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Diversity of marine bryozoans inhabiting demosponges in northeastern Brazil

Diversity of marine bryozoans inhabiting demosponges in northeastern Brazil

Profile image of Facelucia BarrosFacelucia BarrosProfile image of Ana Carolina AlmeidaAna Carolina Almeida

As primary or obligate sessile organisms, bryozoans depend upon a substratum resource that affects their abundance, distribution and diversity. These animals can colonize virtually any type of substratum, including other organisms and artificial structures. Associations between bryozoans and sponges are commonly reported in the literature, but there are few studies discussing the association between these two taxa in detail. Here we present data on the bryozoan community found on shallow-water sponges from Bahia coast, northeastern Brazil, including their taxonomic status, colony form and adaptative structures utilized by these bryozoans to grow on sponges. Twenty-one bryozoan species were found attached to the surface of sixteen species of sponges. Five new species of cheilostome bryozoans are described. A total of 105 colonies were studied and most of them are erect delicate branching (44 colonies) and encrusting patches (34 colonies). The majority of bryozoan colonies were attached to the surface of rugose-textured sponges (67 colonies; 61%). This suggests that bryozoans are more likely to settle on irregular and rough surfaces. Patches colonies were mainly attached to the portion of the sponge that was in contact with the seabed, and spot colonies were particularly found in spatial refuges, showing the preference of larvae to settle on shaded and less exposed substrata. Most erect bryozoans were attached to the lateral sponge surface, other colonies grew on the underside and few on the upper surface of the sponges. These colonies were attached either using anchoring rhizoids, rigid bases, or stolons. The bryozoan species and genera reported here are common in northeastern Brazil and considered generalists in terms of larval settlement requirements. The bryozoan-sponge association studied is considered a non-obligatory commensalism (inquilinism).

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Zootaxa 4290 (2): 281–323 http://www.mapress.com/j/zt/ ISSN 1175-5326 (print edition) Article Copyright © 2017 Magnolia Press ZOOTAXA ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4290.2.3 http://zoobank.org/urn:lsid:zoobank.org:pub:0AE2706B-F77D-4903-B3A6-BB11891CD67B Diversity of marine bryozoans inhabiting demosponges in northeastern Brazil ANA C.S. ALMEIDA1,2, FACELUCIA B.C. SOUZA2, CARLA MENEGOLA3 & LEANDRO M. VIEIRA1 1 Laboratório de Estudos de Bryozoa—LAEBry, Departamento de Zoologia, Centro de Ciências Biológicas, Universidade Federal de Pernambuco, Recife, PE, 50670–810, Brazil. E-mail: carol.salmeida@gmail.com 2 Museu de Zoologia da Universidade Federal da Bahia, Universidade Federal da Bahia, Salvador, BA 40170–290, Brazil. 3 Laboratório de Biologia de Porifera e Fauna Associada, Departamento de Zoologia, Instituto de Biologia, Universidade Federal da Bahia, Salvador, BA, 40170-290, Brazil. Abstract As primary or obligate sessile organisms, bryozoans depend upon a substratum resource that affects their abundance, distribution and diversity. These animals can colonize virtually any type of substratum, including other organisms and artificial structures. Associations between bryozoans and sponges are commonly reported in the literature, but there are few studies discussing the association between these two taxa in detail. Here we present data on the bryozoan community found on shallow-water sponges from Bahia coast, northeastern Brazil, including their taxonomic status, colony form and adaptative structures utilized by these bryozoans to grow on sponges. Twenty-one bryozoan species were found attached to the surface of sixteen species of sponges. Five new species of cheilostome bryozoans are described. A total of 105 colonies were studied and most of them are erect delicate branching (44 colonies) and encrusting patches (34 colonies). The majority of bryozoan colonies were attached to the surface of rugose-textured sponges (67 colonies; 61%). This suggests that bryozoans are more likely to settle on irregular and rough surfaces. Patches colonies were mainly attached to the portion of the sponge that was in contact with the seabed, and spot colonies were particularly found in spatial refuges, showing the preference of larvae to settle on shaded and less exposed substrata. Most erect bryozoans were attached to the lateral sponge surface, other colonies grew on the underside and few on the upper surface of the sponges. These colonies were attached either using anchoring rhizoids, rigid bases, or stolons. The bryozoan species and genera reported here are common in northeastern Brazil and considered generalists in terms of larval settlement requirements. The bryozoan-sponge association studied is considered a non-obligatory commensalism (inquilinism). Key words: Atlantic Ocean, Bryozoa, Cheilostomata, commensalism, Cyclostomata, new species Introduction Bryozoans are primary or obligate sessile organisms ubiquitous in the marine benthic environment. Thus, the occurrence of these animals is directly dependent upon a substratum