[artigo] gomes et al [1998] distribuição adaptação anemona pernambuco

8/3/2019 [Artigo] GOMES et al [1998] Distribuio Adaptao Anemona Pernambuco

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Dis t r ibut io n, a bund a nc e a n da d a p t a t io ns o f thr ee s pec ies o fA c t in i id a e ( C n id a r ia , A c t in ia r ia )on an intert idal beach rock inCam eiro s beach, Pernambu co, BrazilP. B. Gomes, M. J. Belem & E. Schlenz

Gomes, P. B., Bel6m, M. J. & Schlenz, E., 1998. Distribution, abundance and adaptations of threespecies of Actiniidae (Cnidaria, Actiniaria) on an intertidal beach rock in Carneiros beach, Pernambuco,Brazil .Misc. Zoo!., 21.2: 65-72.Distribution, abun dance and adaptations of three species of Actiniidae (Cnidaria, Actiniaria) on anintertidal beach rock in Carneiros beach, Pernambuco, Brazil- Bunodosoma cangicum Correa in Bel6m &Preslercravo, 1973; Actinia bermudensis (McMurrich, 1889) and Anthopleura krebsi Duchassaing &Michelotti, 1860 were studied in an intertidal beach rock at Carneiro's beach, Pernambuco, Brazil.According to the environmental factors five microhabitats occupied by the species were identified.Random surveys were made and a framework w as used to calculate the mean specific density of eachspecies in the different habitats. Ten specimens of each species were measured. The Mann-Whitney testwas used to compare size and density of B. cangicum from different habitats. Specimens of B. cangicumwere found in three different habitats, with the lowest density (1.1 ind/m2) and the lowest size(1.2x3.8 cm, diam x height) in poolson the flats of thebeach rock, exposed to the sun and to the rainfall.In this habitat, there was an increase in the temperature and salinity of water during dry seasons and thespecies was almost absent during rainy seasons. In habitats 4 and 5, which were more protected, themean density was 6.0 and 6.4 and the mean size of 4.5x6.3 cm and 5.3x6.4 cm, respectively. A. krebsiwas present in two habitats with high density (24.5 ind/m 2) in the habitat with longest air exposure timeand this density seems to be associated with asexual reproduction. A. bermudensis was present in onlyone habitat with low density (1.4 ind/m2) and size of 2.0x2.2 cm. No differences were observed in thestatistical comparison of the density of B. cangicum in two protected zones (P> 0.05) which were thusused as one and com pared with another unprotected zone. The results showed significant differences(P < 0.05). The comparison of size showed differences among the three zones (P < 0.05).Keyw ords: Actiniaria, Ecology, Pernambuco, Brazil.(Rebut: 74 X 97; Acceptacio condicional: 27IV 98; Acc. definitiva: 1 XII98)Paula Braga Gomes, Laboratorio de Biologia de Cnidarios, Depto. Cs. Mar., Fac. Cs. Ex. y Nat., UNMdp Funes3250-7600, Mar del Plata, Argentina (Argentine).- Maria Julia da Costa Belem, Retired Professor/UFR], CaixaPostal 24.030, CEP.20.522-970, Rio de Janeiro-RJ, Brasil (Brazil).- Erika Schlenz, Depto. deZoologia, Inst,de Biociencias, USP. Caixa Postal 11.461, CEP.05422-970, Sa o Paulo-SR Brasil (Brazil).

