547
P r : B , i s - 2007
Introduction
The genus Cinachyra Sollas, 1886, of the demosponge family Tetillidae Sollas, 1886, comprises globular sponges with a radial skeleton of oxeas and triaenes, sigmaspires, a cortex reinforced by auxiliary oxeas, and flask-shaped porocalyces (van Soest and Rützler 2002). There is considerable confusion in the literature distinguishing between the closely related genera Cinachyra and Cinachyrella Wilson, 1925, because many authors disregarded the presence or absence of a distinctive cortex, which is now considered the most important character to distinguish between the two genera (Rützler 1987, van Soest and Rützler 2002). Rützler (1987) and Rützler and Smith (1992) revised the Western Atlantic species of Cinachyra, and concluded that they all belong instead to Cinachyrella, in which the cortex is absent. In the most recent revision of the family Tetillidae, van Soest and Rützler (2002) recognized only a single valid species as belonging with certainty to the genus Cinachyra, viz., C. barbata Sollas, 1886.
So far, twelve valid species of tetillid sponges have been described from Brazil: Acanthotetilla rocasensis Peixinho et al., 2007; Acanthotetilla walteri Peixinho et al., 2007; Cinachyrella alloclada (Uliczka, 1929), Cinachyrella apion (Uliczka, 1929), Cinachyrella kuekenthali (Uliczka, 1929), Craniella carteri Sollas, 1886, Craniella corticata (Boury-Esnault, 1973), Craniella quirimure Peixinho, Cosme and Hajdu, 2005, Tetilla euplocamus Schmidt, 1868, and Tetilla radiata Selenka, 1879 (Schmidt 1868, Selenka 1879, Sollas 1886, 1888, Fischel-Johnson 1971, Boury-Esnault 1973, Rützler and Smith 1992, Lazoski et al. 1999, Santos and Hajdu 2003; Peixinho et al. 2005; Muricy and Hajdu 2006).
Craniella cranium Müller, 1776 was quoted from Fernando de Noronha Archipelago (Carter 1890); however, Carter’s description was insufficient, and this record was considered questionable and unrecognisable (Hechtel 1976, Moraes et al. 2006). Cinachyra rhizophyta Uliczka 1929, recorded from Ceará State (Fischel-Johnson 1971) was put in synonym with Cinacyhrella apion (cf. Rützler and Smith 1992). Unfortunately, many Brazilian records of these species, particularly of Cinachyrella spp., did not include descriptions (Mello Leitão et al. 1961, Hechtel 1976, Collette and Rützler 1977, Atta et al. 1989, Muricy et al. 1991, 1993, 2006, Muricy and Moraes 1998, Lobo-Hajdu et al. 1999, Santos et al. 1999, 2002, Muricy and Silva 1999, Moraes et al. 2003, 2006, Santos et al. 2004, Hajdu et al. 2004). Two of such more recent records are of undescribed species of Cinachyra (Cinachyra sp.; Hajdu et al. 2004, Muricy et al. 2006). Since the genus Cinachyra was until recently considered to have a cold-temperate to polar distribution (Kerguelen Islands, Patagonia and Antarctica; van Soest and Rützler 2002), we decided to investigate in more detail the two records of Cinachyra from the subtropical SW Atlantic. Both species turned out to be new, but only one of them really belongs to Cinachyra. In this study, we describe the new species of Cinachyra collected off Espirito Santo State, SE Brazil, by Project REVIZEE Central SCORE (Muricy et al. 2006), and discuss the identity of the record of Cinachyra sp. from São Paulo State, Southern Brazil (24°20’527’’S – 43°46’759’’W), collected by Project REVIZEE South SCORE (Hajdu et al. 2004).