resource that, ultimately, affects their abundance, distribution and diversity (Hayward & Ryland 1998; Kuklinski & Barnes 2005). Virtually any type of substratum can be colonized by bryozoans, including other marine organisms, rocks, shells and rhodoliths (e.g., Gordon 1972; Lindberg & Stanton 1988; Creary 2002; Dick et al. 2006; Kuklinski & Barnes 2005; Denisenko et al. 2013; Souto et al. 2014), as well as artificial substrata, such as marine debris and other man-made structures (e.g., Winston 1982a; Carter & Gregory 2005; Vieira & Migotto 2014; Almeida et al. 2015a). Many studies deal with the association between bryozoans and algae (e.g., Ryland 1962; O'Connor et al. 1979; Manríquez & Cancino 1996; Lippert et al. 2001; Yagunova & Ostrovsky 2008), and investigations of the bryozoan fauna found on corals and inorganic substrata have also been carried out (e.g., Hayward 1974; Winston 1986; Hughes & Jackson 1992; Winston & Vieira 2013). Within many benthic communities in tropical and temperate regions, sponges often represent a dominant component that commonly serves as a living substratum for other organisms, including bryozoans (Klitgaard 1995; Cerrano et al. 2006). Several types of relationships between these animals are known, from epibiosis to commensalisms and mutualism (Cerrano et al. 2006), and the composition of the sponge-associated fauna is Accepted by K. Fehlauer-Ale: 5 May 2017; published: 7 Jul. 2017 281 influenced by intrinsic and extrinsic factors, including the internal and external sponge morphology and environmental conditions such as seafloor characteristics and water depth (e.g., Klitgaard 1995; Ávila & OrtegaBastida 2014). Owing to the intense spatial competition between members of the sessile benthic community, which is frequently dominated by fast-growing organisms such as sponges or algae, most bryozoans live in cryptic habitats. Bryozoans may also use the surface of invertebrates, such as sponges, as a protective suitable substratum for larval settlement and colonial development (Gordon 1972; Jackson 1977; Creary 2002; Berning et al. 2009). The association between bryozoans and sponges is frequently reported in the literature (e.g., Cook 1968; Winston 1982b; Wendt et al. 1985; Gischler & Ginsburg 1996; Ribeiro et al. 2003; Tilbrook 2001, 2006; Vieira et al. 2012), although the publications often consist merely of records of bryozoans attached to the sponge surface. For example, Vieira et al. (2012) listed 10 species of bryozoans from the Brazilian coast attached to sponges, but the identity of the sponges remained unknown. Harmelin et al. (1994) first described in detail the association between a bryozoan and a sponge but, in that case, the bryozoan Smittina cervicornis (Pallas, 1766) was serving as a substratum to the encrusting species of Demospongiae Halisarca cf. dujardini. Klitgaard (1995) first discussed a bryozoan-sponge association based on the morphology of these two taxa and suggested that bryozoans with erect colonies, rather than encrusting species, are commonly found attached to sponge surfaces, and also that sponges represent a hard substratum in areas with soft bottom. Results from Padua et al. (2013) corroborate these notions. Despite providing valuable information between bryozoan-sponge associations, these studies provide no descriptions or illustrations of the studied bryozoans, and the discussion of the relationships is very brief. Here we characterize 21 species of bryozoans inhabiting sponges from Bahia State, northeastern Brazil. Five new species of cheilostomes are also described. The bryozoan-sponge association is discussed in view of the bryozoan colony growth form and the rugosity of the sponge surface. Material and methods Study area and sampling procedures. We analyzed sponges deposited in the Porifera Collection of the Museu de Zoologia at Universidade Federal da Bahia (UFBA). These sponges specimens were collected in two bays along the coast of Bahia coast, northeastern Brazil—Todos os Santos Bay (12º35’S–13º07’S and 38º29’W–38º48’W) and Camamu Bay (13º50’S–14º06’S and 38º57’W–38º4’W) (Fig. 1). At Todos os Santos Bay the bottom is composed mainly by terrigenous and biogenic components originated from rock breakdown and metabolic activity of marine organisms, respectively (Alves et al. 2006; Lessa & Dias 2009). Siliciclastic mud from continental origin is the main component of the seabed at Camamu Bay, but quartz sand and coralline algae are also common (Hatje et al. 2008; Paixão et al. 2010). Sponges are important components of the seabed in both bays, leading to the recognition of these areas as “paradise of sponges” (Hajdu et al. 2011). All sponges were manually collected between 2012–2013 by personnel at LABPOR-UFBA (Laboratório de Biologia de Porifera, Universidade Federal da Bahia) by scuba diving between 3–20 m depth and preserved in plastic bags with 80% ethanol. At the laboratory and under a stereomicroscope, bryozoan colonies were carefully removed from the sponge surface and provisionally preserved in 70% ethanol before drying for taxonomic identification. Among all sampled sponges, we selected only one specimen of each sponge species where bryozoan colonies were visible to the naked eye. Sponges were identified after detailed study of their spicules and skeleton architecture, following