Fishes other than anemonefishes that associate with sea anemones

The obligatory relationship of sea anemones andanemoneﬁshes(Amphiprionspp.andPremnasbiaculea-tus) and damselﬁshes of the genus Dascyllus is wellknown.Wesummarizetheliteratureandaddpersonalobservationoffacultativeassociationswithseaanemo-nesbycardinalﬁshes(Apogonidae),wrasses(Labridae),hawkﬁshes (Cirrhitidae), butterﬂyﬁshes (Chaetodonti-dae),ascaledblenny(Clinidae),andevenatemperategreenling (Hexagrammidae). Although most of theseﬁshesappeartoavoidthetentacles,somemakecontact.The behavioral and physiological facets of these rela-tionshipsremaintobestudied.The27speciesoftheIndo-PaciﬁcanemoneﬁshgenusAmphiprion and the monotypic anemoneﬁsh genusPremnasare well known for their obligate associationwithseaanemones.AtleasttwootherspeciesofIndo-PaciﬁcdamselﬁshesofthegenusDascyllusalsoassociatewith anemones as juveniles (Fautin and Allen 1992).Thesepomacentridﬁshesliveamongthetentaclesoftheanemoneswhichbearnematocysts–microscopicstingingcapsules–thatprovideforthedefenseoftheanemoneand capture prey, including small ﬁshes, that come incontact with the tentacles. Because sea anemones, likeothercnidarians,canbedeadlytoﬁshes,earlyreportsofassociations between them were subject to skepticism.Collingwood(1868a,1868b)ﬁrstdescribedtherelation-shipmorethanacenturyaftersomeoftheanemoneﬁshesandtheirhostswereindividuallydescribed.Lesswellknownareotherﬁshesthatshelterattimesamong the stinging tentacles of large sea anemones.ColinandHeiser(1973)reportedtwoWestIndiancar-dinalﬁshes(Apogonidae),ApogonquadrisquamatusandA.aurolineatus,thatoccasionallyassociatewiththeseaanemonesBartholomeaannulataandCondylactisgigan-tea.Theyaddedthattheﬁsheswerenotimmunetothenematocysts,andmanyhadwhitespotsandlesionsthattheybelievedwerecausedbycontactwiththetentacles.Hanlon et al. (1983) reported more than 30 species ofWestIndianreefﬁshesthatoccasionallydwellwithinthesphereofthetentaclesofseaanemones,principallyasjuveniles.OfsevenspeciesdiscussedindetailbyHanlonet al., all but one avoided touching the tentacles. TheexceptionwasthelabrisomidStarksiahassiwhichmadefullcontactwiththetentacleswithnoapparentharm.J.E.R. took several photographs of a group of theWestIndiancardinalﬁshApogonquadrisquamatusshel-teringamongthetentaclesofaCondylactisgiganteaatBonaire in the southern Caribbean Sea, one of whichwaspublished(Randall1996:Fig.149).Theseﬁshwereusing the anemone during the day for shelter and notjustretreatingtoitfortemporaryasylum,anditseemedthatatleastoccasionaltentaclecontactwasmade.Cardinalﬁshes associate with sea anemones in thetropical Indo-Paciﬁc as well the West Indian region.Somehovernearananemonefortheprotectionitpro-videsbutappeartoavoidthetentacles.AnexampleisApogonnanuswithatubeanemone(orderCeriantharia)at Flores, Indonesia (Fig.1). Another is Pterapogonkaudernifrom Sulawesi (Fig.2) withwhat is probablytheanemoneHeteractiscrispawhichalsohosts‘‘true’’anemoneﬁshes. D.G.F. has seen apogonid ﬁshes ineastern Indonesia around members of the anemonefamily Actinodendronidae, termed Hell’s Fire Anemo-nesbecauseoftheirpowerfulsting.The cardinalﬁsh Apogonmoluccensis(A.ventrifasci-atus is a synonym) deﬁnitely makes contact with the