ProfaOepto. de

Fernanda Ama-al- Area He ZoMo?iU F R P E

ISSN: 0211-6529 1998 Museu de Zoologia


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I n t r od u c t i onBenthic organisms l ike sea-anemones are sub-mitted to environmental factors that influenceor determine their distribution, such as thetype of substrate and competitivity for it. Thefluctuation of tides is also important becauseit influences food availabil ity, garnets and lar-val dispersion and some adaptation is requiredto resist or avoid wave and current action( W A I N W R I G H & K O E HL , 1 9 7 6 ) . Several studies haveevaluated the actions of these factors. STOTZ( 1 9 7 9 ) analysed the zonation of three speciesof sea-anemones correlating with their mor-pholog ical adaptat ions. HA R R I S ( 1 9 9 1 ) consid-ered that water flow is a factor of speciationin some populations of Metridium senileLinneus, 1767. These studies were intensifiedon organisms of the intertidal zone, whichpresent higher stress due to desiccation thanthose that are submerged (BARNES etal . , 1 9 6 3 ) .The intertidal zone is a special place becausethe intertidal organisms need to develop mor-phological, physiological and behavioural ad-aptations to survive in this environment. Theprincipal adaptations developed by intertidalsea-anemones are related to: 1. Reproductivebiology with different reproductive patterns(SHICK et al, 1 9 7 9 ) ; 2 . Different strategies forc o l o n i za t i o n ( S E B E N S , 1 9 8 2 ) ; 3 . Intra andinterspecific a ggression ge nerally associated tothe presence of marginal spherulae (BIGGER,1 9 8 0 ; B R A C E, 1 9 8 1 ) ; 4 . The search for newhabitats by displacem ent capacity (FUJII, 1985;W I L L I A M S , 1 9 9 2 ) an d 5. Several mechanisms todecrease the wave and desiccation action(OTTAWAY, 1973; ZAMPONI, 1981 ).The present paper examines how envi -ronmental condi t ions inf luence the d istribu-tion of three intertidal species of sea-anemo-nes f rom a beach rock at Carnei ros beach(State of Pernambuco, Brazi l ) and the adap-tat ions developed by each spec ies in d i f fer-ent m icrohabi tats.

S t u d y a r e aThe area studied was an intertidal beach rock(sensu GUILCHER, 1985) at Carneiros beach(845' S 3505' W), State of Pernambuco, Bra-zi l . It was 46 m long, 11 m wide and 0.60 mhigh (maximum height) . It was submerged a l lyear round during h igh t ide. When uncov-

ered during low t ide, f ive zones were ident i -f ied, according to envi ronmental condi t ions,forming d i f ferent microhabi tats. These zonesdid not represent the total area of the beachrock but the more conspicuous habi tats oc-cupied by sea-anemones.The study area was located on the southcoast of the State of Pernambuco, which haswarm humid weather wi th temperatures vary-ing from 2 5 C to 3 0 C with an average about

2 6 C; tw o defined seasons throughout the year,one dry (September-February) and one rainy(March-August) and a high mean annual rain-fal l of about 2 , 5 0 0 mm. The water temperaturevar i ed f rom 25C to 32C r e a c h i n g 34C i n sum-mer in the shallow rock pool in the flats ofbeach rock, and salinity fluctuated in the ap-proximate range of 36.4%o - 37%o.

M a t e r i a l a n d m e t h o d sOn the basis of previous work ( G O M E S & M A Y A L ,1997) and according to observations of envi-ronmental factors, f ive zones were identifiedduring low t ides represe nt ing d i f fer entmicrohabi tats on the beach rock. The char-acteri st i cs of each microhabi tat were ana-lysed and correlated with the species presentsand thei r s i ze and densi ty .

Three spec ies were studied: Bunodosomacangicum Correa in Belem & Preslercravo,1973; Anthopleura krebsi Duchassaing &Michelotti , 1860 and Actinia bermudensis(McMurrich, 1889).

Random surveys were carried out in thestudy area on the fol lowing dates: 3 V II I 93,18 IX 93, 14 X 93 and 12 I 94. The tem pera-ture and sa l in i ty of water were measured inAugust 93 ( ra iny season) and Janua ry 94 (dryseason) . The fol lowing measurements weremade at each t ime point and in each of thefive microhabitats:

1. The d iame ter just above the p edal d i skand the height of ten spec imens of eachspec ies was measured. The spec imens wererandomly se lected among sea anemones wi thcommon aspect , avoid ing spec imens con-tracted or wi th d istended column.2. A f ramework (50x50 cm) was randomlyposi t ioned ten t imes and a l l sea anemonespresent were counted.For each species in each microhabitat, thespecific density (De) was calculated, accor d-