the protocols described by Hajdu et al. (2011). The rugosity of the external surface of the sponges was inferred based on the general aspect of preserved specimens, being designated as rugose or smooth-textured (following Ward & Thorpe 1989). Rugose-textured sponges have an irregular external surface, bearing reentrances and concavities, and/or have a hispid to conular general aspect. Smooth-textured sponges have uniform external surfaces without any type of irregularities. Bryozoan identification and classification of colony growth forms. Bryozoan colonies were first examined under a stereomicroscope. Selected portions were mounted on stubs and coated with gold for examination by scanning electron microscopy (JEOL JSM-6390LV) and correspond to the figured colonies. The ctenostome Amathia distans Busk, 1886 was photographed under a stereomicroscope using a coupled camera (Olympus E330). When necessary, measurements were taken from digital SEM images using the software ImageJ®. Only species poorly known or new were fully described, and remarks for taxa recently described are provided. Specimens are 282 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. FIGURE 1. Map of the study area in Bahia State (BA), northeastern Brazil. Sponges were collected at Todos os Santos Bay and Camamu Bay. BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 283 deposited at the Bryozoa collection of the Museu de Zoologia at Universidade Federal da Bahia (UFBA). Each lot contains a single bryozoan colony. The growth form of each bryozoan colony was classified following Bishop (1989) and Taylor & James (2013) (Figs. 2–7). Encrusting species were classified as patches, to medium-sized layers of incrustation that never occupy the whole available substrata, and spots, to multiserial colonies sexually mature with few millimetres in diameter. Erect species were assigned as articulated, delicate branching, fenestrate and stoloniferan. Articulated bryozoans are characterized by skeletal elements joined by elastic joints or nodes; delicate branching are colonies formed by bifurcated branches that are usually bushy; fenestrate colonies are narrow branches linked at regular intervals and with feeding zooids opening on only one side of the colony; and stoloniferan was used in cases that the autozooids are linked by stolons (Taylor & James 2013). FIGURES 2–7. Bryozoan colony growth forms. 2, encrusting patches; 3, encrusting spots; 4, erect articulated; 5, erect delicate branching; 6, erect fenestrate; 7, erect stoloniferan. Scale bars: 5 mm. 284 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. TABLE 1. Bryozoans associated with sponges from Bahia State, northeastern Brazil. New species are marked with an asterisk. Numbers in parentheses represent the number of bryozoan colonies found attached to the studied sponges. Sponges with rugose-textured surface: Bu, Bubaris sp.; Ca, Callyspongia sp.; De, Dysidea etheria de Laubenfels, 1936; If, Ircinia felix (Duchassaing & Michelotti, 1864); Ma, Mycale angulosa (Duchassaing & Michelotti, 1864); Pw, Petrosia (Petrosia) weinbergi van Soest, 1980; Sp, Spongosorites sp.; Ti, Tedania ignis (Duchassaing & Michelotti, 1864). Sponges with smooth-textured surface: Cn, Chondrilla nucula Schmidt, 1862; Da, Desmapsamma anchorata (Carter, 1882); Ha, Halichondria sp.; Hm, Haliclona (Soestella) melana Muricy & Ribeiro, 1999; Mm, Myrmekioderma sp.; Tm, Timea sp.; To, Topsentia ophiraphidites (de Laubenfels, 1934); Tp, Topsentia sp. Bryozoans Sponges Rugose surface Smooth surface Bu Ca De If Ma Pw Sp Ti Cn Da Ha Hm Mm Tm To Tp Celleporaria atlantica (7) 3 0 0 0 0 0 0 0 0 0 0 3 0 0 1 0 Celleporaria carvalhoi (4) 0 0 0 0 0 0 0 0 0 0 2 1 0 0 0 1 Calyptotheca ornatissima n. comb. (3) 0 0 0 0 0 0 0 0 1 0 0 0 0 2 0 0 Hippaliosina imperfecta (1) 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 Metrarabdotos jani (4) 0 0 0 0 0 1 0 0 0 0 1 1 0 1 0 0 *Microporella curta (2) 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Poricella frigorosa (2) 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 Encrusting species (41) Patches (34) Reptadeonella leilae (1) 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 *Rhynchozoon brasiliensis (4) 2 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 Stylopoma aurantiacum (3) 0 0 0 1 0 0 0 0 0 0 0 2 0 0 0 0 *Turbicellepora iarae (3) 0 0 0 0 0 0 0 0 0 0 0 3 0 0 0 0 Calyptooecia conuma (1) 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 *Celleporina joannae (5) 0 0 0 0 0 0 5 0 0 0 0 0 0 0 0 0 *Marcusadorea pinheroi (1) 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 Crisia pseudosolena (5) 2 0 0 0 0 0 0 0 0 3 0 0 0 0 0 0 Nellia tenella (5) 3 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 Synnotum aegyptiacum (7) 5 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 Canda alsia (10) 3 0 0 0 0 0 0 4 0 0 0 3 0 0 0 0 Licornia aff. diadema (27) 7 6 0 0 3 0 0 3 0 5 0 3 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 Amathia distans (8) 0 3 2 0 3 0 0 0 0 0 0 0 0 0 0 0 Total (105) 25 13 2 1 10 1 6 9 1 8 4 17 2 4 1 1 Spots (7) Erect species (64) Articulated (10) Delicate branching (44) Fenestrate (2) Triphyllozoon arcuatum (2) Stoloniferans (8) Results Twenty-one bryozoan species were found associated to sixteen species of sponges (Table 1; Figs. 8–17). A single bryozoan