, principal component analysis (PCA) and a regression tree were used. According to the PCA analysis, the three gastropod shells most frequently used by L. loxochelis varied in size. The regression tree successfully explained the relationship between the hermit crab's characteristics and the internal volume of the inhabited shell. It can be inferred that the relationship between the morphometry of an individual hermit crab and its shell is not straightforward and it is impossible to explain only on the basis of direct correlations between the body's and the shell's attributes. Several factors (such as the morphometry and the availability of the shell, environmental conditions and inter- and intraspecific competition) interact and seem to be taken into consideration by the hermit crabs when they choose a shell, resulting in the diversified pattern of shell occupancy shown here and elsewhere.

Diogenidae; shell utilization; PCA; regression tree; Atlantic

O presente estudo teve como objetivo verificar de forma comparativa a relação entre ermitões e conchas utilizadas por duas populações de Loxopagurus loxochelis. Foram realizadas coletas mensais, de julho 2002 a junho 2003, nas Enseadas de Caraguatatuba e de Ubatuba, São Paulo, Brasil. Os animais amostrados foram identificados quanto ao sexo, pesados, e mensurados; as conchas de gastrópodes foram identificadas, mensuradas, pesadas e tiveram seu volume interno determinado. Para relacionar as variáveis dos ermitões com as conchas foram utilizadas a análise de componentes principais (PCA) e a árvore de regressão. As três conchas de gastrópodes utilizadas mais intensamente diferiram em tamanho de acordo com a PCA. A árvore de regressão explicou com sucesso a relação entre características do ermitão e volume interno das conchas habitadas. Inferiu-se que a relação entre a morfometria de um ermitão e sua concha não é simples e não é possível de ser explicada com base somente em correlações diretas entre atributos do corpo e da concha. Inúmeros fatores (como morfometria e disponibilidade da concha, condições ambientais e competição inter e intra-específica) interagem e devem ser levados em consideração na escolha de uma concha pelos ermitões, resultando no diversificado padrão de ocupação de conchas demonstrado neste e em outros trabalhos correlatos.