The Project REVIZEE (in portuguese, “Programa de Avaliação do Potencial Sustentável dos REcursos VIvos da Zona Econômica Exclusiva”, or Program of Evaluation of the
A new species of Cinachyra (Demospongiae:
Tetillidae) collected by Project REVIZEE off Espírito
Santo State, SE Brazil
Pablo R.D. Rodriguez, Guilherme Muricy(*)
Laboratório de Porifera, Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro. Quinta da Boa Vista s/n, 20940-040, Rio de Janeiro, RJ, Brasil. muricy@acd.ufrj.br
Abstract: A new species of Cinachyra Sollas (Porifera, Spirophorida) was dredged from 500 m depth off Espirito Santo
State, SE Brazil, by Project REVIZEE Central SCORE. Cinachyra helena sp. nov. is characterized by the presence of a single category each, of protriaenes, anatriaenes, anisoactinal choanosomal oxeas, isoactinal cortical oxeas, and sigmaspires. The cortex is formed by small isoactinal oxeas arranged obliquely to the surface, and an anchoring basal spicule mass is absent. This is the first valid record of Cinachyra from the Atlantic; all similar species lacking a cortex should be transferred to Cinachyrella or other genera of Tetillidae. At the present state of knowledge, there are four valid species in the genus Cinachyra: C. barbata Sollas, C. crustata (Wilson), C. uteoides Dendy, and C. helena sp. nov.
Sustainable Resources of the Economic Exclusive Zone) is an initiative of the Brazilian Government to comply with the United Nations Convention on the Law of the Sea (UNCLS). The Brazilian continental shelf and slope (down to 2,076 m depth) were divided in four sectors (called “SCORES”: North, Northeast, Central, and South), in which extensive surveys were done to estimate the diversity and abundance of planktonic, nectonic and benthic organisms and their sustainable exploitation potential (eg, Amaral and Rossi-Wongtschowski 2004, Costa et al. 2005, Lavrado and Ignacio 2006).
Material and methods
The area sampled by Project REVIZEE Central SCORE ranges from Salvador (11°S), Bahia state, to Cabo de São Tomé (22°S), in Northern Rio de Janeiro state, including the islands and seamounts of the Vitória-Trindade chain (Fig. 1). The continental shelf breaks at approximately 70 m depth, and the seafloor is dominated by carbonatic sediments, corals and calcareous algae. The slope is mostly covered by foraminiferan deposits. Sponges have the greatest biomass among benthic organims in this area (Lavrado 2006). A complete description of the environment and benthic communities of the Central SCORE region is given by Lavrado and Ignacio (2006).
Collections were done by dredging and bottom trawling on board of the RV Astro Garoupa, between 20 and 2,076 m depth, from 19/X/1997 a 24/XI/2003. Sponges were fixed in ethanol 70% or formalin 4%, and deposited in the Porifera collection of Museu Nacional (Rio de Janeiro, Brazil). Photographs of both preserved specimens and of the skeleton by light microscopy were taken with a Nikon Coolpix digital camera. Spicule slides were prepared by dissociation of a small fragment of sponge in boiling nitric acid. Sponge fragments were dehydrated in an alcohol series (50-100%) with a final xylene step and included in paraffin. Transverse sections of the skeleton were mounted on microscope slides for identification. Twenty spicules of each kind were measured per specimen. Size ranges and means (underlined) are given in the text.
Systematics
Phylum Porifera Grant, 1836 Class Demospongiae Sollas, 1885 Subclass Tetractinomorpha Lévi, 1953 Order Spirophorida Bergquist and Hogg, 1969
Family Tetillidae Sollas, 1886 Genus Cinachyra Sollas, 1886
Definition: Tetillidae with cortex reinforced by auxiliary
oxeas, with flask-shaped porocalyces (van Soest and Rützler 2002).
Cinachyra helena sp. nov.
(Figs. 2–3, Tab. 1)
Synonyms: Cinachyra sp., Muricy et al. 2006: 115
Diagnosis: Cinachyra with a single category each of
protri-aenes, anatriprotri-aenes, anisoactinal choanosomal oxeas, isoacti-nal cortical oxeas, and sigmaspires. Cortex formed by small isoactinal oxeas arranged obliquely to the surface. Anchoring spicule tufts absent.
Material examined: Holotype: MNRJ 3635C, Espírito Santo
State, Brazil; dredging, project REVIZEE Central SCORE II, station 20C-deep, RV Astro Garoupa, 22/XI/1997, 19°17`S – 37°57`W, 500 m depth. Paratype: MNRJ 3658B, same lo-cality, date and collector.
Material examined for comparison: Cinachyra sp. sensu
Hajdu et al. (2004): MNRJ 2814 A-H, São Paulo state, Bra-zil; dredging, project REVIZEE South SCORE, station 6659, RV Prof. W. Besnard, 09/I/1998, 24°20’S – 43°46’W, 505 m depth.