species, Crisia pseudosolena (Marcus, 1937), belongs to the Class Stenolaemata Borg, 1926 and Order Cyclostomata Busk, 1852. The other 20 species are of the Class Gymnolaemata Allman, 1856—Amathia distans BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 285 Busk, 1886 belongs to the Order Ctenostomata Busk, 1852, and the remaining 19 species belong to the Order Cheilostomata Busk, 1852. We found 20 bryozoan species growing on sponges in Camamu Bay (15 new records), and only six species in Todos os Santos Bay (two new records) (Table 2). Thus, only 9 bryozoan species (43%) were previously reported from the studied area. Fourteen bryozoan species have encrusting colonies, from those 11 species are patches and only three species are spots (see Table 1). Seven bryozoan species have erect colonies, two of which are articulated (Crisia pseudosolena and Nellia tenella (Lamarck, 1816)), three are delicate branching (Synnotum aegyptiacum (Audouin, 1826)), Canda alsia Winston, Vieira & Woollacott, 2014 and Licornia aff. diadema (Busk, 1852)), one is fenestrate (Triphyllozoon arcuatum (MacGillivray, 1889)), and one species in stoloniferan (Amathia distans). FIGURES 8–12. Some rugose-textured sponges analyzed in the study. Arrows indicate the bryozoan attached to the sponges. 8, Bubaris sp.; 9, Mycale angulosa; 10, Petrosia (Petrosia) weinbergi; 11, Spongosorites sp.; 12, Tedanias ignis. Scale bars: 1 cm. 286 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. FIGURES 13–17. Some smooth-textured sponges analyzed in the study. Arrows indicate the bryozoans attached to the sponges. 13, Chondrilla nuculla; 14, Halichondria sp.; 15, Haliclona (Soestella) melana; 16, Timea sp.; 17, Topsentia sp. Scale bars: 1 cm. A total of 105 bryozoan colonies were studied: 41 (39%) are encrusting and 64 (61%) are erect. Most of the encrusting colonies form patches (34 colonies) and only seven colonies form spots. Within erect colonies, most of them are delicate branching (44 colonies), followed by articulated (10 colonies), stoloniferan (eight colonies), and fenestrate (two colonies). So, in general, in those taxa with erect species the number of colonies was greater (5–33 colonies per species) than those of encrusting species (1–8 colonies per species). The majority of the bryozoan colonies (64%; 67 of 105 colonies) were found on the surface of rugose-textured sponges. Most of the bryozoans are erect delicate branching (33 colonies) and encrusting patches (11 colonies). BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 287 Although with fewer colonies, encrusting spots (6 colonies), erect stoloniferan (8 colonies), articulated (7 colonies) and fenestrate (2 colonies) colonies were also found. Among the 38 bryozoan colonies found on smooth-textured sponges, the majority are encrusting patches (23 colonies), followed by erect delicate branching (11 colonies). Encrusting spots and erect articulated colonies were occasionally found on smooth-textured sponges with one and three colonies, respectively. TABLE 2. Bryozoans associated with sponges from Todos os Santos and Camamu bays. New records to the studied area are marked with an asterisk. Todos os Santos Bay Camamu Bay Amathia distans Busk, 1886 *Amathia distans Busk, 1886 Crisia pseudosolena (Marcus, 1937) Calyptotheca ornatissima n. comb. (Canu & Bassler, 1928a) Licornia aff. diadema (Busk, 1852) *Calyptooecia conuma Almeida & Souza, 2014 *Microporella curta n. sp. *Canda alsia Winston, Vieira & Woollacott, 2014 Poricella frigorosa Winston, Vieira & Wollaccott, 2014 *Celleporaria atlantica (Busk, 1884) *Rhynchozoon brasiliensis n. sp. Celleporaria carvalhoi (Marcus, 1939) Celleporina joannae n. sp. *Crisia pseudosolena (Marcus, 1937) Hippaliosina imperfecta (Canu & Bassler, 1928a) *Licornia aff. diadema (Busk, 1852) *Marcusadorea pinheroi n. sp. *Metrarabdotos jani Winston, Vieira & Woollacott, 2014 *Nellia tenella (Lamarck, 1816) *Poricella frigorosa Winston, Vieira & Wollaccott, 2014 *Reptadeonella leilae Almeida, Souza, Sanner & Vieira, 2015 *Rhynchozoon brasiliensis n. sp. *Stylopoma aurantiacum Canu & Bassler, 1928a *Synnotum aegyptiacum (Audouin, 1826) Triphyllozoon arcuatum (MacGillivray, 1889) *Turbicellepora iarae n. sp. Encrusting colonies were mostly found attached on the underside of the sponge (thus, in contact with the seabed), here assigned as the lower surface of the sponge. Whereas patches were found growing on the external lower surface, the spots colonies tend to occupy cryptic habitats in deep fissures at the lower surface. The species Calyptotheca ornatissima n. comb. was commonly found at either lower and upper surface of the sponge, frequently around it. The majority of the erect colonies were found attached laterally at the sponge surface (38 colonies), 19 colonies were found at the lower surface and only seven colonies were at the upper surface of the sponges. The greatest number of bryozoan colonies was registered on the rugose sponge Bubaris sp. (25 colonies) and on the smooth-textured sponge Haliclona (Soestella) melana Muricy & Ribeiro, 1999 (17 colonies). These species also hosted the highest bryozoan diversity, serving as a substratum to seven (Bubaris sp.) and eight bryozoan species (H. (S.) melana). Systematic account Phylum Bryozoa Ehrenberg, 1831 288 