Biotemas, 24 (2): 37-46, junho de 2011 doi: 10.5007/2175-7925.2011v24n2p3737 ISSNe 2175-7925

Reef bioconstructors of the Rocas Atoll, Equatorial South Atlantic

Marcelo de Oliveira Soares1* Carlos Augusto Oliveira de Meirelles2 Valesca Brasil Lemos3

1Instituto de Ciências do Mar – Labomar, Universidade Federal do Av. da Abolição, 3207, CEP 60165-081, Fortaleza – CE, Brasil 2Departamento de Biologia, Centro de Ciências, Universidade Federal do Ceará Av. Mister Hull, s/n, CEP 60455-760, Fortaleza – CE, Brasil 3 !" $%&&'$(%&$)$&&)" *Autor para correspondência [email protected]

Submetido em 30/11/2010 Aceito para publicação 02/04/2011

Resumo Bioconstrutores recifais do Atol das Rocas, Atlântico Sul Equatorial. Este estudo apresenta uma análise dos bioconstrutores recifais do Atol das Rocas, Nordeste do Brasil, amostrados em janeiro e fevereiro de 2008. Os resultados demonstraram a presença de uma comunidade constituída principalmente por algas calcárias (Porolithon pachydermum, Sporolithon sp., Lithoporella sp., Lithophyllum sp.), corais (Siderastrea stellata, Favia gravida), moluscos vermetídeos (Dendropoma irregulare, Petaloconchus varians) e foraminíferos (Homotrema rubrum) na formação do recife. Este levantamento taxonômico demonstra a importância de diferentes espécies :>! Palavras-chave: Algas calcárias, Corais escleractíneos, Foraminíferos, Moluscos vermetídeos, Recifes

Abstract This study presents an analysis of reef bioconstructors at the Rocas Atoll, ’s Northeast, sampled in January and February 2008. The results showed the presence of a community made up mainly of calcareous algae (Porolithon pachydermum, Sporolithon sp., Lithoporella sp., Lithophyllum sp.), corals (Siderastrea stellata, Favia gravida), vermetid mollusks (Dendropoma irregulare, Petaloconchus varians), and foraminifers (Homotrema rubrum) in the reef formation. This taxonomic survey shows the importance of a different species during the :?GKGQG>G! Key words: 'KKSV

Revista Biotemas, 24 (2), junho de 2011 38 M. O. Soares et al.

Introduction archipelago, in the Brazilian YGXY($%$[‡‡'\[`I] ":K ($$‰{!GKGZ&th century, with the sedimentary deposits, accretions or accumulations effective implementation of research activities at REBIO by organic processes and it often involves active (Biological Reserve), the Atoll has been studied more :\!: K>Q?GK GKKG geology, corals, foraminifers, and calcareous algae elevations or reefs from materials produced internally XŠ'\][]‹($$q[]!($$[ (e.g., deposition of biogenic carbonate), removed ‡‡'\[`I]($$‰[Š[($$‰[ from other materials (e.g., organic cementation) or \["]Y'($$$[Z&&([Š` :K:GG! !Z&&Z[]!Z&&$[Z&(&[ bioconstructions are short-term phenomena, due to al., 2010). nonlinear growth cycles or decay standards, while other bioconstructions suffer secondary diagenesis Despite these studies, a series of geobiological GKXY($$Z[\[ aspects (like the bioconstructors species) of the Rocas "]Y'Z&&([Y^`]!,Z&&Z['\`[ Atoll remain poorly studied, despite its importance and \]YZ&&q{! >GGQKG> GX‡‡'\[`I]($$‰[] There are basically three ways of bioconstructive !Z&&${!‚GQGK: XY^`]Z&&%{|({?GG towards the taxonomic survey of the bioconstructors themselves produce biomineralized material, such species in the Rocas Atoll (NE Brazil). as secretion from carbonate skeletons by vermetid V:GK[Z{?G G:QG}K Material and Methods particulate matter, as in the case of sabellarid polychaetes that build tubes with beach sand through the production Study area K[~{?G :V\K! The atoll has a volcanic origin and carbonate formation in the reef, with a slightly ellipsoidal structure, :K`?X($%&{ }X)†{?GG: “[...] the product of constructive and aggregating/ ~!‰VG}XY){GZ!%V cementing activity of biotic constituents that, due to X‡‡'\Z&&Z{!‚GQ:Q‡VG` their potential to resist the waves, erect topographic X($$‰{:GK structures resistant to them.” According to Cloud submarine mount of the guyot type. This ring is made X($%${QK}QG up, from the morphological point of view, of layered of its chain of nutrients, the breakdown product, and terraces that dam the waters during the low tides. These GQX($${!‚G waters form a system of pools (with the maximum construction is therefore the result of organic activity in depth of 5m), leaving only two islands visible. In the response to environmental factors (hydrophysical and northeastern portion of the ring, we notice an area that is hydrochemical) that are incorporated to the geological always submerged and it gives origin to the reef lagoon. X'\`[\]YZ&&q{! The ocean waters have access to the reef interior through The Rocas Atoll is located at the top of a chain the canals (reef pass) (Figure 1). K:GG] whose base is located at a depth of 4,000 m in the :~ %(R`~~ q$R†` 266 km from the city of Natal and 150 km west of the