Description: External form hemispherical to subspherical
(Fig. 2). Size up to 2.5 cm in diameter by 1.5 cm high. Color in vivo unknown; beige after fixation in alcohol 70%. The ho-lotype has irregular porocalyces, up to 3 x 1 mm wide by 2.5 mm deep (Fig. 2A). A thin, whitish cortex is clearly visible in sectioned specimens (Fig. 2B). The paratype was fragmented, and the part of its surface that is left has no porocalyces (Fig. 2C). Oscules are not visibe in preserved specimens, probably contracted. Surface even, strongly hispid, with adhering sand grains. The sponge is fixed by a broad attachment base, with-out anchoring basal spicule mass (Fig. 2D). Consistency firm, slightly compressible, inelastic.
Skeleton: Choanosomal skeleton with radial tracts of
anisoac-tinal coanossomal oxeas, anatriaenes and protriaenes, which project through the cortex and the surface of the sponge (Fig. 3A). The cladomes of both pro- and anatriaenes are directed outwards. Tracts are wider (580-733 µm thick) in the ecto-some than in the choanoecto-some (76-202 µm thick), and are 50-530 µm apart. The cortex is composed of two layers (Fig. 3A): the internal layer (505-884 µm thick) is formed by small isoactinal oxeas arranged obliquely to the surface; the outer layer (253-1137 µm thick) is formed only by sigmaspires and the extremities of the radial tracts of megascleres. Sig-maspires are dispersed randomly both in the cortex and in the choanosome.
Spicules: Choanosomal anisoactinal oxeas, slightly curved at
the thinner extremity: 2075-3305-4300 x 40-43-50 µm (Fig. 3B). Cortical isoactinal oxeas short, robust, fusiform: 425-650-850 x 26-34-43 µm (Fig. 3C). Anatriaenes rare, vary-ing from very thin, reduced forms to long and robust spic-ules, with clads short and slightly curved, forming an almost straight angle with the rhabdome: clads 19-72-116 x 2-18-31 µm; rhabdome 1310-3421-8381 x 5-24-34 µm (Fig. 3D, E). Protriaenes abundant, occasionally irregular, with clads long and straight and rhabdome long, ending abruptly: clads 130-214-275 x 12-16-21 µm, rhabdome 1500-3258-6250 x 17-23-34 µm (Figs. 3F, G). Sigmaspires C- or S-shaped, with spines
large, recurved, and more abundant close to the tips, which are enlarged, irregular: 10-14-21 µm (Fig. 3H).
Ecology: The paratype (MNRJ 3658B) was growing over
a foliaceous sponge, Phakellia sp. The two specimens were partially covered by sand grains.
Geographical and bathymetric distribution: Off Espirito
Santo State, Brazil (19°17`14``S – 37°57`13``W), 500 m depth (Fig. 1).
Etymology: This species was named in honour of Dr. Helena
Passeri Lavrado, in recognition of her successful efforts to coordinate the Benthic Ecology Group of Project REVIZEE Central SCORE.
Discussion
Only two specimens of the new species were found; of these, only the holotype had porocalyces (Fig. 2A). This is an important issue, since the presence of porocalyces is the main character that separates Cinachyra from Craniella Schmidt, 1870 (van Soest and Rützler 2002). The paratype was fragmented, and most of its surface was lost (Fig. 2C). Since the spicules and all other characters of the two
specimens were nearly identical in shape and size (Table 1), we assumed that the porocalyces were present in the lost part of the paratype’s surface.
The genus Cinachyra is currently considered to contain a single species, Cinachyra barbata Sollas, 1886, and most other species previously described as Cinachyra were transferred to Cinachyrella (van Soest and Rützler 2002). However, a literature survey indicates that at least one other species described as Cinachyra (C. uteoides Dendy, 1924) and another described as Tetilla (Tetilla (Cinachyrella) crustata Wilson, 1925) do have both porocalyces and a cortex made up of small oxeas, and should therefore be considered as valid species of Cinachyra. Bergquist (1968) expressed doubts whether C. novae-zealandiae Brondsted, 1924 has true porocalyces, since Brondsted’s (1924) description is not clear about the nature of its aquiferous openings. It is clear however from the original description that C. novae-zealandiae has a cortex of small curved oxeas forming a palisade. We agree with Bergquist (1968) that a revision of the holotype is needed to clarify the best taxonomic placement of C. novae-zealandiae, if in Cinachyra or in Craniella (but not in Tetilla, as suggested by Bergquist, loc. cit.). For the time being, our interpretation is that the aquiferous openings
Fig. 1: Area studied by the Project
REVIZEE, Central SCORE, between Salvador and Cabo de São Tomé, Brazil, with the location of the collecting site of Cinachyra
helena sp n. (BA, Bahia State; ES,
Espírito Santo State; RJ, Rio de Janeiro State).
of C. novae-zealandiae are not porocalyces, and therefore we suggest to classify it as Craniella novae-zealandiae. A detailed taxonomic revision of the family Tetillidae, including observation of type specimens of as many species as possible, is clearly needed for a better understanding of the scope of the genus Cinachyra.