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. Class Stenolaemata Borg, 1926 Order Cyclostomata Busk, 1852 Family Crisiidae Johnston, 1838 Genus Crisia Lamouroux, 1812 Crisia pseudosolena (Marcus, 1937) (Figs. 18–21) Crisevia pseudosolena Marcus, 1937: 19, pl. 1, fig. 4B, pl. 2, figs. 4A–D. Crisia pseudosolena: Vieira et al. 2008: 34; Ramalho et al. 2009: 35, figs. 2A–F, 3A (cum syn.). Material examined. UFBA 1577, UFBA 2346–47, Todos os Santos Bay, 13°00’S, 38°32’W, 3–8 m, coll. 2013 (on sponge Desmapsamma anchorata); UFBA 1191, UFBA 2348, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Bubaris sp.). Remarks. Crisia pseudosolena is characterized by having zooids with longitudinally elongate pseudopores, transverse wrinkles in calcification near distal ends of peristomes, circular autozooidal apertures (Fig. 19) and a pear-shaped gonozooid lacking an ooeciostome (Fig. 21) (Marcus 1937; Ramalho et al. 2009). Ramalho et al. (2009) gave a complete characterization of C. pseudosolena based on specimens found attached to several invertebrates and also to unidentified sponges. In Brazil, at least a second species, Crisia cylindrica Busk, 1886, were previously reported growing on an unidentified sponge (Vieira et al. 2012). Here we found colonies of C. pseudosolena encrusting the sponges Desmapsamma anchorata and Bubaris sp. This is the first record of C. pseudosolena in Bahia State. Distribution. Atlantic: endemic to Brazil (Pernambuco, Bahia, Rio de Janeiro, São Paulo and Paraná) (Ramalho et al. 2009; present study). Class Gymnolaemata Allman, 1856 Order Ctenostomata Busk, 1852 Family Vesiculariidae Johnston, 1838 Genus Amathia Lamouroux, 1812 Amathia distans Busk, 1886 (Figs. 22–23) Amathia distans Busk, 1886: 33, pl. 7, fig. 1; Vieira et al. 2008: 10; Fehlauer-Ale et al. 2011: 56, figs. 3, 4, 6, 8, 10 (cum syn.); Migotto et al. 2011: 269; Marques et al. 2013: 271; Vieira et al. 2014: 514; Almeida et al. 2015b: 3. Material examined. UFBA 1581, UFBA 2349–50, Todos os Santos Bay, 13°00’S, 38°32’W, 3–8 m, coll. 2013 (on sponge Callyspongia sp.); UFBA 1600, UFBA 2351, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Dysidea etheria); UFBA 1617, UFBA 2352–53, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on Mycale angulosa). Remarks. Fehlauer-Ale et al. (2011) redescribed A. distans and stated the bright yellow pigment spots in stolonal and zooidal surfaces, the thickly cuticularised slender stolon (Fig. 22), and the autozooids organized in clockwise and anticlockwise directions (Fig. 23) as distinctive characters of the species. Amathia distans has been reported on a variety of substrata such as algae, bryozoans and anthropogenic surfaces (Fehlauer-Ale et al. 2011). Here we present the first record of A. distans associated with the sponges Callyspongia sp., Dysidea etheria and Mycale angulosa. BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 289 Distribution. Atlantic: Brazil (Alagoas, Bahia, Espírito Santo, Rio de Janeiro, São Paulo and Paraná) (Fehlauer-Ale et al. 2011). FIGURES 18–23. 18–21. Crisia pseudosolena, UFBA 1577. 18, Branching colony and rhizoid; 19, Close-up of autozooids; 20, Entire colony with gonozooid; 21, Close-up of gonozooid. 22–23. Amathia distans, UFBA 1578. 22, Entire colony; 23, Close-up of stolon and autozooids. Scale bars: 18, 20 = 500 µm; 19, 21–23 = 100 µm. Order Cheilostomata Busk, 1852 Family Quadricellariidae Gordon, 1984 290 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. Genus Nellia Busk, 1852 Nellia tenella (Lamarck, 1816) (Figs. 24–26) Cellaria tenella Lamarck, 1816: 135. Nellia oculata Busk, 1852: 18, pl. 64, fig. 6; pl. 65, fig. 4; Winston et al. 2014: 161, fig. 3 (cum syn.); Almeida et al. 2015b: 3; Taylor & Tan 2015: 14, fig. 7B–E. Material examined. UFBA 1602, UFBA 2354, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Mycale angulosa); UFBA 1616, UFBA 1666, UFBA 2355, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on Bubaris sp.). Comparative material. MNHN IB-2008-4546, Cellaria tenella, syntype, Lamarck det., Mers Australes?. Remarks. The taxonomic assignment of this species was recently discussed by Winston et al. (2014), who argued in favour of the usage of the name N. oculata instead of Nellia tenella (Lamarck, 1816) to specimens from Brazil, pending on the revision of type specimens of Cellaria tenella Lamarck. However, the examination of Lamarck’s Cellaria tenella (MNHN 4546; Fig. 26) shows that the main morphological characters are shared between both species, including the subquandrangular zooids, opesia occupying about three-quarters of surface, gymnocyst widest proximally, cryptocyst at proximal end of opesia, pores visible at the basal wall, a pair of very small oval avicularia placed proximolaterally on each zooid (Figs. 24 and 25) and ovicells endozooidal. Accordingly, as already notice by some authors (e.g., Cheetham 1966, Winston 2005) and following the principle of priority of ICZN (1999, Art. 23), Nellia oculata Busk must be considered junior synonym of Nellia tenella (Lamarck). Nellia tenella has a wide tropical and subtropical distribution (Taylor & Tan 2015) and is found in a broad range of marine environments (Winston et al. 2014). It was already found associated to unidentified sponges on the Brazilian coast (Vieira et al. 2012). Distribution. Circumtropical. Brazil: Rocas Atoll, Pernambuco, Bahia and Rio de Janeiro (Winston et al. 2014). Family Epistomiidae Gregory, 1893 Genus Synnotum Pieper, 1881 Synnotum aegyptiacum (Audouin, 1826) (Figs. 27–28) Loricaria aegyptiaca Audouin, 1826: 243; Savigny, [1817]: pl. 13, figs. 4.1–4.5. Synnotum aegyptiacum: Tilbrook 2006: 65, plate 8E–F (cum syn.); Gluhak et al. 2007: 399, figs. 3A–B; Vieira et al. 2008: 18; Almeida et al. 2015b: 3. Material examined. UFBA 1601, UFBA 2356–59 Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Bubaris sp.); UFBA 1615, UFBA 2360, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Mycale angulosa). Remarks. Synnotum aegyptiacum has erect articulated colonies formed by autozooids more or less fusiform with a large uncalcified frontal area, sessile short and robust avicularia placed at distal zooidal corners and pedunculate bulbous distal avicularia, placed between autozooids. This species is supposedly widely distributed in warm tropical waters, frequently found associated with algae, hydrozoans and other bryozoans, shell fragments and rocks (Osburn 1927; Marcus 1955; Shier 1964; Winston 1982b). Marcus (1955) reported colonies of S. aegyptiacum from southeast Brazil that were found anchored inside unidentified sponges. Colonies examined here were attached by rhizoids at the lateral surface of the rugose-textured sponges Bubaris sp. and Mycale angulosa. Distribution. Circumtropical. Brazil: Fernando de Noronha, Alagoas, Bahia, Espírito Santo and São Paulo (Vieira et al. 2008; Almeida et al. 2015b). BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 291 FIGURES 24–29. 24–25. Nellia tenella, UFBA 1602. 24, Close-up of autozooids; 25, Detail of autozooid and paired avicularia. 26. Nellia tenella, MNHN IB-2008-4546, syntype. Detail of autozooids showing gymnocystal and cryptocystal calcification, pores at the basal wall and paired small oval avicularia. 27–28. Synnotum aegyptiacum, UFBA 1601. 27, Autozooids with sessile and pedunculated avicularia; 28, Close-up of sessile avicularia. 29. Canda alsia, UFBA 1192, erect branch showing autozooids, scuta, spines, vibracular chambers and an ovicell. Scale bars: 24, 26, 29 = 200 µm; 25 = 50 µm; 27–28 = 100 µm. Family Candidae d’Orbigny, 1851 Genus Canda Lamouroux, 1816 292 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. Canda alsia Winston, Vieira & Woollacott, 2014 (Fig. 29) Canda philippinensis: Souza 1989: 497. Canda alsia Winston, Vieira & Woollacott, 2014: 164, figs. 15, 16 (cum syn.); Almeida et al. 2015b: 3. Material examined. UFBA 1192, UFBA 2361–62, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Bubaris sp.); UFBA 1663, UFBA 2363–64, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Haliclona (Soestella) melana); UFBA 1670, UFBA 2365–67 Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Tedania ignis). Remarks. Canda alsia has erect rooted colonies with branches connected by transverse tubular kenozooids, autozooids with one inner distal spine and a narrow mushroom-shaped scutum, short curving vibracula and ooecia with single and large fenestra (Fig. 29) (Winston et al. 2014). Despite the other records of Canda from the Brazilian coast—Canda philippinensis Canu & Bassler, 1929; Canda retiformis Pourtalès, 1867 and Canda simplex Busk, 1884 (Vieira et al. 2008)—Winston et al. (2014) suggested they belong to C. alsia (see Winston et al. 2014 for more information). Some specimens of C. alsia from Brazil were found attached on cable (Winston et al. 2014). Caribbean species of Canda are commonly found in coral reefs (e.g., Winston 1984, 1986; Hughes & Jackson 1992). Here we present the first record of a Canda species associated with sponges. Distribution. Atlantic: endemic to Brazil (Bahia, Espírito Santo and Rio de Janeiro) (Winston et al. 2014; present study). Genus Licornia van Beneden, 1850 Licornia aff. diadema (Busk, 1852) (Figs. 30–33; Table 3) ? Scrupocellaria diadema: Ramalho et al. 2005: 236; Ramalho et al. 2009: 36. Licornia diadema: Almeida et al. 2015b: 3. Material examined. UFBA 1172, UFBA 2368–69, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Mycale angulosa); UFBA 1173, UFBA 1190, UFBA 2370–74, Camamu Bay, 13°53’S, 38°59’W, 18– 20 m, coll. October 2012 (on sponge Bubaris sp.); UFBA 1574, UFBA 2375–79, Todos os Santos Bay, 13°00’S, 38°32’W, 3–8 m, coll. 2013 (on sponge Callyspongia sp.); UFBA 1576, UFBA 2380–83, Todos os Santos Bay, 13°00’S, 38°32’W, 3–8 m, coll. 2013 (on sponge Desmapsamma anchorata); UFBA 1613, UFBA 2384–85 Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Haliclona (Soestella) melana); UFBA 1614, UFBA 2386–87, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Tedania ignis). Additional comparative material. UFBA 223, Litoral Norte, Camaçari, 12º41’S, 38º05’W, 23 m, coll. February 2008; UFBA 288, Todos os Santos Bay, 13°00’S, 38°32’W, coll. 1997 by Orane Alves (specimens studied by Almeida et al. 2015b). Description. Colony erect, branched, with internodes comprising 6–16 zooids, living specimens pale brown to yellowish. Internode almost straight, with two series of autozooids, except at the axis of the bifurcation (triserial). Autozooids subrectangular, longer than wide, slightly tapering proximally, with rounded distal edges. Opesia oval, occupying more than three-quarters of the zooidal length, broader distally, surrounded