Revista Biotemas, 24 (2), junho de 2011 Reef bioconstructors of the Rocas Atoll 39

(| ‚GX>G{G?G???|` •`[•\['•Q[•K[•KK!:K activities during the low tide.

Field and laboratory The geological samples were obtained from KGGG'" The collections were made at REBIO (Biological various geoenvironmental compartments of the Rocas Reserve), at the Rocas Atoll, from January 30 to February Atoll. The samples were described macroscopically 28, 2008. The Atoll was accessed by lifebuoy through and microscopically, photographed and parts of them GGY†KGKG were selected for taxonomic description of the main high tide. bioconstructors. The approach to the general aspect of the Rocas ]G?V >KKGK q‘KGQK! map images from the area and visualization both of in situ The bioconstructors were identified through the characteristics of the bioconstructors and the occurrence comparison with samples from other reefs in Brazil’s zones in the reef. The observation was performed during Northeast which belong to the Instituto de Ciências do walks at the top of the reef, which is exposed in the low Mar and the Laboratório de Invertebrados Marinhos, XK)'"{G :GK'M'[G submersed parts. The data obtained in these description KG ?|{GKG –"GKG :[:{G:! M! These data were completed with detailed photographic ‚G?V:QGX($‰({“X($”q{` X($”{ documentation and samplings of the main constructors YX($”‰{‡VGXZ&&Z{?! of the reef structure.

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‚G:Q†QX($‰‰{†V The corals that compose the reefs described X($””{``XZ&&&{\XZ&&Z{? in this work differ from the Caribbean reefs, in KGKG which the latter are largely composed of Acropora, of calcareous algae. Montastraea, Diploria and Porites. They also differ from the southern Abrolhos reef complex, which is mainly built by Mussismilia braziliensis S(”” Results and Discussion (`I]et al.,($””{!‚?K): ?|Siderastrea stellata The reef geology of the Rocas Atoll is a carbonate S(””X~{Favia gravidaS deposit that resulted from the bioconstructor activity 1868 (‡‡'\, 2002). of benthonic organisms in response to environmental factors like availability of light, sea hydrodynamics, Encrusting foraminiferans (homotrematids and subsidence, and eustatic variations. Paleontologically, acervulinids) are concentrated in the reef-ridge and the the taphocoenosis is dominantly made up of calcareous K)\::QKK algae, vermetid mollusks (Figure 2B), foraminifers by sedimentation, or maybe it is not high enough. and only secondarily by corals (Figure 2D). This fact X‡‡'\Z&&Z{:Q Systematic paleontology accepted that coralline algae lack potential to erect or to be primary constructors of reefs in the neogene Siderastrea stellata Verrill, 1868 (Figure 3F) (Figure 2A). Description|'?GQ From the paleobiological point of view, coralline spheric in outline. Live colonies which live in very reefs, like the Rocas Atoll, are formations created by shallow waters have a reddish color, are usually small the action of several types of organisms like corals, (diameter from 5 to 10 mm), have calyxes with diameter calcareous algae, vermetid mollusks, foraminifers, from 2 to 3 mm, roundish, deep and the fourth cycle of GXZ'{!‡VG` X($$‰{ septa is incomplete. ‡VGXZ&&Z{:GG Remarks|‚GG" KGGVK> and in the oceanic islands in the Brazilian Northeast and that there are circumstances when coralline algae and KGG!S. stellata is an endemic coral, associated encrusting organisms behave not only as ?Q:G"XYS[ binders but, also, as reef builders. In the studied area, `SZ&&~{!Q':: coralline algae are the dominant reef-builders between species, but with bigger calyxes, septa more delicate 0 and 15 m depth (Figure 2E). and wider interseptal spaces. This coral displays a In reefs of the Caribbean in the , morphological variation that distinguishes colonies the corals correspond to the greater volumetric part of of shallow waters from those of deep waters (deeper depositions of the holocenic section, however, one notices G%{X`"]`($$($$:[`I] :K|Halimeda ($”[!($$Z[\‚‹`['‚]($$q[ sp., fragments of mollusks and incrusting foraminifers ($$%($$‰['\S[($$‰[ X'\`[\]YZ&&q{!' ‡‡'\[`I]($$‰[‡‡'\Z&&Z[Y‚] common in the description of carbonate sedimentary !Z&&q[Š`!Z&&‰[YS! facies and in the bioconstruction of reef environments. 2010). Modern reefs have a thickness of over 10 m and many are built primarily by calcareous algae (]‚‚ŠY($~[ ‡‡'\[`I]($$‰[‡‡'\Z&&Z[] et al.,Z&&$).

