Anomuran Species (Crustacea, Decapoda) And Their Ecological Distribution At Fortaleza Bay Sublittoral, Ubatuba, Sao Paulo, Brazil

8 Pages·1997·3.4 MB·English

by  Adilson Fransozo

ANOMURAN SPECIES (CRUSTÁCEA, DECAPODA) AND THEIR
ECOLOGICALDISTRIBUTIONATFORTALEZA BAY SUBLITTORAL,
UBATUBA, SÃO PAULO, BRAZIL
MariaLuciaNegreiros-Fransozo^
AdilsonFransozo^
Fernando Luis MedinaMantelatto^
Marcelo Antonio Amaro Pinheiro^
Sandro Santos'*
ABSTRACT
Thecompositionanddistributionoftheanomuranspecieswhichoccuralongthenon-consoli-
dated sublittoral ofFortaleza bay, Ubatuba, São Paulo, Brazil, were determined. Samplings were
monthlycarriedoutfromNovember1988toOctober1989in7transectsof1kmeachusingashrimp
fishing boat equipped with two otter-trawls. Seven anomuran species of the families Diogenidae,
PaguridaeandPorcellanidaewererecorded.ThemostabundantspecieswereLoxopagurusloxochelis
(Moreira, 1901),Dardanusinsignis(Saussure,1858) andIsochelessawayai(Forest &SaintLaurent,
1967).Thecrabswerefoundin11generaofgastropodshells,withapredominanceofThaisRöding,1798
andOlivanciliariaOrbigny,1839.Thespatialdistributionoftheanomuranswasheterogeneous.The
highestincidenceofthoseanimaisoccurringinthetransectII isprobablyassociatedwiththefeatures
ofbottomSediment.
KEYWORDS.Anomura,ecologicaldistribution,diversity,Braziliancoast.
INTRODUCTION
According to McLaughlin (1983), pagurid crabs comprise more than 800
species in 86 genera and six families being 12 species semiterrestrial and the
remainder, marine. Theyrepresent aparcel ofnotorious importance in the inter-
tidalandmoderatelydeepbenthicCommunity,wheretheyplayanimportantrole
in the marine food chain.
1. DepartamentodeZoologia,InstitutodeBiociênciaseCentrodeAquiculturadaUNESP,18618-000Botucatu,SãoPaulo,Brasil.
2. DepartamentodeBiologia,FaculdadedeFilosofia,CiênciaseLetrasdeRibeirãoPreto, USP,14040-901RibeirãoPreto,SãoPaulo,
Brasil.
3. DepartamentodeBiologiaAplicada,FaculdadedeCiênciasAgráriaseVeterinárias,UNESP,14870-000Jaboticabal,SãoPaulo,Brasil.
4. DepartamentodeBiologia,CentrodeCiênciasNaturaiseExatas,UniversidadeFederaldeSantaMaria,97119-800SantaMaria,Rio
GrandedoSul,Brasil.
Iheringia,Sér.Zool.,PortoAlegre, (83): 187-194,30out. 1997
188 Negreiros-Fransozo etai.
In Brazil, few studies are available on hermit crabs: some dealing with
systematics and distribution (Forest & Saint Laurent, 1967; Tommasi, 1967;
Narchi & Hebung, 1976; Hebung & Rieger, 1986; Hebung et al. 1994); others
dealing with larval development (Hebung & Fransozo, 1982; N.-Fransozo &
Hebling, 1983 andHebung & Mansur, 1995); andstill others are relatedtoshell
choice(N.-Fransozoetal.,1991a;N.-Fransozo&Fransozo,1992;PiNHEiROetal.,
1993). AWidestudyonmacrocrustaceanfrombenthicCommunitywascarriedout
in Fortaleza bay (23° 31' S and 45° 09' W), Brazil by N.-Fransozo et al. (1991b),
Fransozo et al., 1992; Santos et al. (1994); Mantelatto et al. (1995) and N.-
Fransozo & Fransozo (1995) inwhich the brachyuranswere emphasized.