Cinachyra barbata presents two categories of protriaenes, one of anatriaenes, two of isoactinal oxeas, and one of sigmaspires. The anatriaenes only occur in the basal spicule mass. Cinachyra helena sp. nov. differs from Cinachyra barbata by the possession of only one category of protriaenes, choanosomal anatriaenes, and the absence of a basal mass. Furthermore, its choanosomal anisoactinal oxeas and cortical isoactinal oxeas are smaller than those of Cinachyra barbata, which has choanosomal isoactinal oxeas of approximately 8000 x 70 µm and cortical isoactinal oxeas around 900 x 36 µm (Sollas 1888).
Cinachyra helena sp. nov. is similar to C. uteoides in the absence of a basal (anchoring) spicule mass: both species are more or less flattened basally, with a broad attachment base. Cinachyra uteoides differs from the new species however by the presence of protriaenes in two size classes, the isoactinal choanosomal oxeas, the larger cortical oxeas (2600 x 80 µm), and the smaller rhabdome of the anatriaenes (2400-2900 x 12-16 µm) (Dendy 1924, Bergquist 1968).
Cinachyra crustata (Wilson 1925) differs from C. helena sp. nov. in many characters, such as the flattened body shape, presence of root-like spicule tracts (= anchoring basal spicule mass), larger porocalyces, thinner cortex with tangentially-disposed rather than obliquely-tangentially-disposed oxeas, and larger, isoactinal choanosomal oxeas (7000-9000 µm).
Although it is uncertain whether it really has porocalyces (cf. Bergquist 1968), Craniella novae-zealandiae Brondsted 1924 shares with the new species the globular shape without spicule basal mass and a strongly hispid surface. They differ,
Spicules Holotype MNRJ 3635C Paratype MNRJ 3658B
Choanosomal oxeas 3050-3635-4300 / 34-41-50 2075-2976-4050 / 40-44-50 Cortical oxeas 425-734-850 / 26-34-43 450-566-775 / 26-34-41 Protriaene clads 130-198-240 / 15-17-20 175-231-275 / 14-17-22 Protriaene rhabdome 2150-3615-6250 / 19-23-26 1500-2901-5000 / 17-22-34 Anatriene clads 38-75-97 / 10-20-29 19-70-117 / 2-17-31 Anatriaene rhabdome 2425-4315-8381 / 14-26-34 1310-2527-6383 / 5-21-44 Sigmaspires 12-14-22 / 0.5-2 10-14-18 / 0.5-2
Table 1: Measurements (min-med-max length / min-med-max width in µm) of the spicules of Cinachyra helena sp. nov. (n=20). Fig. 2: Cinachyra helena sp. nov.
A. upper view of holotype (MNRJ
3635C) showing the irregular porocalyces in the surface (arrows). B. side view of holotype, showing the radial arrangement of the skeleton and the lighter cortex (arrow). C. upper view of paratype (MNRJ 3658B), in which most of the surface was lost. D. basal view of paratype, showing the broad, flattened attachement surface (scale bars = 1 cm).
however, in that the cortical oxeas of C. novae-zealandiae are curved, its choanosomal oxeas are isoactinal, all megascleres are smaller, and it apparently has no sigmaspires (Brondsted 1924).
We reexamined the specimens of Cinachyra sp. recorded from Southern Brazil (Hajdu et al. 2004). They are flattened, discoidal, with a fringe of projecting spicules.
Both porocalyces and a spiculated cortex are absent, and therefore they would be better classified in Tetilla Schulze 1868, or maybe in a new genus defined by a discoidal shape, rather than in Cinachyra. Cinachyra rhizophyta, recorded from Ceará state, NE Brazil (Fischel-Johnson 1971), was synonymized with Cinachyrella apion by Rützler and Smith (1992). However, the description of C. rhizophyta from
Fig. 3: Skeletal characters of
Cinachyra helena sp. nov. A.
transverse section of the skeleton showing the cortex of smaller oxeas and the radial megasclere tracts.