Revista Biotemas, 24 (2), junho de 2011 Reef bioconstructors of the Rocas Atoll 41

Z| X{ :G Q ! ‚G ) : K G }KGVKG[X"{SG?G)QXS{! Presence of Porolithon pachydermumX($&${KGX{Q—:X'{[ (C) Polytypic subfossiliferous assembly formed by scleractinian corals (Co), bentonic foraminifers (arrows), red coralline X{Q—::Q:X'{[X{KK:Q G:[X{KG:G?GX{ :Q?[X{"X{KGXDendropoma irregulare Orbigny, 1842) on top of :X{!GKG:KGK of the variations of the sea level due to being in intertidal or immediately subtidal zones.

Revista Biotemas, 24 (2), junho de 2011 42 M. O. Soares et al.

~| " G K G | X{ ' : : XSporolithon sp., Lithophyllum, Lithoporella!{??[X"{"?GG!" )Š[ X'{ K X??{ Q : :Q Porolithon pachydermum ($&$!‚GGKKGGG'::)[ (D) Macroscopic aspect of several bioaccumulations (globes) of foraminifera Homotrema rubrum (Lamarck, 1816) in the [X{KKGKKGG?GK:!S bioconstructor Petaloconchus varians (arrow) is indicator of intertidal or immediate sub-tidal environment, Central part with X{!ŠK?V|'!':KK)')Š[X{ Siderastrea stellataKG>QKKG†G!

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Favia gravida Verrill, 1868 plane to protuberant surface, smooth texture and strongly G!ŠQX($$‰{G?GG Description|‚GGG extensive crustose coralline algal pavements studied hemispherical colonies, bearing a light brown or G?GK!‚G Q?G!'Q}>G pavements consist mainly of Porolithon spp. long, often deformed. Due to the great variation on shape KGQ}`:X($${:GKK Lithoporella sp. (Figure 3A) shapes, types 1, 2 and 3, each one of them with three morphological variations, named A, B and C, which Description: Plants are epigenous with variate according to the depth where the polyps live. pseudoparenchymatous thalli, thin applanate crustose thalli, dorsiventral and monomerous. Primigenous Remarks: This species is very resistant to environmental KQK variations, particularly regarding the temperature, cells. Conceptacles were not found in the inspected salinity and water turbidity. Together with Siderastrea specimens. stellata, they are the most common corals in shallow pool tides on the emersed top of the coastal reefs and Remarks|‡VG` X($$‰{G even on non-reefy substrata along the beaches. It is an Lithoporella is one of the main calcareous algae in reefs endemic species of the Brazilian fauna in neogene reefs K:G"RYG!G" X`"]`($$($$:[`I]($”[ X($$${GG::Q? !($$Z[\‚‹`['‚]($$q['\S[ G!†VX($””{ ($$‰[‡‡'\[`I]($$‰[Š`[ a combination of diagnostic features that differentiate Š]Z&&‰{! Lithoporella from other genera of Mastophoroidae as K?|{GQGVG[:{ Porolithon pachydermum (Foslie) Foslie, 1909 K[{ (Figures 2E and 3C) VK}K! Description: Plants are epigenous with Lithophyllum sp. (Figure 3A) pseudoparenchymatous thalli, applanate crustose to protuberant thalli. Crustose portion of the plant Description: Plants epigenous, pseudoparen- dorsiventral, monomerous composed of a noncoaxial chymatous thalli, varying in form from applanate to G?G columnar thalli. Columnar plants are monomerous with curve upwards from the core towards the thallus surface. K}} It has typical shapes of crest and slots observed in the G?GK !S)". (2005) comment on cellular or their derivates curve outwards towards the thallus characteristics that are diagnoses of the species as single- surface. pore conceptacles, trichocysts in horizontal groups, Remarks: †V':X($$Z{K perithalial cells connected by fusion and hypothalial cells dimerous thalli in crustose portions of Lithophyllum arranged horizontally parallel to the substrate. KG!G" Remarks: The species is common in the Caribbean, X($$${KGGKG mainly in the reefs of the Bahamas and Antilles. In the Rocas Atoll. The genus Lithophyllum is restricted to the Northeastern region of Brazil, it is very common in the lower intertidal and uppermost subtidal in the study area. KK:GGX\[ "]Y'($$${!GG:?GK>Q Sporolithon sp. (Figure 3A) bored by polychaetes and associated with the Homotrema Description: Plants epigenous, pseudoparen- rubrum foraminifer. It is found in all the reef habitats chymatous thalli, with applanate crustose thalli described. Color varies from light brown to dark purple, sometimes forming spherical thalli. Monomerous in