,
The objective of this work was to determine the anomuran crab species
occurring alongthe non-consoHdated sublittoralofFortalezabay,Ubatuba (SP),
andtheirecological distribution.
MATERIALANDMETHODS
TheanomuransweresampledinFortalezabay,Ubatuba,SãoPaulousingashrimpfishingboat
equippedwithtwootter-trawls,eachonemeasuring7.5minlengthand10mmofcodmesh.SampHngs
were monthlyperformedovera 12months (fromNovember 1988to October 1989). EachsampHng
consistedofseventransectsof1kmeach(fig.1).Fortalezabayhasameandepthof9m(4.4to13.3m),
andpresentedthefollowingmeanannualvaluesforthehydrologicfactorsanalyzedduringthestudy
period:temperature,23.5°C(21 to28.1°C);sahnity,34.4°/^(32.4to35.6°/J,anddissolvedoxygen,5.46
mg/1(4.2to6.33mg/1).ThemeanannualcontentoforganicmatterintheSedimentwas4.07% (2.3to
6.6%)andthegranulometriccompositionofthesedimenthad,duringmostseasons,predominanceof
fineandveryfinesandfractions.MeanannualrainfallfortheUbatubaregionduringthestudyperiod
was 154.3mm(27.4to282.6mm).Adetaileddescriptionofsamplingandanalysisofenvironmental
factorsatFortalezabay,inthesameperiodwaspreviouslypubhshedby N.-Fransozoetal.(1991b).
TheShellsgenerawereidentifiedasindicatedbyRios(1970)andthehermitcrabsasindicated
byForest&SaintLaurent(1967)andHEBLiNO&Rieger(1986).Thebiologicalmaterialisdeposited
inthescientificcollectionoftheDepartamentodeZoologia,InstitutodeBiociências,UNESP,Campus
deBotucatu.Thepercentageofassociationofhermitcrabswiththegenus ofeachgastropodshellwas
determined.
ThediversityindexofShannon-WeaverwascalculatedaccordingtoPooLE(1974),wherethenull
hypothesiswasH^:H^=¥i^,anditwastestedbyattest(P<0.05). Thesimilaritybetweenthetransects
wasanalyzedbytheJaccardindexasindicatedbyKrebs(1989),wherethelevelofsignificancewasset
at5%.
RESULTS
Atotalof161specimensoftheinfraorderAnomurawererecorded.Ofthese,
126 belonging to the Diogenidae family: Petrochirus diogenes (Linné, 1758),
Dardanusinsignis (Saussure, 1^5^),Loxopagurusloxochelis (Moreira, 1901),and
Isochelessawayai (Forest & SaintLaurent, 1967). Nineteen specimens are repre-
sentativesofthefamilyPaguridae:Pagwrw^criniticornis(Dana,1852)dináPagurus
leptonyx(Forest&SaintLaurent,1967).ThePorcellanidaewererepresentedonly
byPorcellanasayana(Leach,1820)with16individuais.Theabsoluteabundanceof
these species (tab. I) shows predominance of diogenids over pagurids and
porcellanids. The most abundant specieswas L. loxochelis.
The anomuran species showed differential distribution in the studied area
with allthe seven recorded species present onlyinthe transect IL The species of
Iheringia,Sér.Zool.,PortoAlegre, (83): 187-194,30out. 1997
Anomuranspecies(Crustácea,Decapoda)... 189
II' 10' 9' 8' T 45°
Fig. 1. FortalezabayinUbatuba,SãoPaulo,showingthepositionofthesamplingtransectsandstaions.
widest spatial distribution was L. loxochelis (not detected only in transect VI)
followedbyP. sayana,whichwas absentintransectIV. Allthe anomuranspecies
presented discontinuous continuous occurrence alongthe collectingmonths.
