I. Introductory Section1. IntroductionThe phylum <strong>Porifera</strong> includes approximately 8350valid species known worldwide (van Soest et al., 2011).Of these, 443 species have been identified in Brazil.<strong>Brazilian</strong> sponges thus comprise approximately 5.3%<strong>of</strong> the diversity <strong>of</strong> <strong>Porifera</strong>, but they still are in a poorstate <strong>of</strong> knowledge. Besides outdated surveys such asthose <strong>of</strong> Mello-Leitão (1961), who listed all spongegroups, and Hechtel (1976), who listed exclusively themarine Demospongiae, there are few recent but partialcompilations, such as those <strong>of</strong> Moraes et al. (2006) forspecies <strong>of</strong> oceanic islands, Azevedo & Klautau (2007)for calcareous sponges, and Hajdu & Lopes (2007)for deep-water species. In the last 30 years many newrecords and new species have been described fromBrazil, and there have been important changes in thecomposition <strong>of</strong> supra-generic groups. Synonyms areabundant and <strong>of</strong>ten hidden within the systematicliterature; their misinterpretation is a major source <strong>of</strong>confusion in the systematics <strong>of</strong> <strong>Brazilian</strong> sponges. Inthis catalogue the existing and proposed synonyms aredetailed and a list <strong>of</strong> names once reported from Brazilbut currently not considered valid is presented. Thisshould help the reader to find out what happened toold names that “disappear” from the recent literatureand to avoid repetition <strong>of</strong> the same species underdifferent names in checklists or other studies.The main goals <strong>of</strong> this catalogue are to tabulateall the literature <strong>of</strong> the Phylum <strong>Porifera</strong> <strong>of</strong> Brazil upto year 2010 and to generate a thorough annotatedclassification <strong>of</strong> <strong>Brazilian</strong> sponges with their currentlyaccepted names and distributions. It provides a guideto the specialized literature and synonyms. Eventualmistakes in the literature (species and localities names,coordinates, depth, systematic misinterpretations,etc.) were corrected. Our results show the specieswith taxonomic problems that should be revised, aswell as the areas underexplored. This catalogue willbe also useful as a basis for diversity estimation andbiogeographic studies.All taxons are arranged in alphabetical order.Taxon diagnoses are not given; the reader can findthem in the Systema <strong>Porifera</strong> (Hooper & van Soest,2002a) or in more recent revisions. Species identifiedonly to supraspecific levels or as “sp.”, “aff.”, or “cf.” weretabulated in a separate list, by alphabetical order andwith less information than the main list. Additionally,we provide a list <strong>of</strong> species by locality and a list <strong>of</strong>nomenclatorial acts here proposed.2. Taxonomical history <strong>of</strong> the<strong>Brazilian</strong> spongesThe date <strong>of</strong> the first study on <strong>Brazilian</strong> sponges isuncertain. In 1766, Pallas described Spongia cavernosa(now Hyattella cavernosa), Spongia fulva (now Aplysinafulva) and Spongia fistularis (now Aplysina fistularis)from “Mare Americanum” (American Sea). This isusually interpreted as corresponding to the TropicalWestern Atlantic, but the country and locality areunknown. Pinheiro et al. (2007) designated a neotypefor Aplysina fulva from SE Brazil and for Aplysinafistularis from Puerto Rico. Here we follow Pinheiroet al. (2007) and assumed that the collection site <strong>of</strong>Spongia fulva could have been in Brazil. Later on,Lamarck (1814) described Spongia carbonaria (nowNeopetrosia carbonaria), also from “Mare Americanum”,maintaining the uncertainty on the beginning <strong>of</strong> thestudy <strong>of</strong> sponges <strong>of</strong> Brazil.Only in 1863, in his monograph <strong>of</strong> freshwatersponges, Bowerbank described seven speciesunequivocally collected in Brazil, all from the Amazonregion. Seven other studies with one or a few speciesfrom Brazil were published from 1860-1880 (e. g.,Schultze, 1865; Müller, 1865; Hyatt, 1877; Selenka,1879). The first major input <strong>of</strong> knowledge on <strong>Brazilian</strong>sponges came in the 1880’s, with the results <strong>of</strong> theH.M.S. Challenger expedition (Poléjaeff, 1883, 1884;Schulze, 1885, 1887a; Ridley & Dendy, 1886, 1887;Sollas, 1886a, 1888). These reports added 40 species tothe list <strong>of</strong> known sponges in Brazil.From 1890 to 1960 relatively few studies werepublished by different authors (average 2.6 articles perdecade), mostly on freshwater sponges (e.g., Cordero,1923, 1924, 1928; Arndt, 1930; Carvalho, 1942;Machado, 1947a,b) but also reporting a few marinesponges (e.g., Carter, 1890; Schulze, 1899; Burton, 1940,1954). In 1956, de Laubenfels reported 13 species <strong>of</strong>marine sponges from Brazil. In 1961, Mello-Leitão andcollaborators made the first compilation <strong>of</strong> <strong>Brazilian</strong>sponges, including 82 species from all three classes andboth marine and freshwater demosponges.The number <strong>of</strong> studies on <strong>Brazilian</strong> spongesbegan to increase in the 1960’s, when Volkmer-Ribeiro11