Revista Biotemas, 24 (2), junho de 2011 44 M. O. Soares et al.

!'K}?G G"G[G distal portions curve outwards towards thallus surface. YG:GK‚X] Lacks palisade cells. ($$q[]!Z&(&{! Remarks: It belongs to the Melobesioideae subfamily. Petaloconchus varians (Orbigny, 1841) (Figure ‡VG` X($$‰{GSporolithon sp. is 3E) one of the main calcareous algae of the reefs of Abrolhos, "RYG!†VX($””{ Description|SKVK account on the historical controversy that surrounded X(&&GZ{!SQ the application of the names Archaeolithothamnium variable growing in all directions. Color dark brown, GX(”$({Archaeolithothamnium Rothpletz G:?!?G exX(”$”{GSporolithon. ribs sometimes reticulate. Operculum concave, translucent and multispiral. Aperture circular. Homotrema rubrum (Lamarck, 1816) (Figure 3D) V|‚G"† Description: Homotrema is a calcareous encrusting Indies and Brazil along the coast and oceanic islands foraminifera genus, with a large test (usually more than 8 X]($$q[]!Z&(&{! mm diameter), with varied shape (globular, hemispheric) ‚GKGKGQG KKQK| Atoll was studied revealing the presence of calcareous !‚GG::G— algae (Porolithon pachydermum, Sporolithon sp., or spiraled, and the last ones are distributed in layers. Lithoporella sp., Lithophyllum sp.), corals (Siderastrea Irregular openings between lateral walls occur when stellata and Favia gravida), vermetids (Dendropoma new chambers are added. These unilocular microfossils irregulare and Petaloconchus varians) and foraminifers have, when alive, a common reddish color, caused by (Homotrema rubrum) in the bioconstruction of this the pigment decomposition of hosted endossimbionts neogene ocean reef. Later taxonomic studies may X`]"`'\[‚Y($q($””{:: identify the calcareous algae species of the genus G)V?GGKGXŠ`Y($%({! proposed here. A paleoecological model should be Remarks: The high wave energy on this reef environment elaborated to explain the predominance of these is responsible for the fragmentation of the reef structure, organisms in the reef construction, along the Holocene, which is composed by corals, coralline algae and different from the coral reefs around the world. several encrusting organisms, e.g. Homotrema and vermetids (Dendropoma irregulare) in the Rocas Atoll X\["]Y'Z&&({"GK Acknowledgments XY]`']($”‰{! ‚GKGGVG'Y>XY KG"{K Dendropoma irregulare (Orbigny, 1842) (Figures granting his Doctorate scholarship. Petrobras and 2F and 3B) ]ŠKGG Description|G?):?G”! G?V!'Š"]KGG }QG:?!Q! GXQG ?G:!'K coordinator of Reserva Biológica Maurizélia Brito). Q[GV other shells. Dendropoma nebulosa (Dillwyn, 1817) is the same. References Remarks|`KK Š` ! Š! ![ "]\‚ Š![ 'Y ! ![ '\`Y‹!VGQKMillepora species occuring the Rocas Atoll (Brazil’s Northeast). Other occurrences in Brazil, including a previously new species. Proceedings of the