Diogenidae; utilização de conchas; PCA; árvore de regressão; Atlântico

AYRES-PERES, L.; MANTELATTO, F. L. Análise comparativa da estrutura populacional do ermitão endêmico do Atlântico Ocidental Loxopagurus loxochelis (Decapoda, Anomura) em duas regiões do Estado de São Paulo, Brasil. Iheringia, Sér. Zool., v. 98, n. 1, p. 28-35, 2008.
AYRES-PERES, L.; MANTELATTO, F. L. Patterns of distribution of the hermit crab Loxopagurus loxochelis (Moreira, 1901) (Decapoda, Diogenidae) in two coastal areas of southern Brazil. Rev. Biol. Mar. Oceanogr., v. 43, n. 2, p. 399-411, 2008a.
AYRES-PERES, L.; MANTELATTO, F. L. Epibiont occurrence on gastropod shells used by the hermit crab Loxopagurus loxochelis (Anomura: Diogenidae) on the northern coast of São Paulo, Brazil. Zoologia, v. 27, n. 2, p. 222–227, 2010.
AYRES-PERES, L.; SOKOLOWICZ, C. C.; KOTZIAN, C. B.; RIEGER, P. J.; SANTOS, S. Ocupação de conchas de gastrópodes por ermitões (Decapoda, Anomura) no litoral de Rio Grande, Rio Grande do Sul, Brasil. Iheringia, Sér. Zool., v. 98, n. 2, p. 218-24, 2008.
BARNES, D. K. A. Ecology of tropical hermit crabs at Quirimba Island, Mozambique: shell characteristics and utilization. Mar. Ecol. Prog. Ser., v. 183, p. 241-251, 1999.
BERTINI, G.; FRANSOZO, A. Patterns of shell utilization in Petrochirus diogenes (Decapoda, Anomura, Diogenidae) in the Ubatuba region, São Paulo, Brazil. J. Crust. Biol., v. 20, n. 3, p. 468-473, 2000.
BERTNESS, M. D. The influence of shell-type on hermit crab growth rate and clutch size. Crustaceana, v. 40, n. 2, p. 197-205, 1981.
BERTNESS, M. D. Shell utilization, predation pressure, and thermal stress in Panamanian hermit crabs: an interoceanic comparison. J. exp. mar. Biol. Ecol., v. 64, n. 2, p. 159-187, 1982.
BIAGI, R.; MEIRELES, A. L.; SCELZO, M. A.; MANTELATTO, F. L. Comparative study of shell choice by the southern endemic hermit crab Loxopagurus loxochelis from Brazil and Argentina. Rev. Chil. Hist. Nat., v. 79, n. 4, p. 481-487, 2006.
BOLLAY, M. Distribution and utilization of gastropod shells by the hermit crabs Pagurus samuelis, Pagurus granosimanus and Pagurus hirsutiusculus at Pacific Grove, California. Veliger, v. 6 (suppl.), p. 71-76, 1964.
BOTELHO, Z. B.; COSTA, A. C. Shell occupancy of the intertidal hermit crab Clibanarius erythropus (Decapoda, Diogenidae) on São Miguel (Azores). Hydrobiologia, v. 440, p. 111-117, 2000.
CARRANZA, A.; SEGURA, A.; LOPEZ, J.; RUBIO, L. Shell use patterns of the hermit crab Loxopagurus loxochelis (Decapoda: Diogenidae) in Cerro Verde - La Coronilla (Rocha, Uruguay). Comun. Soc. Malacol. Urug., v. 9, n. 91, p. 139-145, 2008.
CHILDRESS, J. R. Behavioral ecology and fitness theory in a tropical hermit crab. Ecology, v. 53, n. 3, p. 960-964, 1972.
CONOVER, M. R. The importance of various shell characteristics to the shell - selection behavior on the hermit crabs. J. exp. mar. Biol. Ecol., v. 32, n. 2, p. 131-142, 1978.
DEATH, G. Multivariate regression trees: a new technique for modelling species-environment relationships. Ecology, v. 83, n. 4, p. 1105-1117, 2002.
DEATH, G.; FABRICIUS, K. E. Classification and regression trees: A powerful yet simple technique for ecological data analysis. Ecology, v. 81, n. 11, p. 3178-3192, 2000.
DOMINCIANO, L. C. C.; SANT'ANNA, B. S.; TURRA, A. Are the preference and selection patterns of hermit crabs for gastropod shells species or site-specific? J. exp. Mar. Biol. Ecol., v. 378, n. 1, p. 15-21, 2009.
ELWOOD, R. W.; MARKS, N.; DICK, J. T. A. Consequences of shell - species preferences for female reproductive success in the hermit crab Pagurus bernhardus Mar. Biol., v. 123, n. 3, p. 431-434, 1995.