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ing to O D U M (1959) . Only the mean spec i f i cdensi ty i s presented since the seasonal f luc-tuat ions are usual ly caused by varied repro-duct ive patterns or d i f ferent mechanisms ofsettlement of larvas and juveniles.Finally, a Mann-Whitney test (SOKAL & ROHLF,1981) was appl ied w i th a s igni f i cant leve l of0.05 to compare the s i ze and densi ty of B.

cangicum in d i f ferent microhabi tats at eacht ime point .This was done to determine whether therewere d i f ferences between the protected andunprotected zone. B. cangicum was the onlyspecies used because of its distribution inthree habi tats. A. krebsi was present in twohabi tats but the densi ty and si ze of th i s spe-

cies is influenced by its capacity for asexualreproduct ion. For the comparison of s i ze ,only d iameter was used since i t i s more sta-ble than the height of the sea-anem one.

R e s u l t sA c c o rd i n g t o e n v i ro n m e n t a l c o n d i t i o n s ,f ive d i f ferent microhabi tats occupied by seaa n e m o n e s w e re i d e n t i f i e d o n t h e b e a c hrock.Habi tat 1Characteri st i cs: smal l caverns in the lateralwal l on the sea s ide.Wave act ion: very strong during the ri seof the t ide, but the caverns provided protec-t ion for sea-anemones.

Time of a i r exposure: short , as i t was thef i rst sector of the beach rock covered bywater.Species present: A. bermudensis an d A.krebsi; both were randomly d istributed withlow specific density (De = 1.4 ind/m 2 an dDe = 3.0 ind/m 2, respectively).Observat ions: spec ies showed no prefer-ence for d i f ferent he ights on the wal l .Habi tat 2Characterist i cs: shal low rock pools formed atlow t ide in the f lats of the beach rock.Wave act ion: there was no di rect waveact ion. During the ri se of t ide there was noimpact of waves in the area.Time of a i r exposure: th i s habi tat was ex-

posed for two or three hours, but sea-a n e m o n e s w e re p e rm a n e n t l y s u b m e rg e d i nthe pools, not suf fering desiccat ion. Thesepools had the temperature and sa l in i ty ofwater af fected by exposure to the sun andrainfa l l , during low t ides.Species present: B. cangicum (De = 1.1 ind/m2).Observat ions: during ra iny seasons themean temperature in these pools was 3 2 Cw h e n s e a w a t e r w a s 2 9 C . During dry sea-sons there was an increase in water tempera-ture ( 3 4 C in pools and 3 2 C in sea-water)with a sl ight increase in sal inity in the poolsf ro m 3 6 . 7 % o to 3 6 . 9 % o . The number of sea-anemones in th is habi tat decreased duringthe ra iny season.

Habi tat 3Characteristics: horizontal crevices on the lateralwal l on the shore s ide.Wave action: no wa ve action in this habitat.Time of a i r exposure: very prolonged; th iswas the last sector to be submerged.Species present: A. krebsi (De = 24.5 ind/m 2).Observations: density of this anemone washigh, forming a gregar ious populat ion inthe crev ices wi th 0 . 4 0 - 0 . 5 0 m h i g h .Habi tat 4Characteristics: great crevice of 0.60 m d e e pperpendicular to the ocean. The sea-anemo-nes were low on the lateral wal l s .Wave action: very l ittle; there was a smallchannel at low t ide and ri s ing t ide.Time of a i r exposure: indiv iduals locatedat the bottom of the crev ice were covered bythe water of the smal l channel . Other indi -v iduals located on the lateral wal l s of thecrev ice suf fered varied degrees of desicca-t ion according to thei r he ight on the wal l .

Species present: B. cangicum (De = 6.0 ind/m2).Observat ions: some anemones on the lowsector of the lateral wal l extended thei r ten-tacular c rown towards the water channel .Habi tat 5Characteristics: the low sector of the verticalwal l on the shore s ide near remnant wateron the bottom.