The most abundant hermit crab species was L. loxochelis followed by D.
insignisand/. sawayai. The abundance ofthe anomuranspecieswashigherinthe
transectswith higher organicmattercontent (%) (fig. 2).
Eleven gastropod genera Shells were found to be inhabited by the hermit
crabs (tab. III). Thepredominantgenerawere Thais Röding, 1798 (32.14 %) and
Olivancillaria Orbigny, 1839 (27.14 %).
The similarityindex (fig. 2) revealsthattransectsII andVmaybe grouped.
Thediversityindex(tab.III)foreachtransectevidencedslightlydifferences
V
inrelationtohermitcrabs.TransectsIIand presentedthehighest diversityindex
ofShannon-Weaver, where there was no significant difference at the 5% level.
DISCUSSION
ThesevensampledareasintheFortalezabaywerereasonablyhomogeneous
intermsoftemperature,salinityanddissolvedoxygen(N.-Fransozoetah,1991b).
Theenvironmentalfactorsofwidest Variation,producingecologicallycontrasting
areas, were depth, granulometriccomposition, and organicmatter content in the
Sediment.Amongthem,the Sedimentfeaturesarethemostrelatedtothebenthic
organisms distribution (Fransozo etal., 1992).
The heterogeneous distribution of anomurans in the Fortaleza bay with
respecttothesamplingtransectsisprobablyassociatedwiththecontrastingleveis
of organic matter and granulometric composition of the sediment, reflecting
specific habitats. The action of organic matter is particularly evident and the
abundance ofanomuranswasnegativelycorrelatedwiththisparameter. Thisfact
Iheringia,Sér.Zool.,PortoAlegre, (83): 187-194,30out. 1997
I
190 Negreiros-Fransozo etai. I
TableI.ListoftheAnomuranspeciesandnumberofindividuaissampledpertransectatFortalezabay,
SP,fromNovember 1988toOctober1989. i
FAMILIES TRANSECTS
Species I II III IV V VI VI Total
DIOGENIDAE
Loxopagurusloxochelis 22 28 8 4 14 2 78
Isochelessawayai 1 1 19 - - 21
Petrochirusdiogenes 1 - - 2 - 5
Dardanusinsignis 3 8 - 2 9 - 22
PAGURIDAE
Paguruscriniticornis 13 _ _ _ . 13
Pagurusleptonyx 3 - - 2 - 6
PORCELLANIDAE
Porcellanasayana 1 1 1 - 10 1 2 16
Totalofspecies 3 7 3 3 5 3 2 7
Totalofindividuais 26 55 10 25 37 4 4 161
TableII.RelationshipofthehermitcrabspecieswiththegeneraofgastropodshellssampledinFortaleza
Bay,SP,fromNovember1988toOctober1989.(1=Loxopagurusloxochelis;2=Isochelessawayai;3
=Petrochirusdiogenes;4=Dardanusinsignis;5=Pagurus criniticornis;6=Pagurusleptonyx).The
absolutevaluesrecordedaregiveninbrackets;5ofthe145specimensrecordedhadnoshell.
