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Feeding habits of Calma glaucoides (Alder & Hancock, 1854): its adaptive structures and behaviour PDF

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BollettinoMalacologico,Roma,37(5-8):177-180,2001 -SocietàItalianadìMalacologìa Feeding habits of Calma glaucoides (Alder & Hancock, 1854): its adaptive structures and behaviour Gommalo Calado & Victoriano Urgorri KEYwords: Aeoliclacea;Calmagla/midey.Feeding ABSTRACT Calmaglaucoides(Alder& Hancock, 1854) isan aeolid nudibranch inhabiting European (Atlantic and Mediterranean)Westernand Southern coasts. Adultsfeedonfisheggs, thusbeingfound nearspawningsitesontheundersideofbouldersorinsmallcrevices,wheremalefish nest.Thisaeolidhas an atypical uniseriate radula. Littleis knownaboutformationorfunctionofthis radula, but it isbelieved that such apeculiarmorphology is related toadult'sfeedinghabits.Thefeedingbehaviourofindividualscouldbeobservedinaquaria. Itwasalsopossibletostudynotonlyradulaefromindivi- duals ofdifferent sizes but also the damage caused by radula action upon fish egg membranes. These were visible under SEM after critical point dryingofemptyeggs. SEM imagessuggest that the radula is usedasasaw. Aftertheegg is "opened" theyolkandembryoaresuckedoutwith the help ofstrong buccal musculature, leaving the external membrane ofthe egg almost intact. The way these animals puncture fish eggs, associated withtheearlydevelopmentoftheirradula,leadustoconclude,thatthereseemstobenofoodsizeconstraint,aspreviouslyhadbeenthought. RIASSUNTO Cí?/wí?g/íí/i'íWíí'er(Alder&Hancock, 1854)èuneolidiaceopresentelungolecostedell’Atlanticooccidentaleedelmediterraneo.L'adultosi nutredi uova dipescievienespessoritrovatovicinoalleovature,sullatoinferioredei massioinpiccolefessuredellaroccia.Questoeolidiaceohaunaradulauniseriata moltoatipica. Pococonosciutesonolasuaformazioneefunzione, masi pensachelatipicamorfologiadebbaesserecorrelataalcomportamentotrofico dellaspecie.Graziealmantenimentodialcuniesemplari inacquario,èstatopossibilestudiareilcomportamentodiquestaspecie,lamorfologiaradulare in individui di diversatagliae i danni causati dalla radulastessasullamembranadelle uovadei pesci. Immagini al SEM,dopo il criticai point, delle uova indicanoche la radulavieneusatacome unasega. Dopochel’uovoèstatotagliato, il tuorloe l’embrionevengono risucchiati grazieall'azionedi unatortemuscolatura, lasciandopraticamente intattalamembranaesternadell’uovo. Il modoconcui questi nudibranchi taglianoleuova, in rapporto anchealprimosviluppodellaradula,suggeriscechelatagliadelleuovanonsiaunfattorelimitante,comeprecedentementesupposto. G. CALADO. V. URGORRI 1InstitutoPortuguésdeMalacologia,ApartadoNo. 52,S. PedrodoEstoril,2766-501 Estoril,Portugal.E-mail: bacongasís’mail.telepac.it 2LaboratoriodeZooloxíaMariña,DepartamentodeBioloxíaAnimal,UniversidadedeSantiagodeCompostela, 15706SantiagodeCompostela,Spain. INTRODUCTION collected individuals ofC. glaucoides were kept in small beakers Most aeolid nudibranchs feed on cnidarians and therefore they with small stones containing portions of Parablennius pilicornis possess strong jaws and uniseriare or triseriate radulae, with lew spawn. After consumption of the egg contents, empty egg exceptions in both characteristics. Calma glaucoides (Alder & membranes were dehydrated in alcohol, and observed in SEM after Hancock, 1854), an aeolid nudibranch inhabiting European criticalpoint treatment. (Atlantic and Mediterranean) Western and Southern coasts, is precisely one such exception. Adults (7 to 18 mm) are found on RESULTS their food supply, the spawn of teleost fishes such as Lepadogaster Radularmorphology lepadogaster, Lepadogaster candollei, Parablennius gattorugine and SEM images allowed a closer examination of seemingly Parablenniuspilkomis, which deposit their eggs on the underside of important details. Small teeth ofthe serration measure about 1mm boulders or empty bivalve shells. C. glaucoides has a very atypical (Eig. 1) and in high magnification (top image) it is possible to see “uniseriate” radula, previouslydescribed as arowofverysmall teeth some erosion on their outer edges (Eig. 2). In small animals, the and a group ol lew bigger teeth at the distal end. Little is known first-formed “big” teeth were smaller than in later ones, as ifthey about theformationorfunctionofthisradula, but it isbelieved that were in formation after differentiation of the distal edges. Radula such a peculiar morphology is related with adult’s feeding habits size differs relating little between animals having a body length of (E'VANS, 1922; Ro'WETT, 1946). The extent ofsuch a relation is the 1.4 mm from those of 2.5 mm (Eig. 3). This suggests that the subjectmatterofthisstudy. radular morphology, developed early in the ontogeny, is preserved throughout adult