FANTUCCI, M. Z.; BIAGI, R.; MANTELATTO, F. L. Shell occupation by the endemic western Atlantic hermit crab Isocheles sawayai (Diogenidae) from Caraguatatuba, Brazil. Braz. J. Biol, v. 68, n. 4, p. 859-867, 2008.
FOTHERINGHAM, N. Hermit crabs as a limiting resource (Decapoda, Paguridae). Crustaceana, v. 31, n. 2, p. 193-199, 1976.
FOTHERINGHAM, N. Effects of shell stress on the growth of hermit crabs. J. exp. mar. Biol. Ecol., v. 23, n. 3, p. 299-305, 1976a.
GARCIA, R. B.; MANTELATTO, F. L. Variability of shell occupation by intertidal and infralittoral Calcinus tibicen (Anomura, Diogenidae) populations. Nauplius, v. 8, n. 1, p. 99-105, 2000.
GARCIA, R. B.; MANTELATTO, F. L. Shell selection by the tropical hermit crab Calcinus tibicen (Herbst, 1791) (Anomura, Diogenidae) from Southern Brazil. J. exp. mar. Biol. Ecol., v. 265, n. 1, p. 1-14, 2001.
GHERARDI, F. Relative growth, population structure, and shell utilization of the hermit crab Clibanarius erythropus in the Mediterranean. Oebalia, v. 17, p. 181 – 196, 1991.
GRANT, W. C.; ULMER, K. M. Shell selection and aggressive behavior in two sympatric species of hermit crabs. Biol. Bull., v. 146, n. 1, p. 32-43, 1974.
HAMMER, Ø.; HARPER, D. A. T.; RYAN, P. D. PAST: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontologia Electronica 4(1):9pp. http://palaeo-electronica.org/2001_1/past/issue1_01.htm, 2001.
HAZLETT, B. A. The behavioral ecology of hermit crabs. Ann. Rev. Ecol. Syst., v. 12, n. 1, p. 1-22, 1981.
KELLOGG, C. W. Gastropod shells: a potentially limiting resource for hermit crabs. J. exp. mar. Biol. Ecol., v. 22, n. 1, p. 101-111, 1976.
LEITE, F. P. P.; TURRA, A.; GANDOLFI, S. M. Hermit crabs (Crustacea: Decapoda: Anomura), gastropod shells and environment structure: their relationship in southeastern Brazil. J. Nat. Hist., v. 32, n. 10-11, p. 1599-1608, 1998.
MANTELATTO, F. L.; GARCIA, R. B. Shell utilization pattern of the hermit crab Calcinus tibicen (Diogenidae) from Southern Brazil. J. Crust. Biol., v. 20, n. 3, p. 460-467, 2000.
MANTELATTO, F. L.; MEIRELES, A. L. The importance of shell occupation and shell availability in the hermit crab Pagurus brevidactylus (Stimpson, 1859) (Paguridae) population from the southern Atlantic. Bull. mar. Sci., v. 75, n. 1, p. 27-35, 2004.
MANTELATTO, F. L.; ALARCON, V. F.; GARCIA, R. B. Egg production strategies of the tropical hermit crab Paguristes tortugae from Brazil. J. Crust. Biol., v. 22, n. 2, p. 390-397, 2002.
MANTELATTO, F. L.; BIAGI, R.; MEIRELES, A. L.; SCELZO, M. A. Shell preference of the hermit crab Pagurus exilis (Anomura: Paguridae) from Brazil and Argentina: a comparative study. Rev. Biol. Trop., v. 55, p. 153-161, 2007.
MANTELATTO, F. L.; FERNANDES-GÓES, L. C.; FANTUCCI, M. Z.; BIAGI, R.; PARDO, L. M.; GÓES, J. M. A comparative study of population traits between two South American populations of the striped-legged hermit crab Clibanarius vittatus Acta Oecol., v. 36, n. 1, p. 10-15, 2010.
MARKHAM, J. Notes on the growth patterns and shell utilization of the hermit crab Pagurus bernhardus (L.). Ophelia, v. 5, p. 189-205, 1968.
MARTINELLI, J. M.; MANTELATTO, F. L. Occurrence of exobionts in gastropod shells occupied by the hermit crab Loxopagurus loxochelis (Anomura: Diogenidae) in Ubatuba Bay (SP) Brazil. In: Anais do IV Simpósio de Ecossistemas Brasileiros, v. 2, n. 104. Publicações ACIESP, São Paulo, p. 221-226, 1998.
MARTINELLI, J. M.; MANTELATTO, F. L. Shell utilization by the hermit crab Loxopagurus loxochelis (Diogenidae) in Ubatuba Bay, Brazil. In: SCHRAM, F. R. & VANPEL KLEIN, J. R. (eds.). Crust. Biodiv. Crisis, v. 1, p. 719-731, 1999.
MEIRELES, A. L.; MANTELATTO, F. L. Shell use by the Pagurus brevidactylus (Anomura, Paguridae): a comparison between laboratory and field conditions. Acta Zool. Sin., v. 51, n. 5, p. 813-820, 2005.