B. choanosomal, anisoactinal
oxea. C. cortical, isoactinal oxea. D. anatriaene. E. cladome of anatriaene. F. protriaene. G. cladome of protriaene (SEM) H. sigmaspire (SEM).
Ceará does not mention the two categories of protriaenes or the presence of raphides, characters typical of C. apion. We consider the record of Cinachyra rhizophyta sensu Fischel-Johnson (1971) as a synonym of Cinachyrella alloclada, rather than of C. apion. This, however, does not change the list of tetillid species from Brazil, which now includes eleven valid species: Acanthotetilla rocasensis, Acanthotetilla walteri, Cinachyra helena sp. nov., Cinachyrella alloclada, Cinachyrella apion, Cinachyrella kuekenthali, Craniella carteri, Craniella corticata, Craniella quirimure, Tetilla euplocamus, and Tetilla radiata.
The genus Cinachyra appears to be more frequent in deep than in shallow water, with records of C. barbata from 45-549 m depth (Sollas 1886, 1888), of C. uteoides from 55-182 m (Dendy 1924, Bergquist 1968), and of the new species from 500 m depth. Cinachyra helena sp. nov. is the first valid record of the genus from the Brazilian coast and the Atlantic Ocean. The distribution of the genus Cinachyra now includes Kerguelen Island, Patagonia and Antarctica (C. barbata), New Zealand (C. uteoides), Philippines (C. crustata) and SE Brazil (C. helena sp. nov.).
Acknowledgements
We are grateful to Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), Fundação Carlos Chagas Filho de Apoio à Pesquisa do Estado do Rio de Janeiro (FAPERJ), Secretaria da Comissão Interministerial para os Recursos do Mar (SECIRM), and CENPES-PETROBRAS for financial and logistic support. We also thank Dr. Marcia Attias and Noêmia Rodrigues (Laboratório de Ultraestrutura Celular Herta Meyer, Instituto de Biofísica Carlos Chagas Filho, UFRJ) for their help in the use of SEM. We are particularly grateful to Dr. Helena P. Lavrado for inviting us to identify the sponges of Project REVIZEE Central SCORE.
References
Amaral ACZ, Rossi-Wongtschowski C (eds). (2004) Biodiversidade
bentônica da região sudeste-sul do Brasil – plataforma externa e talude superior. Série Documentos REVIZEE – Score Sul.
Instituto Oceanográfico USP, São Paulo
Atta AM, Barral M, Peixinho S, Souza-Atta MLB (1989) Isolation and functional characterization of a mitogenic lectin from the marine sponge Cinachyrella alloclada. Braz J Med Biol Res 22: 379-385
Bergquist PR (1968) The marine fauna of New Zealand: Porifera, Demospongiae, Part I. (Tetractinomorpha and Lithistida). New
Zeal Oceanogr Inst Mem 37(188): 1-105
Bergquist PR, Hogg JJ (1969) Free aminoacid patterns in Demospongiae: a biochemical approach to sponge classification.
Cah Biol Mar 10: 205-220
Boury-Esnault N (1973) Campagne de la Calypso au large des côtes Atlantiques de l´Amérique du Sud (1961-1962) 29. Spongiaires.
Rés Sci Camp Calypso 10: 263-295
Brondsted HV (1924) Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. XXIII. Sponges from New Zealand. Part I.
Vidensk Meddel Dansk Naturh Foren Kjoben 77: 435-483
Carter HJ (1890) Porifera. Notes on the zoology of Fernando de Noronha. J Linn Soc 20: 564-569
Collette B, Rützler K (1977) Reef fishes over sponge bottoms off the mouth of the Amazon River. Proc 3rd Int Coral Reef Symp, Miami 1: 305-310
Costa PAS, Martins AS, Olavo G (eds). Pesca e potenciais de
exploração de recursos vivos na região central da Zona Econômica Exclusiva brasileira. Museu Nacional, série Livros 13, Rio de
Janeiro
Dendy A (1924) Porifera. Part I. Non-Antarctic sponges. British Antarctic (‘Terra Nova’) Expedition, 1910 (Zoology). Nat Hist
Rep 6(3): 269-392
Fischel-Johnson M (1971) Some marine sponges of northeast Brazil.