Hermitcrabs
Gastropods 1 2 3 4 5 6 Total
FusinusRafinesque,1815 - - - - 15.38 - 1.43
(2) (2)
OlivancillariaOrbigny,1839 42.46 23.81 - 4.55 7.69 - 27.14
(31) (5) (1) (1) (38)
ThaisRöding,1798 31.51 66.67 - 9.09 38.45 16.67 32.14
(23) (14) (2) (5) (1) (45)
BuccinanopsOrbigny,1841 19.18 4.76 - 4.55 7.69 66.66 15.0
(14) (1) (1) (1) (4) (21)
ChiroreusMontfort,1810 1.37 - 4.55 23.09 16.67 4.29
(1) (1) (3) (1) (6)
HastulaH.&A.Adams,1853 1.37 - - - - - 0,71
(1) (1)
CerithiumBruguière,1789 - - - 7.69 " 0.71
(1) (1)
Po/m/ce5Montfort,1810 1.37 - - - - - 0.71
(1) (1)
CymatiumRöding,1798 2.74 4.76 - 45.44 - - 9.29
(2) (1) (10) (13)
TonnaBrunnich,1772 40.0 27.27 - - 5.72
(2) (6) (8)
StrombusLinnaeus,1758 ' 60.0 4.55 " 2.86
(3) (1) (4)
Total 73 21 5 22 13 6 140
Iheringia,Sér.Zool.,PortoAlegre, (83): 187-194,30out. 1997
Anomuranspecies(Crustácea,Decapoda). 191
wasobservedforClibanariusvittatus (Bosc, 1802),whichinhabitsareasofmuddy
sand that accumulate more detrital material, which appears to be a major food
source forthis species (Caine, 1980).
A great abundance ofIsochelessawayaiwas observed in transect IV, which
is characterized by smaller amounts of organic matter and by lower sahnity and
depth. Furthermore, the subárea corresponding to this transect presents a pre-
dominanceofveryfinesand(7.58%),afactthatfavoursgreatercompactingofthe
bottom, allowingforanimaltostayburiedandnottobedraggedaway,especially
in the surfzone. The presence ofthe species in this subárea may be because this
05[-1 3.5 3.5 --] 6.5 6.5 —J 9.5
ORGANIC MATTER (%)
Fig.2.Relativefrequencyofhermitcrabsasafunctionofclassoforganicmattercontent.
SIMILARITY
,00% II V VI I III VII IV
7 l%J
67%
53%^
42%
32%
J
Fig.3.DendrogramofsimilaritybetweentransectsobtainedbyapplyingtheJaccardindex.
Iheringia,Sér.Zool.,PortoAlegre, (83): 187-194,30out. 1997
192 Negreiros-Fransozo etai.
hermit crabis aSuspensionfeederone,whichpreferssiteswithlargeramountsof
particlesinSuspension.Thisspeciestolerateslowersalinityasitisoftenfoundnear
sites influenced by freshwater.
Ovigerous females were detected only in Loxopagurus loxochelis (5 speci-
mens),Petrochirusdiogenes(1specimen)andDardanusinsignis(1specimen).The
Table III. Diversity index calculated foreach transect (N = number ofindividuais; H and Hmax =
Shannon-Weaverindexasbits/individual;E=equitability;V=variance).
INDEX I II III IV V VI VII
N 26 55 10 25 37 4 4
H
0.744 1.936 0.639 1.015 1.992 1.5 1
Hmax 1.585 2.807 1.099 1.585 2.322 1.585 1
E 0.469 0.690 0.582 0.641 0.858 0.946 1
V 0.849 0.608 0.528 0.530 094 0.981 0.23
low incidence of ovigerous females suggests a spawning behavior involving
differencialhabitatoccupationbythosefemales.N.-Fransozo &Hebung (1983)
reported that Isocheles sawayai is found in the surf zone during spawning and
incubation, a fact explaining the total absence of this species in the sampling
transects.
The occupation ofgastropodShells ofthe genera Thais andOlivancillariais
propablycorrelatedwithagreaterabundanceofthesespecimensinthearea.This
occurrence was probably due to the action oflocal currents and tides. The lower
plasticityofshelltypesmayexertpressureonthestructureofspeciespopulations
intheregion, aspreviouslyobservedforotherhermitcrabs, alteringgrowthrates
(WiLBER, 1990), aggressiveness (Hazlett, 1967) andfecundity (Bertness, 1981).
According to Pires (1989) who studied the structure and dynamics of the
benthic megafauna at several sites along the continental shelf of the northern
region ofthe State ofSão Paulo, mollusks represented the third most numerous
group in the internal domain, especially in spring, and gastropods were always
dominant among them, representing 60 to 73% of the Mollusca fauna sampled.