life. Similar arrested morphological development MATERIALS AND METHODS of the radula is known, forexample, from the Harpidae (HUGHES& We worked with specimens obtained from the Iberian Atlantic Emerson, 1987). NW Coast (Galicia - Spam (43" 27’ N, 8" 16’ W) and Arrábida, Portugal (38° 27’ N, 9° 0’ W)). They were preserved in 70% Feedingbehaviour ethanol. Buccal masses were extracted and placed in a 5% NaOH In order toconsume an egg, Calmaglaucoides crawls over the fish solution for 24 h at 40" C. When radulae were detached they were spawn. Then it proceeds to enwrap an egg with its oral tentacles; abundantly rinsed in tap water. Six radulae for SEM were dried and immediately afterwards we can see, through the almost transparent coatedwithgoldpriortoexamination. body,theeggcontentsbeingsucked in andrapidlymovedalongrhe In order to observe their feeding behaviour, rhree recently oesophagus to the stomach. During the feeding^process, > 177< GongaloCalado&VictorianoUrgorri Fig. 1 -SEMpictureofaradulawheresaw-liketeetharevisible. Fig. 2-Topimageofserraredbarwithsaw-liketeeth. > 178< FeedinghabisofClamaglaucoides Fig. 3-SEMviewsoftworadulae.Left: froma 1.4mm longspecimen. Right: froma2.5 mm longspecimen.Scalebarforbothpictures. Fig.4- PredatedParahlevnimpilicurniseggs. > 179< GongaloCalado&VictorianoUrgorri Fig. 5 - Detailoftheincision intheeggmembrane.Arrowindicatesouterborder. immediately beforesuction theanimal usually bends forwatds all its SEM imagesofempty fishegg membranesshowopenings, made cerata, letting them return to the "normal” position after the actual by the molluscs, that are consistently regular as ifcut with a suction begins. The opening ofthe egg membrane was observed to "surgical " instrument and not just torn open. The openings be a quick task. If by any chance the egg already contains an observed were morphologically identical and suggest a sawing-like embryo, it is apparently crushed during suctionprocess, as we see it method ofcutting, precise and efficient. The observation ofliving into small pieces. Theprocedure above is repeated withasuccession animals while feeding also indicates that this procedure is a rapid ofeggs, until the animal is satiated, leaving behind avariable band one. We conclude that the saw-like portion ofthe radular tooth is ofemptyegg membranes. Under laboratory conditions weobserved used tocut open the membranewith aminimum effortallowing its up to 30 eggs being continuously consumed, in less than one hour, innercontent(which maycontainhardparts, iffish's larvaisalready byasingleadult individual. formed)tobeextracted. SEM images ofempty fish egg membranes (figs. 4 and 5) allow us to see the size and shape ofthe opening made by the mollusc. It ACKNOWLEDGEMENTS clearlyshowsevidenceofradularactionasdiscussedbelow. Thisworkwascarriedoutwhilefirstauthorwas holderofagrant from "Fundai^ao para a Ciencia e Tecnologia”- Portugal DISCUSSION (PtaxisXXI/BD/11185/97). We thank Ilidio Félix-Alves (I. P. M. - Erosionon theouterborderofteeth,on theserratedportionofthe Portugal) for his critical teview ofthe English version ofrhe text. tadula, made us believe that these teeth ate functional, regardless of We also acknowledge Dr. J.L. Cervera, editor of this special issue, their small size (as compared with those ofother aeolidaceans). and two anonymous referees, whose comments and criticism helped Examination offish egg membranes after consumption seems to toimprovethecurrentversionofthepaper. agree with this view. Moreover, there is a reduced radula size differencebetween animals havinga markedlydifferent body length, REFERENCES making ussuspect that theentire "adult-like" radular morphology is Evans T.J., 1922. Calmaglancoides'. a study in adaptation. QuarterlyJournalof developed early in the ontogenetic process. In fact we have observed MicroscopicalScience,London, 66:439-455. smalleranimalsfeedinginamannersimilartolargerones. HughesR.N. &Emerson W. K., 1987. Anatomical and taxonomiccharacteristics Therefore, one can assume that juveniles already feed on fish ofHarpaandMorum(Neogastropoda:Harpaidae)TheVeliger,Berkeley,29: eggs,contrarilyrothepreviousassumption that juvenilesaregeneral .349-358. carnivores (Evans, 1922). Thus, there seems to be no food size RowettH.G.Q., 1946. Acomparison oftliefeeding mechanismsofCalmaglaucoides constraint. and Nebaliopsistipica.JounialoftheMarine BiologicalAssociationofthe United Terminal larger teeth seem to continue differentiating, from the Kingdom,Plymouth,26:352-357. juvenile phase onwards, making us believe that they may also be useful for the adult’s feeding behaviour. EvANS (1922) suggested that these teeth might be used only in tlie post-settlement phase, beforejuvenilescould findfishspawn. Lavoroaccettalo7febbraio2000 > 180<

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