Arq Ciênc Mar 11(2): 103-116
Grant RE (1836) Animal Kingdom. In: Todd RB (ed). The
cyclopaedia of anatomy and physiology. Vol. 1. Sherwood, Gilbert,
and Piper, London. pp. 107-118
Hajdu E, Santos CP, Lopes DA, Oliveira MV, Moreira MCF, Carvalho MS, Klautau M (2004) Filo Porifera. In: Amaral CZA, Rossi-Wongtschowski CLB (Eds) Biodiversidade bentônica da
região sudeste-sul do Brasil – plataforma externa e talude superior.
Série Documentos REVIZEE Score Sul. Instituto Oceanográfico USP, São Paulo. pp. 49-56
Hechtel GJ (1976) Zoogeography of Brazilian marine Demospongiae.
In: Harrison FW, Cowden RR (eds). Aspects of sponge biology.
Academic Press, New York & London. pp. 237-260
Lavrado HP (2006) Capítulo 1 – Caracterização do ambiente e da comunidade bentônica. In: Lavrado HP, Ignácio BL (eds).
Biodiversidade bentônica da região central da Zona Econômica Exclusiva brasileira. Museu Nacional, Série Livros 18, Rio de
Janeiro. pp. 19-64
Lavrado HP, Ignácio BL (2006) Biodiversidade bentônica da região
central da Zona Econômica Exclusiva brasileira. Museu Nacional,
Série Livros 18, Rio de Janeiro
Lazoski C, Peixinho S, Russo CAM, Solé-Cava AM (1999) Genetic confirmation of the specific status of two sponges of the genus
Cinachyrella (Porífera: Demospongiae: Spirophorida) in the
southwestern Atlantic. Mem Qld Mus 44: 299-305
Lévi C (1953) Sur une nouvelle classification des Démosponges.
Compt Rend Hebdom Séan Acad Sci Paris 236(8): 853-855
Lôbo-Hajdu G, Mansure JJ, Salgado A, Hajdu E, Muricy G, Albano RM (1999) Random amplified polymorphic DNA (RAPD) analysis can reveal intraspecific evolutionary patterns in Porifera. Mem Qld
Mus: 317-328.
Mello-Leitão A, Pêgo AF, Lopes WM (1961) Poríferos assinalados no Brasil. Avul Centr Est Zool Univ Brasil 10: 1-29
Moraes FC, Ventura M, Klautau M, Hajdu E, Muricy G (2006) Biodiversidade de esponjas das ilhas oceânicas brasileiras. In: Alves RJV, Castro, JWA (eds). Ilhas Oceânicas Brasileiras – da
pesquisa ao manejo. Ministério do Meio Ambiente, Brasilia. pp.
147-178
Moraes FC, Vilanova EP, Muricy G (2003) Distribuição das esponjas (Porifera) na Reserva Biológica do Atol das Rocas, Nordeste do Brasil. Arq Mus Nac 61(1): 13-22
Muricy G, Moraes FC (1998) Marine sponges of Pernambuco state, NE Brazil. Rev Bras Oceanogr 46(2): 213-217
Muricy G, Silva OC (1999) Esponjas marinhas do Estado do Rio de Janeiro: um recurso renovável inexplorado. In: Silva SHG, Lavrado HP (eds). Ecologia dos Ambientes Costeiros do Estado
Muricy G, Hajdu E (2006) Porifera Brasilis. Guia de identificação
das esponjas marinhas mais comuns do sudeste do Brasil. Museu
Nacional, Série Livros 17, Rio de Janeiro
Muricy G, Hajdu E, Custódio M, Klautau M, Russo C, Peixinho S (1991) Sponge distribution at Arraial do Cabo, SE Brazil. In: Magoon OT, Converse H, Tippie V, Tobin LT, Clark D (eds).