Other areas alongthe northern littoral ofSão Paulo State also presented greater
abundance ofgastropods comparedthe othergroups ofthe marine malacofauna.
ThisaspecthasbeenwelldefinedinastudyontheSãoSebastiãoCanalconducted
by MiGOTTO et al. (1993). These records may support the relatively successful
presence of hermit crabs in the region. According to Rittschoff (1980), hermit
crabscanobtainnewShellsbysimplychangingthem,byfindingemptyShellsorby
lookingforsitesweregastropods arepreyeduponbyotheranimais. Thefirsttwo
eventsseemtooccurinFortalezabay,butnoreportsofthemexistintheliterature.
Fransozoetal.(1992)recordedagreatabundanceofbrachyuransinFortalezabay
which are known to prey on gastropods (Wear & Haddon, 1987; Choy, 1986;
Haefner,1990).Forinstance,astudymadebyMANTELATTO&Petracco(inpress)
on Hepatuspudibundas (Herbst, 1785) diet revealed that Gastropoda were the
third item in the feeding preference ofthese animais.
Analysis of present data suggests that the occurrence and distribution of
Iheringia,Sér.Zool.,PortoAlegre, (83): 187-194,30out. 1997
.
Anomuranspecies(Crustácea,Decapoda)... 193
anomurans,andhermitcrabsinparticular,dependsnotonlyonfavorableenviron-
mentalfactorsbutalso onbiogenicfactorssuch asshell availability.These results
confirmthoseobtainedalsoforbrachyurans(Fransozoetal.,1992),indicatingthat
enviromentalfactors,whenactingtogether,aredecisiveintheecologicaldistribu-
tion ofthe benthic organisms.
AppHcation ofthe similarity index revealed that transects II and V may be
groupedbecausetheyshare anumberofspecies. This factsupports the similarity
ofthesesubareasobservedbyN.-pRANSOzoetal.(1991b)attheenviromentallevel.
Recently,Hebungetal.(1994)carriedoutasimilarstudyinAnchietaIsland
region. São Paulo State, where 12 anomuran specieswere recorded, for atotal of
323 animais.Thisfactdemonstrates agreaterwelthandabundanceofanomurans
for an area close to Fortaleza bay. Nevertheless, the total diversity index for
Fortalezabay(H-2.2388bits/ind. Equitability=0.7975;Variance=0.0687)andfor
;
AnchietaIsland(H=2.7481bits/ind.;Equitability=0.7666;Variance=0.0309)does
not differs significantly (p>0.05; t = 1.61), which means a similar anomuran
compositioninneighboringareasmonitoredatdifferenttimes. Asobservednear
Anchieta Island and in Fortalezabay, the geographicposition ofthe transects in
relationtothelandcharacterizesdifferentenvironmentalconditions,implyingthat
differentinfluences exist on the distribution ofbenthicspecies.
Acknowledgments.Thisresearchwas supportedbyConselhoNacionaldeDesenvolvimento
CientíficoandTecnológico(CNPq)andbyFundaçãodeAmparoàPesquisadaUNESP(FUNDUNESP)
ToDr.NiltonJoséHeblingforconfirmingtheIdentificationofthehermitcrabs,andMsc.AugustoA.V.
FloresandEduardoA.Geraqueforhelpinginthelaboratorywork.
REFERENCES
BERTNESS, M. D. 1981.The influence of shell-type on hermit crab growth rate and clutch size
(Decapoda,Anomura).Crustaceana,Leiden,40(2):197-205.
CAINE,É.A.1980.Adaptationsofaspeciesofhermitcrab(Decapoda,Paguridea)inhabitingsessile
wormtubes.Crustaceana,Leiden,38(3):306-310.
CHOY,S.C.1986.ReproductivebiologyofLiocarcinuspuberandL.holsatus(Decapoda,Brachyura,
Portunidae) from the Gower Península, south Wales. Mar. Ecol. Prog. Ser., AmeUnghausen,
9(3):227-241.