Revista Biotemas, 24 (2), junho de 2011 Reef bioconstructors of the Rocas Atoll 45

Biological Society of Washington †G ! ((% ! ~ ! LABOREL, J. Les peuplements de madréporaries des côtes ”()$%Z&&Z! tropicales du Brêsil. Annales University d’ Abidjan, Abidjan, v. Š` ! Š! ![ \]Y Š! Š![ ‚Y ! ˜![ ~!()Z(($$:! Š]'!!'!Y?GQK `I]‹!Š!!Y! GŠ`ŠVXKŠG YG |"G($”!%‰! Brazil). Biota Neotropica !‰!()$Z&&‰! `I]‹!Š!!Y![ARAÚJO‚!Š!![Y]`'], M. C. The Š` ! Š! ![ Š] '! ! '! V :Q K coral reefs off the coast of Eastern Brazil. In: INTERNATIONAL Favia gravidaXS(””{K"!Biota Neotropica ']` ^Š]Š  ($”” ' ! Paulo, v. 7, p. 1-7, 2007. Proceedings... '|'($””!S ! ANDRADE ! ]! ] K X LITTLER Š! Š![ LITTLER ! ! Caribean reef plant. An    >{! Anais da Associação dos Geógrafos Brasileiros!((!Z$) Bahamas, Florida and Gulf of Mexico.†G|]KKG (($%$! Z&&&!%qZ! ARAÚJO, T. M. F. Morfologia, composição, sedimentologia e LOEBLICH ! ![ TAPPAN, H. Treatise on Invertebrate história evolutiva do recife de coral da Ilha de Itaparica, Bahia. Paleontology. Y?^V| Q K ($q! ($”q! $Z K! XŠ { M $&&! "G!($”q! LOEBLICH!![TAPPAN, H. Foraminiferal genera and their CLOUD!!QKŠ!q!: Y?^V| Q K ($””! topography and shoal-water ecology. US Geological Survey $‰&! Professional PapersG!Z”&!~()qq%($%$! `]†Y‚Š\!!YKKG`V!The '\S'!![, D. O. Cnidarians of the Atol das Journal of Geology'G!%”!q~&)q”‰($%&! Rocas. |Y‚Y‚]Y`']`^Š]Š” MACHADO ! ›![ ]‹ F. B. C. Principais espécies de ($$‰Q!ProceedingsGQ|'($$‰!!qq~)qq! foraminíferos e briozoários do Atol das Rocas. Revista Brasileira de Geociências':!Zq!q!Zq‰)Z(($$q! EMILIANI, C. On the species Homotrema rubrum (Lamarck). Cushman Found Foraminifera Research Contribution, v. 2, p. Š'Y‚^ ! ‚: G K (q~)(q‰($%(! phytoplankton within the cuphotic zone of Lake Biwa, Japan. Japanese Journal of Limnology, Tokio, !%‰!~$%)q(&($$‰! \ ! ! Š![ "]Y' ! †! ›! Š K G ecological succession of encrusting organisms in recent coralline- MAIDAŠ![FERREIRA"!!'KK"|?! algal frameworks from Atol das Rocas, Brazil. Palaios, Lawrence, | Y‚Y‚]Y` ']` ^Š]Š ” ($$‰ !(q!Z!(q%)(%”($$$! Q!ProceedingsGQ|'($$‰!!Z~)Z‰q! \ ! ! Š![ "]Y' ! †! ›! ' Y^`], L. A. The Contributions of biogeomorphology to community structure of the coralline algal reefs from Atol das K:Q!Palaeogeography, Palaeoclimatology, G"!Coral Reefs\:!($! PalaeoecologyY?^V!Z($!~%)%(Z&&%! Z&%)($Z&&(! Y^`] `! ![ S` \! ![ CARTER, N. E.A. '\`, E![ \]Y, J. H. Holocene development of the "GQ | V ? G K! Belize Barrier Reef. Sedimentary Geology, Fern Barrow, v. 164, Geomorphology, Charlottesville, v. 47, p. 3-14, 2002. p. 223-236, 2004. YS ! ![ `S, F. L. Release of planula larvae, settlement and development of Siderastrea stellata S(”” \‚‹`"!['‚], C. B. Corais do Sul da Bahia. Rio de XG{!Hydrobiologia"!%&(!(~$) ›|Y($$q!(”$! 147, 2003. HORTA ! ! "  YS!![`S!`!['\]YŠ![›]\Y]Y articuladas do litoral brasileiro – uma síntese do conhecimento. !'Siderastrea siderea (Ellis Biotemas, Florianópolis, v. 15, n. 1, p. 7-44, 2002. (‰”{| \ } G K ‡‡'\!‡!!tol das Rocas, litoral do nordeste do Brasil '::G?!Checklist, M  >! | '\]"Y\ Rio Claro, v. 6, n. 4, p. 505-510, 2010. '![ 'Š] !![ ˜]‹ !‚![†Y Š![ ""‚) Y]`'], M. C. Construções carbonáticas da costa norte do BORN, M. L. (Eds). Sítios geológicos e paleontológicos do Brasil. Estado da Bahia (Salvador a Subaúma). ($”‰!(q~K! "|YŠ—'ŠZ&&Z!!~‰$)~”$! XŠ { M "G ‡‡'\ ! ‡! ![ `I] ‹! Š! ! Y! XG? !($”‰! > "{| : Q :Q OTTMAN ! O`R P R> ! algae. | Y‚Y‚]Y` ']` ^Š]Š ” Revue de Géographie Physique et de Géologie Dinamique, ($$‰Q!ProceedingsGQ|'($$‰!!‰~! !Z!(&()(&‰($~! LABOREL, J. Madréporaires et hydrocoralliaires recifaux des Y]![†]`‡`Y†!›!} :!Q> of Hydrolithon Foslie, Porolithon Foslie and Pseudolithophyllum geographie. Annales Institut Oceanographie, Paris, v. 47, p. 171- Lemoine (Corallinaceae, Rhodophyta) and their relationships to ZZ$($$! ‡!Phycologia!Z‰!(%$)(‰($””!