Wave act ion: during the ri s ing t ide, asmal l current was formed due to water en-


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tering on both s ides of the wal l .Time of air exposure: moderate; it was thelast side reached by the water, but sea-anem o-nes, on the bottom, recovered quick ly .Species present: B. cangicum (De = 6.4 ind/m2).Observat ions: some spec imens presenteda distended colum n with the oral d i sk in theremnant water on the bottom.

The annual mean si ze and annual mean spe-cific density of al l species in each micro-habi tat are show n in table 1. Table 2 showsthe mean densi ty and si ze (only d iameter) ofB. cangicum f rom protected (microhabi tats 4and 5) and unprotected (microhabi tat 2)zones at each t ime point . The resul ts of theMann-Whitney test showed no di f ferencesin densi t ies between microhabi tats 4 and 5(P > 0.05 in a l l t ime points) . These two habi -tats were thus used as one protected zoneand compared with microhabi tat 2. At a l lt ime points there were d i f ferences betweenthe two zones (for al l U calculated, P < 0.05).

Table 1. Mean size (diameter of columnjust above the basal d i sk x he ight ofcolumn) and mean specific density (De)of sea-anemones in the f ive habi tats.Tamaho medio (diametro de lacolumna por sobre el disco basal x

altura de la columna) y densidad mediaespecifica (De) de las anemonas de maren los cinco habitats .

Comparison of the anemone si ze showedthat the three m icrohabi tats are stat i st i ca l lyd i f f e re n t ( P< 0 . 0 5 ) .

D i s c u s s i onThe density of al l species fluctuated through-out the year, but no uni formity was ob-served. This f luctuat ion could be caused bydi f ferences in the reproduct ive p atterns ofeach spec ies or in thei r d i splacement capac-i ty in response to unfavourable envi ronmen-tal conditions, as discussed by F U J I I ( 1 9 8 5 ) .

The species B. cangicum was a lmost ab-sent in microhabi tat 2 during the winter. Inthis habi tat the indiv iduals were exposed torainfa l l which d i rect ly af fected the tem pera-ture and salinity.

Wave act ion and desiccat ion caused byai r exposure seem to be the most importantfactors characterizing each microhabitat. Thegreat d iversi ty of microhabi tats probablymeets the needs of a l l species, thus keepin gthem separate. Only microhabi tat 1 wasshared by two spec ies that were randomlydistributed at low densi t ies.B. cangicum occupied a h igh number ofdi f ferent microhabi tats. This could be inter-preted as indicat ing th is spec ies adapts bet-ter than others. One of these adaptat ions i si ts aggressive capac i ty due to the large s i zeand the presence of marg inal spherulae. Forthe same species B E L E M & PR E S LE R C R A V O ( 1 9 7 3 )observed aggressive behaviour in spec imenskept in aquariums. Fu rthermore, th i s spec ies

needs a mechanism to resi st the ri se of tem-perature and sa l in i ty during summer in themicrohabi tat 2. In th is habi tat the v i rtualabsence of B. cangicum in the winter couldbe interpreted as indicat ing movements ofindiv iduals into less exposed microhabi tatsin response to the decrease of sa l in i ty ormortal i ty of anemones in more exposed ar-eas. Microhabi tats 4 and 5 were more pro-tected f rom wind and sun but the anemoneswere exposed to a i r for some t ime andneeded ef f i c ient mechanisms to attenuatethe ef fects of desiccat ion. During low t idessome spec imens acqui red a dome shape,which a l lowed them to keep water in thecoelenteron and decreased thei r exposedarea. This is important due to their large sizeand coinc ides wi th f indings by D A Y T O N ( 1 9 7 1 ) .