FOREST, J. & SAINT LAURENT, M. de 1967. Campagne de la "Calypso" au large des cotes
Atlantiques de TAmérique du sud (1961-1962). 6. CrustacésDécapodes: Pagurides. Annls.Inst.
océanogr.Monaco,Mónaco,45(2):47-169.
FRANSOZO, A.; N.-FRANSOZO, M. L. et al. 1992. Composição e distribuição dos Brachyura
(Crustácea, Decapoda) do subhtoral não-consolidado na Enseada da Fortaleza, Ubatuba (SP).
Revtabras.Biol.,RiodeJaneiro,52(4):667-675.
HAEFNERJr.,P.A.1990.NaturaldietofCallinectesornatus(Brachyura,Portunidae)inBermuda.J.
Crust.Biol.,SanAntonio,10(2):236-246.
HAZLETT,B.A.1967.InterspecificshelfightingbetweenPagurusbernhardusanáPaguruscuanensis
(Decapoda,Paguridea).Sarsia,Oslo,29:215-220.
HEBLING,N.J.&FRANSOZO,A. 1982.Desenvolvimentopós-embrionáriodePaguristeserythrops
Holthuis,1959(Decapoda,Diogenidae),emlaboratório.Revta bras.Biol., RiodeJaneiro,42(1):
117-128.
KEBLING,N.J. &MANSUR,C.B. 1995.DesenvolvimentolarvaldeDardanusinsignis (Saussure,
1858)(Crustácea,Decapoda,Diogenidae),emlaboratório.Revtabras.Zool.,SãoPaulo, 12(3):471-
491.
HEBLING,N.J.;MANTELATTO,F.L.M. etal.1994.Levantamentoedistribuiçãodebraquiúros
Iheringia,Sér.Zool.,PortoAlegre, (83): 187-194,30out. 1997
194 Negreiros-Fransozo etai.
eanomuros(Crustácea,Decapoda)dossedimentossublitoraisdaregiãodaIlhaAnchieta,Ubatuba
(SP). Bolm.Inst.Pesca,SãoPaulo,21:1-9.
HEBLING,N.J.&RIEGER,P.J.1986.Osermitões(Crustácea,Decapoda:PaguridaeeDiogenidae)
dolitoraldoRS,Brasil.Atlântica,RioGrande,8:63-77.
KREBS,C.J. 1989.EcologicalMethodology. NewYork, Harper&Row,654p.
MANTELATTO, F. L. M.; FRANSOZO, A. & N.- FRANSOZO, M. L. 1995. Distribuição do
caranguejoHepatuspudibundas(Herbst, 1785) (Crustácea,Decapoda)naEnseadadaFortaleza,
Ubatuba(SP),Brasil.BolmInst.Oceanogr.,SãoPaulo,43(1):51-61.
MANTELATTO,F.L.M.&PETRACCO,M.(inpress)NaturaldietofthecrabHepatuspudibundas
(Crustácea, Brachyura, Calappidae) in Fortaleza bay, Ubatuba (SP), Brazil. J. Crust. Biol., San
Antonio.
McLAUGHLIN, P. 1983. Hermitcrabs - are theyreallypolyphyletic? J. Crust. Biol., San Antonio,
3(4):608-621.
MIGOTTO,A.E.;TIAGO,C.G. &MAGALHÃES,A.R.M.1993. Malacofaunamarinhadaregião
costeiradoCanaldeSãoSebastião,SP,Brasil:Gastropoda,Bivalvia,PolyplacophoraeScaphopoda.
BolmInst.Pesca,SãoPaulo,41(l/2):13-27.
NARCHI,W.&HEBLING,N.J. 1976.Calcinustibicen(Herbst,1791)inBrazilianlittoral(Decapoda,
Paguridea).PapéisAvulsZool.,SãoPaulo,26(6):67-71.