proximally by a narrow and smooth cryptocyst. Scutum often seen in ovicelled zooids, inserted at midline of inner edge of the opesia, paddleshaped, covering small part of the frontal membrane. Four to five oral spines, frequently two inner, two outer and one median. Each zooid with a lateral avicularium placed at distal outer edge, small, rostrum triangular with serrated margins, directed laterally and slightly downward. Frontal avicularia axial zooid or near ovicelled zooids; avicularia of axial zooids placed below opesial margin, with raised base and obscuring proximal opesia, rostrum directed obliquely forward, with hooked tip; other zooids sometimes with a small avicularium at inner side just below opesia, rostrum triangular and with a hooked tip, directed proximo-medially. Vibracular chamber present on basal surface of each zooid, conspicuous in frontal view, also seen in axial zooid, almost triangular, with straight and oblique setal groove, with one proximal rhizoidal foramen. Vibracular setae smooth, as long as or longer than BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 293 two zooids. Axial zooid with single vibracular chamber. Rhizoids tubular with some retroussé hooks, usually only present at branch bases. Ooecia subglobular, with ectooecium perforated by 5–8 elliptical to circular pores with raised edge. FIGURES 30–35. 30–33. Licornia aff. diadema, UFBA 1190. 30, Frontal surface of branch; 31, Close-up of autozooids showing spines, lateral avicularia and vibracular chambers; 32, Close-up of ovicelled zooids showing scutum and frontal avicularia; 33, Abfrontal surface of branch. 34–35. Poricella frigorosa, UFBA 1585. 34, Overview of encrusting colony; 35, Group of zooids showing orificial spines, anchor-shaped frontal mucrones, frontal pores, avicularia and an ovicell. Scale bars: 30, 34 = 500 µm; 31, 33, 35 = 200 µm; 32 = 100 µm. Remarks. Licornia comprises at least 23 recent species mainly distributed in the Indian and Pacific Oceans (Vieira et al. 2013). Only five Licornia species have been reported in the Western Atlantic: Licornia diadema (Busk, 1852), Licornia drachi (Marcus, 1955), Licornia jolloisii (Audouin, 1826), Licornia micheli (Marcus, 1955) 294 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. and Licornia regularis (Osburn, 1940). The specimens here figured and described may be conspecific with other specimens previously reported from the Atlantic as L. diadema. Licornia diadema is characterized by having zooids with a variably shaped scutum, from club-shaped to rounded, with proximal and distal lobes of equal size, or slightly bifurcated to highly branched (Vieira et al. 2013). Recent studies, however, suggest that L. diadema comprises a species complex, and that many undescribed species were ascribed to this name (Tilbrook & Vieira 2012; Vieira et al. 2013, 2014; Sokolover et al. 2016). Thus, since the specimens here figured are distinct from the type specimen figured by Tilbrook & Vieira (2012), we prefer to use Licornia aff. diadema until the complete review of this group is done. Species of Licornia are commonly found attached to algae, corals and also artificial substrata (e.g., Creary 2002; Tilbrook & Vieira 2012; Vieira et al. 2013). Here we found several colonies of Licornia aff. diadema attached to both rugose (Fig. 9) and smooth-textured sponges. Distribution. Atlantic: Brazil (Bahia and possibly Rio de Janeiro) (Almeida et al. 2015b). TABLE 3. Measurements (mm) of Licornia aff. diadema (Busk, 1852) (UFBA 1574). Min: minimum; Max: maximum; N: number; SE: standard error. Structures N Min Max Mean SE Zooid length 15 0.394 0.534 0.475 0.048 Zooid width 15 0.187 0.239 0.215 0.012 Opesia length 15 0.282 0.330 0.375 0.024 Opesia width 15 0.100 0.178 0.156 0.019 Scuta length 15 0.112 0.167 0.128 0.017 Avicularia length 15 0.051 0.093 0.070 0.012 Avicularia width 15 0.029 0.049 0.040 0.005 Vibracular chamber length 15 0.136 0.210 0.189 0.017 Vibracular chamber width 15 0.063 0.074 0.063 0.009 Ovicell length 15 0.118 0.143 0.133 0.007 Ovicell width 15 0.185 0.226 0.205 0.013 Family Arachnopusiidae Jullien, 1888 Genus Poricella Canu, 1904 Poricella frigorosa Winston, Vieira & Wollaccott, 2014 (Figs. 34–35) Poricella frigorosa Winston, Vieira & Wollaccott, 2014: 186, fig. 30. Poricella mucronata: Almeida et al. 2015b: 4. Material examined. UFBA 1585, Todos os Santos Bay, 13°00’S, 38°32’W, 3–8 m, coll. 2013 (on sponge Callyspongia sp.); UFBA 2066, UFBA 2388, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Myrmekioderma sp.). Remarks. Poricella frigorosa has encrusting colonies (Fig. 34), autozooids with 3–4 orificial spines, an anchor-shaped frontal mucro, a central cluster of two or three round to oval pores, large interzooecial avicularia with a typically straight and pinched distal margin, and with a densely calcified ooecia opening above the operculum (Fig. 35) (Winston et al. 2014). Poricella frigorosa is part of the Poricella mucronata (Smitt, 1873) species complex (Winston et al. 2014). Until the description of P. frigorosa, P. mucronata and Poricella sp. were the only species recorded from the Brazilian coast (Vieira et al. 2008; Almeida et al. 2015b). Part of these specimens may also belong to P. frigorosa. No information about the substrata of the specimens reported from Brazil was