MEIRELES, A. L.; TEROSSI, M.; BIAGI, R.; MANTELATTO, F. L. Spatial and seasonal distribution of the hermit crab Pagurus exilis (Benedict, 1892) (Decapoda: Paguridae) in the southwestern coast of Brazil. Rev. Biol. Mar. Oceanogr., v. 41, n. 1, p. 87-95, 2006.
MELO, G. A. S. Manual de identificação dos Crustacea Decapoda do litoral brasileiro: Anomura, Thalassinidea, Palinuridea, Astacidea. Plêiade, São Paulo, 551 p, 1999.
MITCHELL, K. A. Shell selection in the hermit crab Pagurus bernhardus. Mar. Biol, v. 35, n. 4, p. 335-346, 1976.
OHMORI, H.; WADA, S.; GOSHIMA, S.; NAKAO, S. Effect of body size and shell avaibality on the shell utilization pattern of the hermit crab Pagurus filholi (Anomura: Paguridae). Crust. Res., v. 24, p. 85-92, 1995.
PESSANI, D.; PREMOLI, C. Some aspects of the biology of Cestopagurus timidus (Crustacea, Paguridae) in relation to the occupied shell. Boll. Zool., v. 60, n. 2, p. 199-205, 1993.
PRASAD, A. M.; IVERSON, L. R.; LIAW, A. Newer classification and regression tree techniques: Bagging and random forests for ecological prediction. Ecosystems, v. 9, p. 181-199, 2006.
PIRES-VANIN, A. M. S.; ROSSI-WONGTSCHOWSKI, C. L.; AIDAAR, E.; MESQUITA, H.; SOARES, L. S. H.; KATSURAGAWA, M.; MATSUURA, Y. Estrutura e função do ecossistema de plataforma continental do Atlântico Sul brasileiro: síntese dos resultados. Publ. esp. Inst. Oceanogr., v. 10, p. 217-231, 1993.
PROVENZANO Jr., A. J. Notes on Bermuda hermit crabs (Crustacea: Anomura). Bull. Mar. Sci. Gulf Caribb., v. 10, p. 117-124, 1960.
R Development Core Team. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. ISBN 3-900051-07-0, URL http://www.R-project.org, 2010.
REESE, E. Behavioral adaptation of intertidal hermit crabs. Am. Zool., v. 9, n. 2, p. 343-355, 1969.
RIOS, E. C. Seashells of Brazil. Rio Grande do Sul. Fundação Universidade de Rio Grande, Instituto Acqua, Museu Oceanográfico de Rio Grande, Universidade de Rio Grande. 2ed. 368 p. + 113pl., 1994.
SATO, Y.; SENO, H. A mathematical consideration for the optimal shell change of hermit crab. J. Theor. Biol., v. 240, p. 14-23, 2006.
SCULLY, E. P. The effects of gastropod shell availability and habitat characteristics on shell utilization by the intertidal hermit crab Pagurus longicarpus Say. J. exp. mar. Biol. Ecol., v. 37, n. 2, p. 139-152, 1979.
SPIGHT, T. M. Availability and use of shells by intertidal hermit crabs. Biol. Bull., v. 152, n. 1, p. 120-133, 1977.
SUZUKI, R.; SHIMODAIRA, H. PVCLUST v 1.2-0. R package version 3.1-34. URL http://www.R-project.org, 2006.
TEROSSI, M.; ESPÓSITO, D. L. A.; MEIRELES, A. L.; BIAGI, R.; MANTELATTO, F. L. Pattern of shell occupation by the hermit crab Pagurus exilis (Anomura, Paguridae) in southern Atlantic region. J. Nat. Hist., v. 40, n. 1-2, p. 77-87, 2006.
TORATI, L. S.; MANTELATTO F. L. Uncommon mechanism of egg incubation in the endemic Southern hermit crab Loxopagurus loxochelis: how is this phenomenon related to egg production? Acta Zool., v. 89, n. 1, p.79-85, 2008.
THERNEAU, T. M.; ATKINSON, B. RPART: Recursive Partitioning. Report by B. Ripley. R package version 3.1-34. URL http://www.R-project.org, 2010.
TUNBERG, B. G.; NELSON, W. G.; SMITH, G. Population ecology of Pagurus maclaughlinae Garcia-Gomes (Decapoda: Anomura: Paguridae) in the Indian River Lagoon, Florida. J. Crust. Biol., v. 14, n. 4, p. 686-699, 1994.
VANCE, R. R. The role of shell adequacy in behavior interactions in hermit crabs. Ecology, v. 53, n. 6, p. 1075-1083, 1972.
VANCE, R. R. Competition and mechanisms of coexistence in three sympatric species of intertidal hermit crabs. Ecology, v. 53, n. 6, p.1062-1074, 1972a.