Coastal Zone ’91. Proc 7th Symp Coast Ocean Manag. ASCE Publications, Long Beach. pp. 1183-1196
Muricy G, Hajdu E, Araujo FV, Hagler AN (1993) Antimicrobial activity of Southwestern Atlantic shallow-water marine sponges (Porifera). Sci Mar 57(4): 427-432
Muricy G, Santos CP, Batista D, Lopes DA, Pagnoncelli D, Monteiro LC, Oliveira MV, Moreira MCF, Carvalho MS, Melão M, Klautau M, Rodriguez PRD, Costa RN, Silvano RG, Schientek S, Ribeiro SM, Pinheiro US, Hajdu E (2006) Capítulo 3. Filo Porifera. In: Lavrado HP, Ignácio BL (2006) Biodiversidade bentônica da
região central da Zona Econômica Exclusiva brasileira. Museu
Nacional, Série Livros 18, Rio de Janeiro. pp. 109-145
Peixinho S, Cosme B, Hajdu E (2005) Craniella quirimure sp. nov. from the mangroves of Bahia (Brazil) (Tetillidae, Spirophorida, Demospongiae). Zootaxa 1036: 31-42
Peixinho S, Fernandez J, Oliveira MV, Caíres S, Hajdu E (2007) Description of two new species of Acanthotetilla Burton, 1959 from NE Brazil, Southwestern Atlantic (Tetillidae, Spirophorida, Demospongiae). In: Custódio MR, Lôbo-Hajdu G, Hajdu E, Muricy G (eds). Porifera research: biodiversity, innovation and
sustainability. Série Livros 28. Museu Nacional, Rio de Janeiro.
pp. 509-515
Rützler K (1987) Tetillidae (Spirophorida, Porifera): a taxonomic reevaluation. In: Vacelet J, Boury-Esnault N (eds). Taxonomy of
Porifera from the N.E. Atlantic and Mediterranean Sea. NATO
ASI Series G 13. Springer-Verlag, London. pp.187-203
Rützler K, Smith KP (1992) Guide to western Atlantic species of
Cinachyrella (Porifera: Tetillidae). Proc Biol Soc Wash 105:
148-164
Santos JP, Mothes B, Tenório DO, Cantarelli J (1999) Porifera (Demospongiae, Calcarea) entre os estados do Ceará e Pernambuco, Brasil. Taxonomia e distribuição. Trab Oceanogr UFPE 27(2): 49-60
Santos JP, Cantarelli J, Tenório DO (2002) Porifera do Estado de Pernambuco, Brasil. In: Tabarelli M, Silva JMC (eds). Diagnóstico
da biodiversidade de Pernambuco 2. Editora Massangana, Recife.
pp. 385-404
Santos CP, Hajdu E (2003) Redescription of Tetilla radiata Selenka from the Southwestern Atlantic (Porifera, Spirophorida, Tetillidae) and designation of its neotype. Rev Bras Zool 20(4): 637-642 Santos CP, Hajdu E, Muricy G (2004) An identification system for
common Demospongiae of the São Sebastião Channel Area, SW Atlantic, developed with the Linnaeus II software. Boll Mus Ist
Biol Univ Gen 68: 587-591
Schmidt O (1868) Die Spongien der küste von Algier. Mit nachträgen
zu den Spongien des Adriatischen Meeres (Drittes Supplement).
Wilhelm Englemann, Leipzig
Schmidt O (1870) Grundzüge einer Spongien fauna des Atlantischen
Gebietes. Wilhelm Engelmann, Leipzig
Selenka E (1879) Ueber einem Kieselschwamm von achtstrahligem Bau, und über Entwicklung der Schwammknospen. Zeitsch Wiss
Zool: 472-475
Sollas WJ (1885) A classification of the sponges. Ann Mag Nat Hist 5(16): 95-395
Sollas WJ (1886) Preliminary account of the tetracinellid sponges dredged by H.M.S ‘Challenger’ 1872-76. Part I. The Choristida.
Sci Proc Royal Dublin Soc 5: 177-199
Sollas WJ (1888) Report on the Tetractinellida collected by H.M.S. ‘Challenger’ during the years 1873-1876. Rep Sci Res Voy H.M.S
‘Challenger’, Zool 25: 1-458
Uliczka E (1929) Die tetraxone Schwämme Westindiens (auf Grund der Ergebnisse der reise Kükenthal-Hartmeyer). Zool Jahrb, Suppl 16: 35-62
van Soest RWM, Rützler K (2002) Family Tetillidae Sollas, 1886.
In: Hooper JNA, van Soest RWM (eds). Systema Porifera: a guide to the classification of sponges. Kluwer Academic/Plenum
Publishers, New York. pp. 85-98
Wilson HV (1925) Siliceous and horny sponges collected by the US Fisheries Steamer ‘Albatross’ during the Philippine Expedition, 1907-10. Bull US Nat Mus 100(2): 273-532