N.-FRANSOZO,M.L.&FRANSOZO,A. 1992.Estruturapopulacionalerelaçãocomaconchade
PaguristestortugaeSchmitt,1933(Decapoda,Diogenidae)nolitoralnortedoEstadodeSãoPaulo,
Brasil.Naturalia,RioClaro,17:31-42.
1995. OnthedistributionofCallinectesornatasOrdway,1863and CallinectesdanaeSmith,1869
. . (Brachyura,Portunidae)inFortalezabay,Ubatuba(SP),Brazil.IheringiaSér.Zool.,PortoAlegre,
(79):13-25.
N.-FRANSOZO, M. L.; FRANSOZO, A. & HEBLING, N. J. 1991. Estrutura populacional e
determinaçãodotamanhodaconchaocupadapor4espéciesdeermitões(Crustácea,Decapoda,
Anomura)dolitoraldeSãoPaulo.Biotemas,FlorianópoHs,4(2):135-148.
N.-FRANSOZO,M.L.;FRANSOZO,A.;PINHEIRO,M.A.A.etai. 1991.Caracterizaçãofísicae
químicadaEnseadadaFortaleza,Ubatuba(SP).Revta.bras.Geoc,SãoPaulo,21(2):114-120.
N.-FRANSOZO, M. L. & HEBLING,N. J. 1983. Desenvolvimento pós-embrionário de Isocheles
sawayai,Forest&SaintLaurent,1967(Decapoda,Diogenidae)emlaboratório.PapéisAvulsZool.,
SãoPaulo,35(4):41-53.
PINHEIRO,M.A.A;FRANSOZO,A.&N.-FRANSOZO,M.L.1993.Seleçãoerelaçãocomaconcha
emIsochelessawayaiForest&SaintLaurent,1967(Crustácea,Anomura,Diogenidae).Arq.Biol.
Tecnol.,Curitiba,36(4):745-752.
PIRES,A.M.S.1989.EstruturaedinâmicadamegafaunabênticanaplataformacontinentaldaRegião
NortedoEstadodeSãoPaulo,Brasil.172p.TesedelivredocênciaIO-USP,SãoPaulo,SP. [Não
pubhcada]
POOLE,R.W. 1974.Anintroductiontoquantitativeecology. N.York,McGraw-Hill,532p.
RIOS,E.C,1970.CoastalBrazilianseashells.RioGrande, FundaçãoCidadedoRioGrande,Museu
OceanográficodoRioGrande,255p.
RITTSCHOFF, D. 1980. Chemical attractions of hermit crabs and other attendants to simulated
gastropodpredationssities.J.Chem.EcoL,NewYork,6:103-118.
SANTOS,S.;N.-FRANSOZO,M.L.&FRANSOZO,A.1994.Thedistributionoftheswimmingcrab
Fortunas spinimanus Latreille, 1819 (Crustácea, Brachyura, Portunidae) in the Fortaleza bay.
Atlântica,RioGrande,16:125-141.
TOMMASI,L.R.1967.Observaçõespreliminaressobreafaunabênticadesedimentosmolesdabaía
deSantoseregiõesvizinhas.BolmInst.oceanogr..SãoPaulo,16(l):43-55.
WEAR, R. G. & HADDON, M. 1987. Natural diet of the crab Ovalipes catharus (Crustácea,
Portunidae) aroundcentralandnorthernNewZealand. Mar.Ecol.Prog. Ser., Amelinghausen,
35:39-49.
WILBERJr.,T. P. 1990. Associationsbetweencrab morphologyand gastropod shell speciesinthe
hermitcrabPaguruslongicarpus.J.Crust.Biol.,SanAntonio,10(1):134-138.
Recebidoem06.12.1996;aceitoem02.07.1997.
Iheringia,Sér.Zool.,PortoAlegre, (83): 187-194,30out. 1997