Publication Dates

Publication in this collection
30 Oct 2012
Date of issue
Sept 2012

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

[1] AYRES-PERES, L.; MANTELATTO, F. L. Análise comparativa da estrutura populacional do ermitão endêmico do Atlântico Ocidental Loxopagurus loxochelis (Decapoda, Anomura) em duas regiões do Estado de São Paulo, Brasil. Iheringia, Sér. Zool., v. 98, n. 1, p. 28-35, 2008.

[2] AYRES-PERES, L.; MANTELATTO, F. L. Patterns of distribution of the hermit crab Loxopagurus loxochelis (Moreira, 1901) (Decapoda, Diogenidae) in two coastal areas of southern Brazil. Rev. Biol. Mar. Oceanogr., v. 43, n. 2, p. 399-411, 2008a.

[3] AYRES-PERES, L.; MANTELATTO, F. L. Epibiont occurrence on gastropod shells used by the hermit crab Loxopagurus loxochelis (Anomura: Diogenidae) on the northern coast of São Paulo, Brazil. Zoologia, v. 27, n. 2, p. 222–227, 2010.

[4] AYRES-PERES, L.; SOKOLOWICZ, C. C.; KOTZIAN, C. B.; RIEGER, P. J.; SANTOS, S. Ocupação de conchas de gastrópodes por ermitões (Decapoda, Anomura) no litoral de Rio Grande, Rio Grande do Sul, Brasil. Iheringia, Sér. Zool., v. 98, n. 2, p. 218-24, 2008.

[5] BARNES, D. K. A. Ecology of tropical hermit crabs at Quirimba Island, Mozambique: shell characteristics and utilization. Mar. Ecol. Prog. Ser., v. 183, p. 241-251, 1999.

[6] BERTINI, G.; FRANSOZO, A. Patterns of shell utilization in Petrochirus diogenes (Decapoda, Anomura, Diogenidae) in the Ubatuba region, São Paulo, Brazil. J. Crust. Biol., v. 20, n. 3, p. 468-473, 2000.

[7] BERTNESS, M. D. The influence of shell-type on hermit crab growth rate and clutch size. Crustaceana, v. 40, n. 2, p. 197-205, 1981.

[8] BERTNESS, M. D. Shell utilization, predation pressure, and thermal stress in Panamanian hermit crabs: an interoceanic comparison. J. exp. mar. Biol. Ecol., v. 64, n. 2, p. 159-187, 1982.

[9] BIAGI, R.; MEIRELES, A. L.; SCELZO, M. A.; MANTELATTO, F. L. Comparative study of shell choice by the southern endemic hermit crab Loxopagurus loxochelis from Brazil and Argentina. Rev. Chil. Hist. Nat., v. 79, n. 4, p. 481-487, 2006.

[10] BOLLAY, M. Distribution and utilization of gastropod shells by the hermit crabs Pagurus samuelis, Pagurus granosimanus and Pagurus hirsutiusculus at Pacific Grove, California. Veliger, v. 6 (suppl.), p. 71-76, 1964.

[11] BOTELHO, Z. B.; COSTA, A. C. Shell occupancy of the intertidal hermit crab Clibanarius erythropus (Decapoda, Diogenidae) on São Miguel (Azores). Hydrobiologia, v. 440, p. 111-117, 2000.