<strong>Catalogue</strong> <strong>of</strong> <strong>Brazilian</strong> <strong>Porifera</strong>started her studies on <strong>Brazilian</strong> sponges (Fig. 1). Shepublished since then more than 130 studies includingarticles, book chapters, and conference proceedingsfrom 1963 to 2010, representing a major contributionto the knowledge <strong>of</strong> <strong>Brazilian</strong> sponges, especially <strong>of</strong>freshwater. From 1960 to 1969, a total <strong>of</strong> 10 studieswere published on <strong>Brazilian</strong> sponges (e.g., Mello-Leitão et al., 1961; Volkmer, 1963a, b; Volkmer-Ribeiro,1969; Fig. 1).In the 1970’s there was an increase in the number<strong>of</strong> articles (34) and <strong>of</strong> authors publishing on <strong>Brazilian</strong>sponges. The studies <strong>of</strong> Borojevic (1971) and Borojevic& Peixinho (1976) on calcareous sponges, Volkmer-Ribeiro & de Rosa-Barbosa (1972, 1974, 1978), deRosa-Barbosa (1979) and Bonetto & Ezcurra de Drago(1973) on freshwater sponges, and those <strong>of</strong> Johnson(1971), Collette & Rützler (1977), Mello-Leitão &Pacheco-Coelho (1978), Maggs et al. (1979) andespecially <strong>of</strong> Mothes de Moraes (e.g., Volkmer-Ribeiro& Mothes de Moraes, 1975; Mothes de Moraes, 1977,1978) on marine Demospongiae greatly increased thetaxonomic knowledge <strong>of</strong> <strong>Brazilian</strong> sponges. However,these studies were limited to relatively few taxonomicgroups or geographic areas. The main contribution tothe taxonomy <strong>of</strong> <strong>Brazilian</strong> <strong>Porifera</strong> in this decade wasthe description <strong>of</strong> 60 species <strong>of</strong> marine demospongescollected by the Calypso expedition all along theEastern <strong>Brazilian</strong> coast, from Fernando de NoronhaArchipelago to Rio Grande do Sul State (Boury-Esnault, 1973).The 1970’s also witnessed an increase in thenumber <strong>of</strong> research approaches, which were untilthen limited to taxonomy and faunistics. The firststudies on the ecology <strong>of</strong> sponges in Brazil were those<strong>of</strong> B<strong>of</strong>fi (1972) reporting ophiuroids associated tomarine sponges and <strong>of</strong> Volkmer-Ribeiro & de Rosa-Barbosa (1972, 1974) on sponge-sponge and molluskfreshwatersponge associations. The first studies onsponge chemistry and evolution were published by theend <strong>of</strong> the decade (Kelecom & Kannengiesser, 1979;Volkmer-Ribeiro, 1979). In the first zoogeographicalanalysis <strong>of</strong> <strong>Brazilian</strong> sponges, Hechtel (1976) listed 156marine demosponges occurring in Brazil.The trend <strong>of</strong> increase in authors and areas <strong>of</strong>knowledge <strong>of</strong> studies on <strong>Brazilian</strong> sponges persistedand gained impulse in the 1980’s (30 articles). Themain contributions to the taxonomy were those <strong>of</strong> Solé-Cava et al. (1981), Hechtel (1983), Volkmer-Ribeiro(1984, 1986), Volkmer-Ribeiro et al. (1981, 1983, 1984,1988) and Mothes de Moraes (1980, 1981, 1983, 1985a,b, 1987). More ecological studies were published thanin the previous decades, focusing on fish predation(Volkmer-Ribeiro & Grosser, 1981), associated fauna(Duarte & Morgado, 1983), vertical distribution (Estonet al., 1986) and pollution-biomonitoring (Muricy,1989). The first pharmacological study <strong>of</strong> a <strong>Brazilian</strong>sponge was published by the end <strong>of</strong> the decade (Attaet al., 1989; Fig. 2).In the 1990’s, there was a two-fold increase in thenumber <strong>of</strong> studies published (from 30 to 72 articles; Fig.1). The number <strong>of</strong> authors and <strong>of</strong> areas <strong>of</strong> knowledgealso increased. In this decade were published the firststudies on genetics using both allozyme electrophoresis(Solé-Cava et al., 1991; Klautau et al., 1994a, 1999;Lazoski et al., 1999) and DNA analysis (Lôbo-Hajduet al., 1999), cytology (Boury-Esnault et al., 1994a),biochemistry (Zierer et al., 1995), paleontology(Sifeddine et al., 1994; Ferreira & Fernandes, 1997),phylogeny (Hajdu, 1995, 1999), biomass (Melão &Rocha, 1996b), growth rates (Melão & Rocha, 1998),chemical ecology (Epifanio et al., 1999), and larvalbiology (Vacelet, 1999) <strong>of</strong> sponges from Brazil.The exponential growth <strong>of</strong> the literature on<strong>Brazilian</strong> <strong>Porifera</strong> continued from 2000 to 2009,with 250 