Species Mea n size DeHabitats (cm) (ind/m2)B. cangicum

2 2.2x3.8 1.14 4.5x6.3 6.05 5.3x6.4 6.4

A. krebsi1 0.9x1.1 3.03 1.3x1.4 24.5

A. bermudensis1 2.0x2.2 1.4


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Table 2. Mean densi ty (De, ind/m 2 ) and mean si ze (Ms, cm )(x. Mean; s.d . Standarddesv iat ion) of B. cangicum in protected and unprotected zones at each t ime point .Resul ts of Mann-Whitney test for the comparison of the densi t ies of B.cangicum f romhabi tats 4 and 5, and (4+5) and 2 (U and i ts probabi l i ty) .Densidad media (De, ind/m2> ytalla media (Ms, cm) (x Media; s.d. Desviacion estandar)de B. cangicum en zona protegida ydesprotegida en cada muestreo. Resultad os del testde Mann-Whitney para la comparacion de las densidades de B. cang icum provenientesde los habitats 4y 5 y (4+5) y 2 (valores de U y sus probabilidades).

Protected zone Unprotected zone Habi tats(Habi tats 4 and 5) (Habi tat 2) 4x5 (4+5)x2Dates De Ms (x s.d.) De Ms (x s.d. ) U(p) U(p)3 VIII 93 7.1 4.720.60 0.7 1.980.13 20 P>0.05 0 P0.05 1.5 P0.05 8 P0.05 7 P


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70 Gomes et a l .

i ts great capac i ty for asexual reproduct ionby long i tudinal f i ss ion described by B E LE M &P I NT O ( 1 9 9 0 ) and observed in some indiv idu-als in the study area. It seems that in thisreg ion A. krebsi used a reproduct ive strat-egy based on asexual reproduct ion, prob-ably due to the stress condi t ions to which i ti s submitted. E X C O F F O N & Z A M P O N I ( 1 9 9 5 ) ob -served the same pattern in the a nemo neTricnidactis errans Pires, 1988 in Mar del Plata,Argent ine. A detai led study of the repro-duct ion strateg ies of th i s spec ies i s neededto c lari fy th i s quest ion.

The presence of marg inal spherulae in Akrebsi indicates an aggressive capacity, prob-ably lower than that presented by B. cangicumdue to the d i f ference in s i ze between them.This capac i ty could contribute to the nu-merical superiori ty of th i s spec ies compare dwith others and a l so to i ts i solat ion, wi thapparent ly c lonal behaviour.

Other studies on spec ies f rom the genusAnthopleura ( F R AN C I S , 1 9 7 3 ; S M I T H & PO T T S ,1987) demonstra ted that c lonal behaviour i svery common in th is genus.In both microhabi tats occupied by A.krebsi, this species is in protected zones suchas small caverns (microha bitat 1) or in crevices(microhabi tat 3) . The smal l s i ze and the con-traction capacity al low this species to occupyvery protected zones and a l so decrease theamount of body surface exposed to desicca-tion. In spite of this, individua ls, special ly inmicrohabi tat 3, were exposed to a very longperiod to a i r and needed some adaptat ions.In the column there are numerous verrucaesto which particles, such as pieces of shellsand calcareous a lgae adhere, thereby form-ing a protect ive layer which protects agains tdesiccation.A. bermudensis was only observed inm i c ro h a b i t a t 1 a n d f e w i n d i v i d u a ls w e refound, suggest ing th is spec ies i s not wel ladapted to th is envi ronment. In fact , th i sdark red anem one c learly absorbs solar heatand i ts smoo th column favours desiccat ion.The only adaptat ions presented by th is spe-c ies are the domeshape during low t ideand i ts locat ion in a microhabi tat quick lycovered by water, thus resi st ing the waveimpact and being in protected zones suchas small caverns.The resul ts of the present study conf i rmthe need for sea anemones in the intert idal

zone to develop mechanisms which a l lowthem to occupy microhabi tats on the basisof the i r ecolog ical preference and the envi -ronmental condi t ions.