[12] CARRANZA, A.; SEGURA, A.; LOPEZ, J.; RUBIO, L. Shell use patterns of the hermit crab Loxopagurus loxochelis (Decapoda: Diogenidae) in Cerro Verde - La Coronilla (Rocha, Uruguay). Comun. Soc. Malacol. Urug., v. 9, n. 91, p. 139-145, 2008.

[13] CHILDRESS, J. R. Behavioral ecology and fitness theory in a tropical hermit crab. Ecology, v. 53, n. 3, p. 960-964, 1972.

[14] CONOVER, M. R. The importance of various shell characteristics to the shell - selection behavior on the hermit crabs. J. exp. mar. Biol. Ecol., v. 32, n. 2, p. 131-142, 1978.

[15] DEATH, G. Multivariate regression trees: a new technique for modelling species-environment relationships. Ecology, v. 83, n. 4, p. 1105-1117, 2002.

[16] DEATH, G.; FABRICIUS, K. E. Classification and regression trees: A powerful yet simple technique for ecological data analysis. Ecology, v. 81, n. 11, p. 3178-3192, 2000.

[17] DOMINCIANO, L. C. C.; SANT'ANNA, B. S.; TURRA, A. Are the preference and selection patterns of hermit crabs for gastropod shells species or site-specific? J. exp. Mar. Biol. Ecol., v. 378, n. 1, p. 15-21, 2009.

[18] ELWOOD, R. W.; MARKS, N.; DICK, J. T. A. Consequences of shell - species preferences for female reproductive success in the hermit crab Pagurus bernhardus Mar. Biol., v. 123, n. 3, p. 431-434, 1995.

[19] FANTUCCI, M. Z.; BIAGI, R.; MANTELATTO, F. L. Shell occupation by the endemic western Atlantic hermit crab Isocheles sawayai (Diogenidae) from Caraguatatuba, Brazil. Braz. J. Biol, v. 68, n. 4, p. 859-867, 2008.

[20] FOTHERINGHAM, N. Hermit crabs as a limiting resource (Decapoda, Paguridae). Crustaceana, v. 31, n. 2, p. 193-199, 1976.

[21] FOTHERINGHAM, N. Effects of shell stress on the growth of hermit crabs. J. exp. mar. Biol. Ecol., v. 23, n. 3, p. 299-305, 1976a.

[22] GARCIA, R. B.; MANTELATTO, F. L. Variability of shell occupation by intertidal and infralittoral Calcinus tibicen (Anomura, Diogenidae) populations. Nauplius, v. 8, n. 1, p. 99-105, 2000.

[23] GARCIA, R. B.; MANTELATTO, F. L. Shell selection by the tropical hermit crab Calcinus tibicen (Herbst, 1791) (Anomura, Diogenidae) from Southern Brazil. J. exp. mar. Biol. Ecol., v. 265, n. 1, p. 1-14, 2001.

[24] GHERARDI, F. Relative growth, population structure, and shell utilization of the hermit crab Clibanarius erythropus in the Mediterranean. Oebalia, v. 17, p. 181 – 196, 1991.

[25] GRANT, W. C.; ULMER, K. M. Shell selection and aggressive behavior in two sympatric species of hermit crabs. Biol. Bull., v. 146, n. 1, p. 32-43, 1974.

[26] HAMMER, Ø.; HARPER, D. A. T.; RYAN, P. D. PAST: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontologia Electronica 4(1):9pp. http://palaeo-electronica.org/2001_1/past/issue1_01.htm, 2001.

[27] HAZLETT, B. A. The behavioral ecology of hermit crabs. Ann. Rev. Ecol. Syst., v. 12, n. 1, p. 1-22, 1981.

[28] KELLOGG, C. W. Gastropod shells: a potentially limiting resource for hermit crabs. J. exp. mar. Biol. Ecol., v. 22, n. 1, p. 101-111, 1976.