articles published. New areas <strong>of</strong> knowledgestudied included spiculogenesis (Custódio et al.,2002), bioerosion (Reis & Leão, 2002), bioinvasion(Ferreira et al., 2006), microbiology (Turque et al.,2008; Hardoim et al., 2009; Marinho et al., 2009), DNAsequencing (Cárdenas et al., 2009), and reproduction(Lanna et al., 2007b). All areas previously studied hada great increase in number <strong>of</strong> publications, especiallytaxonomy, chemistry, pharmacology, and ecology. Thestudy <strong>of</strong> <strong>Porifera</strong> in Brazil during the first decade <strong>of</strong> theXXI century was marked by the realization <strong>of</strong> the 7 thInternational Sponge Symposium (7 th ISS) in Armaçãodos Búzios, Rio de Janeiro State, in May 2006. Thisconference had 260 participants from 35 countriesand attracted a great interest from the <strong>Brazilian</strong>scientific community, with 91 <strong>Brazilian</strong> participantsfrom 18 institutions presenting a total <strong>of</strong> 71 abstracts.It is therefore not surprinsing that the year with thehighest number <strong>of</strong> publications in the history <strong>of</strong>spongiology in Brazil so far was 2007 (59 articles),when the proceedings <strong>of</strong> the 7 th ISS were published(Fig. 2; Custódio et al., 2007).Another major accomplishment in the decade from2000 to 2009 was the Project REVIZEE, a governmentalprogram aimed at listing and mapping the biologicalresources <strong>of</strong> the <strong>Brazilian</strong> Exclusive Economic Zone(EEZ; e.g., Amaral et al., 2003; Amaral & Rossi-Wongtschowski, 2004; Lavrado & Ignacio, 2006). The12
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- Page 6 and 7: SUMMARYAbstract ...................
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- Page 28 and 29: Taxonomic ListTYPE LOCALITY: Brazil
- Page 30 and 31: Taxonomic ListREMARKS: The specimen
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Taxonomic List(REVIZEE sta. C5-2R,
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Taxonomic ListH.M.S. Challenger cre
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Taxonomic ListCliona carteri, Hecht
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Taxonomic Liststate: Guarapari (20
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Taxonomic ListFinali, 1993: 274; R
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Taxonomic ListChondrilla phyllodes
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Plates
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Taxonomic ListFig. 6. Type specimen
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Taxonomic ListFig. 8. Type specimen
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Taxonomic ListFig. 10. Type and rep
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Taxonomic ListFig. 12. Type specime
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Taxonomic ListFig. 14. Type specime
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Taxonomic List1973 [description]; H
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Taxonomic ListSão Paulo State: Sã
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Taxonomic ListE Axinella Schmidt, 1
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Taxonomic ListTYPE LOCALITY: Brazil
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Taxonomic ListTYPE SPECIMENS: Holot
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Taxonomic Listthe Lithistida is wid
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Taxonomic Listet al., 2004b [descri
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Taxonomic ListCladochalina armigera
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Taxonomic ListK Chalinula zeae de W
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Taxonomic List& van Soest, 1998: 91
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Taxonomic ListSantos et al., 2002b
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Taxonomic List1988; Lehnert & van S
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Taxonomic ListK Oceanapia fistulosa
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Taxonomic List[description]). ARAGU
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Taxonomic ListState: Caseara, Aragu
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Taxonomic ListRibeiro & Batista, 20
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Taxonomic ListK Metania spinata (Ca
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Taxonomic ListOncosclera navicela,
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Taxonomic Listdescription]). Manaus
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Taxonomic List& Pronzato, 2002), Ve
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Taxonomic ListParaná River (near o
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Taxonomic List2007b [faunistics]).