R e s u m e nDistribucion, abundancia y adaptaciones dealguno s Actiniidae (Cnida ria, Actiniaria) enun arrecife interm areal de la playa de losCarneiros, Pernambuco, BrasilEn este trabajo fue analizada la distribucion yabundanc ia de Bunodosoma cangicum Correain Belem & Preslercravo, 1973; Anthopleurakrebsi Duchass aing & Michelotti , 1860 yActinia bermudensis (McMurrich, 1889) en unarrecife de arenito intermareal de la playa delos Carneiros, Pernambuco, Brasi l . Segun lascondiciones ambientales fueron identificadosc i n c o m i c ro h a b i t a t s d i s t i n t o s d o n d e s eubicaban las especies, con distintos tiemposde exposicion al aire y accion de las olas.

A lo largo de un ano se real i zaron censoscon la uti l izacion de un cuadrado de 50x50 cmpara el calculo de la densidad especifica me-dia de cada espec ie en los d i ferentesmicrohabitats. In situ se midio la tal la (diame-tro y altura) de diez ejemplares de cada espe-cie en cada habitat (tabla 1).La especie B. cangicum estuvo presenteen tres habi tats d i st intos con menor densi -dad (1,1 ind/m 2) y menor ta l la (2,2x3,8 cm,diam x a l t ) en e l habi tat 2, formado porpozas en la p lataforma superior de l arrec i fey el mas expuesto a la accion del sol y de lal luv ia . En estas pozas hubo un incrementode la temperatura y sa l in idad del agua du-rante e l verano y una casi ausenc ia de laespecie en el periodo l luvioso. En los habitats4 y 5, mas proteg idos, tuvo densidad mediade 6,0 y 6,4 ind/m 2 y ta l las medias de 4,5x6,3cm y 5,3x6,4 cm, respectivamente.A. krebsi se presento en dos habitats dife-rentes con ta l la pequena y densidad baja encavernas ubicadas en la pared del arrec i feque miraba al mar (0,9x1,1 cm y 3,0 ind/m 2) ygran densidad en e l ambiente con mayort iempo de exposic ion a l a i re , donde estadensidad parece estar re lac ionada con la ca-pac idad de reproducc ion asexuada presen-tada por d icha espec ie .A bermudensis estuvo presente en un solo


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Miscel lania Zoolog ica 21.2 (1998) 71

habi tat com pe quena densidad (1,4 ind/m 2) yta l la media de 2,0x2,2 cm.El test de Mann -Whitney fue ut i l i zadopara comparar las ta l las y densidades de laespecies B. cangicum en los distintos habitats(tabla 2). La densidad de esta especie en lasdos zonas proteg idas no mostro n ingunadi ferenc ia s igni f i cat iva (P > 0,05) . Estas doszo n a s f u e ro n c o m p a ra d a s c o n o t ra zo n adesproteg ida y se observaron di ferenc ias s ig -n i f ic a t iv a s ( P< 0 ,0 5 ) .Con relacion a la tal la, las tres zonas pre-sentaron di ferenc ias entre s i (P < 0,05) . Seobservo que la d i stribuc ion y densidad delas especies estan relacionadas principalmen-te con la incidencia de las olas y el tiempo

de exposicion al aire. Se discuten las adapta-c iones morfolog icas, f i s io log icas y compor-tamentales de las especies segun su distribu-cion en los distintos habitats.

A c k n o w l e d g e m e n t sW e w a n t t o t h a n k t o D r . M a u r i c i o O .Zamponi for h i s suggest ions and rev is ion ofthe paper and Prof . Marie l Amato for therevis ion of the Eng l i sh version.

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krebsi Duchassaing & Michelott i, 1860(Cnidaria, Anthozoa, A ctiniidae), a new recordin Brazil. An. Acad. Bras. Ci., 6 2 ( 2 ) : 1 8 3 - 1 9 2 .B E L M , M . J . & P R E S L E RC R A V O , J . C . , 1 9 7 3 .C o n t r i b u t e s a o co n h e ci m e n to d a f a u n ade Cnidarios do Espi ri to Santo, Brasi l . I .Considera^oes sobre Act in iaria do Muni -c ip io de Aracruz, ES. Bol. Mus. Biol. Prof.Mello Leitao, S. Zoologia, 8 0 : 1 - 1 4 .BIGGER, C. H.( 1980. Interspecific and intras-

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