[29] LEITE, F. P. P.; TURRA, A.; GANDOLFI, S. M. Hermit crabs (Crustacea: Decapoda: Anomura), gastropod shells and environment structure: their relationship in southeastern Brazil. J. Nat. Hist., v. 32, n. 10-11, p. 1599-1608, 1998.

[30] MANTELATTO, F. L.; GARCIA, R. B. Shell utilization pattern of the hermit crab Calcinus tibicen (Diogenidae) from Southern Brazil. J. Crust. Biol., v. 20, n. 3, p. 460-467, 2000.

[31] MANTELATTO, F. L.; MEIRELES, A. L. The importance of shell occupation and shell availability in the hermit crab Pagurus brevidactylus (Stimpson, 1859) (Paguridae) population from the southern Atlantic. Bull. mar. Sci., v. 75, n. 1, p. 27-35, 2004.

[32] MANTELATTO, F. L.; ALARCON, V. F.; GARCIA, R. B. Egg production strategies of the tropical hermit crab Paguristes tortugae from Brazil. J. Crust. Biol., v. 22, n. 2, p. 390-397, 2002.

[33] MANTELATTO, F. L.; BIAGI, R.; MEIRELES, A. L.; SCELZO, M. A. Shell preference of the hermit crab Pagurus exilis (Anomura: Paguridae) from Brazil and Argentina: a comparative study. Rev. Biol. Trop., v. 55, p. 153-161, 2007.

[34] MANTELATTO, F. L.; FERNANDES-GÓES, L. C.; FANTUCCI, M. Z.; BIAGI, R.; PARDO, L. M.; GÓES, J. M. A comparative study of population traits between two South American populations of the striped-legged hermit crab Clibanarius vittatus Acta Oecol., v. 36, n. 1, p. 10-15, 2010.

[35] MARKHAM, J. Notes on the growth patterns and shell utilization of the hermit crab Pagurus bernhardus (L.). Ophelia, v. 5, p. 189-205, 1968.

[36] MARTINELLI, J. M.; MANTELATTO, F. L. Occurrence of exobionts in gastropod shells occupied by the hermit crab Loxopagurus loxochelis (Anomura: Diogenidae) in Ubatuba Bay (SP) Brazil. In: Anais do IV Simpósio de Ecossistemas Brasileiros, v. 2, n. 104. Publicações ACIESP, São Paulo, p. 221-226, 1998.

[37] MARTINELLI, J. M.; MANTELATTO, F. L. Shell utilization by the hermit crab Loxopagurus loxochelis (Diogenidae) in Ubatuba Bay, Brazil. In: SCHRAM, F. R. & VANPEL KLEIN, J. R. (eds.). Crust. Biodiv. Crisis, v. 1, p. 719-731, 1999.

[38] MEIRELES, A. L.; MANTELATTO, F. L. Shell use by the Pagurus brevidactylus (Anomura, Paguridae): a comparison between laboratory and field conditions. Acta Zool. Sin., v. 51, n. 5, p. 813-820, 2005.

[39] MEIRELES, A. L.; TEROSSI, M.; BIAGI, R.; MANTELATTO, F. L. Spatial and seasonal distribution of the hermit crab Pagurus exilis (Benedict, 1892) (Decapoda: Paguridae) in the southwestern coast of Brazil. Rev. Biol. Mar. Oceanogr., v. 41, n. 1, p. 87-95, 2006.

[40] MELO, G. A. S. Manual de identificação dos Crustacea Decapoda do litoral brasileiro: Anomura, Thalassinidea, Palinuridea, Astacidea. Plêiade, São Paulo, 551 p, 1999.

[41] MITCHELL, K. A. Shell selection in the hermit crab Pagurus bernhardus. Mar. Biol, v. 35, n. 4, p. 335-346, 1976.

[42] OHMORI, H.; WADA, S.; GOSHIMA, S.; NAKAO, S. Effect of body size and shell avaibality on the shell utilization pattern of the hermit crab Pagurus filholi (Anomura: Paguridae). Crust. Res., v. 24, p. 85-92, 1995.