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Taxonomic ListK Heteromeyenia insig
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Taxonomic ListType species: Heterom
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Taxonomic Listal., 2003 [faunistics
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Taxonomic ListWORLD DISTRIBUTION: B
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Taxonomic ListPará State: Santaré
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Taxonomic ListRiver (Itaipu H.P.S.;
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Taxonomic ListMoraes et al., 2006:
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Taxonomic ListBRAZILIAN RECORDS: So
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Taxonomic Listfrom those presented
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Taxonomic ListFortaleza (Johnson, 1
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Taxonomic ListTYPE SPECIMENS: Holot
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Taxonomic ListState: Off Guarujá (
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Taxonomic ListFamily Guitarridae De
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Taxonomic ListSchizolectotypes USNM
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Taxonomic List1994: 569; Hajdu, 199
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Taxonomic ListSuborder Myxillina Ha
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Taxonomic ListTYPE SPECIMENS: Synty
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Taxonomic ListWORLD DISTRIBUTION: C
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Taxonomic List1998: 88; Muricy & Si
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Taxonomic ListHajdu & Lopes, 2007:
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Taxonomic Listal., 2006: 166; Cedro
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Taxonomic ListWORLD DISTRIBUTION: E
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Taxonomic Listal., 2008 [pharmacolo
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Taxonomic List(Amapá or Pará (Col
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Taxonomic ListRegion: Espírito San
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Taxonomic ListP Hyalonema (Cylicone
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Taxonomic ListC2-20F (off Linhares,
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Taxonomic ListFor synonyms see Taba
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Records Identified Only to Genus Le
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Records Identified Only to Genus Le
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Records Identified Only to Genus Le
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Records Identified Only to Genus Le
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Records Identified Only to Genus Le
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Records Identified Only to Genus Le
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Records Identified Only to Genus Le
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Records Identified Only to Genus Le
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IV. LIST OF NOMENCLATORIAL ACTSA sy
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List of Nomenclatorial ActsHaliclon
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V. SPECIES NAMES PREVIOUSLY REPORTE
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Species Names Previously Reported F
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Species Names Previously Reported F
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Species Names Previously Reported F
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Species Names Previously Reported F
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Species Names Previously Reported F
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VI. LIST OF SPECIES BY LOCALITYThe
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List of Species by LocalityPottsiel
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List of Species by Localityfloridan
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List of Species by LocalityOff São
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List of Species by LocalityAcarnus
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List of Species by LocalityJoão Pi
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List of Species by LocalityHymeniac
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List of Species by LocalityEunapius
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VII. Taxonomic IndexThis index incl
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Taxonomic Indexarmigera, Callyspong
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Taxonomic Indexcarteri, Craniella .
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Taxonomic Indexcunctatrix, Spirastr
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Taxonomic IndexForcepia (Ectoforcep
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Taxonomic Indexjanusi, Verongia ...
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Taxonomic Indexmucronata, Myxilla .
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Taxonomic Indexplumosa var. palmeri
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Taxonomic Indexschmidti, Cliona ...
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Taxonomic Indextoxiformis, Erylus .
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VIII. REFERENCESRemarks: The articl
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Referencesdo Cabo - RJ. Revista Pes
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ReferencesDémosponges du plateau c
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ReferencesCarter, H.J. (1869) On Gr
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ReferencesSudamericanas. Comunicaci
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ReferencesDickinson, M.G. (1945) Sp
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ReferencesPark, Ceará State, Brazi
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Referencesinvertebrados marinhos da
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ReferencesHooper, J.N.A. & van Soes
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ReferencesSouza, A.O., Galetti, F.C
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ReferencesRaspailia (Raspaxilla) bo
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Referencesof the West Indies: Disco
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Referencese futuro. In: Viana, D.L.
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References*Muricy, G. & Minervino,
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References(Brazil) (Tetillidae, Spi
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Referencesfive new species. Zoologi
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ReferencesRose, C.S. & Risk, M.J. (
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ReferencesMit Nachträgen zu den Sp
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ReferencesSystema Porifera: A guide
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ReferencesSollas, W.J. (1889) On th
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ReferencesSociété Scientifique et
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ReferencesBiodiversity, Innovation
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References*Volkmer-Ribeiro, C. & Ta
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ReferencesWiedenmayer, F. (1977) Sh