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Y!![ŠY]S!!S![`S!Š!'!Š Š‚\ ! ! †! Atlantic reef corals. | Q K KK! Š($‰(!(q! Revista da Gestão Costeira Integrada, Itajaí, v. 10, p. 331-345, ] Š! ]![ `Š] S! "![ ‡‡'\ ! ‡! ! 2010. >| : PERRIN '! K do recife biogênico. Revista Brasileira de Geociências, Curitiba, acervulinid foraminifera, their contribution to reef construction and !~$!Z~”)Zq~Z&&$! other biogenic facies. GeobiosY?^V!Z%!!‰Z%)‰‰( ] Š! ]![ Š`` '! ! ]![ `Š] S! "! ($$Z! Distribuição de vermetídeos incrustantes no Atol das Rocas, ! ]! ' ] X'  >! Boletim do Museu Paraense Emílio Anthozoa). ($$%! ZZ‰ K! ‚ X ‹{ M Goeldi. Série Ciências Naturais, Belém, v. 5, p. 225-231, 2010. !($$%! ‚]‚, D. R. Ecology and morphology of recent coral reefs. ! ]! ' K ! Proceedings of the Biological Reviews, Cambridge, v. 44, !q~~)q$”($$! Biological Society of Washington †G ! ((& ! Z ! †]`‡`Y†!›!The coralline red algae|QK (‰)(”%($$‰! G:KK'!]}K| !]!['‚], C. B![Š]‚‚]!![Š˜, ]}K($””!Z”! !'! ' : > †]`‡`Y†!›![CAMPBELL!›!KG Noronha, Brasil. Boletim do Museu Nacional de Zoologia, Rio de Australia species of Lythophyllum (Corallinaceae, Rhodophyta). ›!~%q!()Z(($$Z! Bulletin of the British Museum (Natural History), Botany ], E. de C. Seashells of Brazil. | series`!ZZ!()(&‰($$Z! ($$q!~Z$! †^, J. L. Development in Paleontology and Stratigraphy. ] ! ![ `Y‚‹ ! ![ `Y‚‹, B. Foraminíferos YG|($‰‰!(”%! Holocênicos do Atol das Rocas. Acta Geologica Leopoldensia, `!($!q!‰‰)$(($$! Y‚] Š! ![ Š` ! ![ \YY‹ Š! ! Š![ ‡Y]†`‚]Y Y![ ›¤ ›! S K Favia fravida S($”Siderastrea stellata S(””X' {|> !Boletim do Museu Nacional, , v. 517, !()$Z&&q!

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