[43] PESSANI, D.; PREMOLI, C. Some aspects of the biology of Cestopagurus timidus (Crustacea, Paguridae) in relation to the occupied shell. Boll. Zool., v. 60, n. 2, p. 199-205, 1993.

[44] PRASAD, A. M.; IVERSON, L. R.; LIAW, A. Newer classification and regression tree techniques: Bagging and random forests for ecological prediction. Ecosystems, v. 9, p. 181-199, 2006.

[45] PIRES-VANIN, A. M. S.; ROSSI-WONGTSCHOWSKI, C. L.; AIDAAR, E.; MESQUITA, H.; SOARES, L. S. H.; KATSURAGAWA, M.; MATSUURA, Y. Estrutura e função do ecossistema de plataforma continental do Atlântico Sul brasileiro: síntese dos resultados. Publ. esp. Inst. Oceanogr., v. 10, p. 217-231, 1993.

[46] PROVENZANO Jr., A. J. Notes on Bermuda hermit crabs (Crustacea: Anomura). Bull. Mar. Sci. Gulf Caribb., v. 10, p. 117-124, 1960.

[47] R Development Core Team. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. ISBN 3-900051-07-0, URL http://www.R-project.org, 2010.

[48] REESE, E. Behavioral adaptation of intertidal hermit crabs. Am. Zool., v. 9, n. 2, p. 343-355, 1969.

[49] RIOS, E. C. Seashells of Brazil. Rio Grande do Sul. Fundação Universidade de Rio Grande, Instituto Acqua, Museu Oceanográfico de Rio Grande, Universidade de Rio Grande. 2ed. 368 p. + 113pl., 1994.

[50] SATO, Y.; SENO, H. A mathematical consideration for the optimal shell change of hermit crab. J. Theor. Biol., v. 240, p. 14-23, 2006.

[51] SCULLY, E. P. The effects of gastropod shell availability and habitat characteristics on shell utilization by the intertidal hermit crab Pagurus longicarpus Say. J. exp. mar. Biol. Ecol., v. 37, n. 2, p. 139-152, 1979.

[52] SPIGHT, T. M. Availability and use of shells by intertidal hermit crabs. Biol. Bull., v. 152, n. 1, p. 120-133, 1977.

[53] SUZUKI, R.; SHIMODAIRA, H. PVCLUST v 1.2-0. R package version 3.1-34. URL http://www.R-project.org, 2006.

[54] TEROSSI, M.; ESPÓSITO, D. L. A.; MEIRELES, A. L.; BIAGI, R.; MANTELATTO, F. L. Pattern of shell occupation by the hermit crab Pagurus exilis (Anomura, Paguridae) in southern Atlantic region. J. Nat. Hist., v. 40, n. 1-2, p. 77-87, 2006.

[55] TORATI, L. S.; MANTELATTO F. L. Uncommon mechanism of egg incubation in the endemic Southern hermit crab Loxopagurus loxochelis: how is this phenomenon related to egg production? Acta Zool., v. 89, n. 1, p.79-85, 2008.

[56] THERNEAU, T. M.; ATKINSON, B. RPART: Recursive Partitioning. Report by B. Ripley. R package version 3.1-34. URL http://www.R-project.org, 2010.

[57] TUNBERG, B. G.; NELSON, W. G.; SMITH, G. Population ecology of Pagurus maclaughlinae Garcia-Gomes (Decapoda: Anomura: Paguridae) in the Indian River Lagoon, Florida. J. Crust. Biol., v. 14, n. 4, p. 686-699, 1994.

[58] VANCE, R. R. The role of shell adequacy in behavior interactions in hermit crabs. Ecology, v. 53, n. 6, p. 1075-1083, 1972.

[59] VANCE, R. R. Competition and mechanisms of coexistence in three sympatric species of intertidal hermit crabs. Ecology, v. 53, n. 6, p.1062-1074, 1972a.