given. Species of Poricella are known to encrust other bryozoans, coral skeletons, red algae, lava boulders and serpulid tubes (Tilbrook et al. 2001; Berning 2006; Dick et al. 2006; Moissette et al. 2007). BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 295 Distribution. Atlantic: endemic to Brazil (Bahia and Rio de Janeiro) (Winston et al. 2014). Family Adeonidae Busk, 1884 Genus Reptadeonella Busk, 1884 Reptadeonella leilae Almeida, Souza, Sanner & Vieira, 2015 (Figs. 36–37) Reptadeonella leilae Almeida, Souza, Sanner & Vieira, 2015c: 362, figs. 33–36, 42 (cum syn.). Material examined. UFBA 1174, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Halichondria sp.). Remarks. Reptadeonella leilae is characterized by unilaminar encrusting colonies (Fig. 36), autozooids with a tubular peristome, a circular spiramen and no avicularia (Fig. 37) (Almeida et al. 2015c). The species grows on small substrata such as calcareous nodules (Almeida et al. 2015c). Other Reptadeonella species are frequently found encrusting rocks, other bryozoans, calcareous nodules and algae, corals and shells (e.g., Marcus 1939; Soule 1961; Winston 1982b; Hayward & Ryland 1999; Tilbrook et al. 2001; Tilbrook 2006; Winston & Vieira 2013; Almeida et al. 2015c). Reptadeonella leilae was found attached to the smooth-textured sponge Halichondria sp., representing the first record of an association with sponges in the genus. Distribution. Atlantic: endemic to Brazil (Bahia) (Almeida et al. 2015c). Family Lepraliellidae Vigneaux, 1949 Genus Celleporaria Lamouroux, 1821 Celleporaria atlantica (Busk, 1884) (Figs. 38–41) Cellepora mamillata var. atlantica Busk, 1884: 199 (part), pl. 35, fig. 4. Celleporaria atlantica: Winston et al. 2014: 191, figs. 33A-D, 34, 35 (cum syn.); Almeida et al. 2015b: 4. Material examined. UFBA 1180, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Topsentia ophiraphidites); UFBA 1181, UFBA 2389–90, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Haliclona (Soestella) melana); UFBA 1188, UFBA 2391–92, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Bubaris sp.). Remarks. Celleporaria atlantica was recently redescribed by Winston et al. (2014), who studied original specimens from Busk (1884). Although Busk (1884) described C. atlantica from Brazil (Bahia state) and Australia (Possession Island), Winston et al. (2014) elucidated that specimens from Possession Island truly belong to Celleporaria fusca (Busk, 1854) and thus C. atlantica is restricted to Brazilian waters. Busk (1884) found C. atlantica on seafloors composed of gravel and shells. Other species of Celleporaria are commonly found associated with corals and rocks (e.g., Winston 1986; Winston 2005). Celleporaria atlantica has encrusting colonies, autozooids with almost rounded orifice with a shallow and wide sinus, suboral avicularium placed on a pointed umbo (Fig. 40), small leaf-shaped avicularia associated with marginal areolar pores and spatulate avicularia occurring between zooids (Fig. 41) (Winston et al. 2014). Celleporaria atlantica is often found at Bahia state forming nodular colonies attached to hard substrata such as rocks and calcareous nodules. Colonies of C. atlantica here studied are unilaminar and were encrusting the rugose sponge Bubaris sp. (Fig. 8) as well as the smooth-textured sponges Haliclona (Soestella) melana and Topsentia ophiraphidites. Distribution. Atlantic: endemic to Brazil (Bahia and Espírito Santo) (Winston et al. 2014). 296 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. FIGURES 36–41. Figures 36–37. Reptadeonella leilae, UFBA 1174. 36, Overview of encrusting colony with some gonozooids; 37, Close-up of autozooids showing tubular peristome. 38–41, Celleporaria atlantica, UFBA 1188. 38, Overview of encrusting colony; 39–40, Close-up of autozooids showing suboral avicularia, pointed umbos and leaf-shaped frontal avicularia; 41, Spatulate avicularium. Scale bars: 42 = 500 µm; 43–45 = 100 µm. Scale bars: 36, 38 = 500 µm; 37 = 200 µm; 39–41 = µm. Celleporaria carvalhoi (Marcus, 1939) (Figures 42–45; Table 4) Holoporella carvalhoi Marcus, 1939: 158, pl. 12, fig. 23A–D; Marcus 1949: 2. Celleporaria carvalhoi: Souza 1989: 499; Vieira et al. 2008: 16; Almeida et al. 2015b: 4. BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 297 FIGURES 42–47. 42–45. Celleporaria carvalhoi, UFBA 1610. 42, Group of autozooids with suboral umbos and frontal avicularia; 43, Autozooid showing D-shaped primary orifice and frontal avicularia with lanceolate rostrum; 44, Autozooids showing primary orifices and oral spines; 45, Autozooids and large interzooidal avicularia. 46–47. Metrarabdotos jani, UFBA 1606. 46, Overview of encrusting colony; 47, Autozooids showing pseudosinus, latero-oral avicularia, and a gonozooid (left). Scale bars: 42, 46–47 = 500 µm; 43 = 100 µm; 44 = 200 µm; 45 = 250 µm. Material examined. UFBA 1610, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Haliclona (Soestella) melana); UFBA 1611, UFBA 2393, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Halichondria sp.); UFBA 1945, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Topsentia sp.). Description. Colony encrusting, multilaminar. Zooids primarily subrectangular becoming irregularly polygonal, convex, limited by slightly raised lateral walls, frontal budded zooids irregularly oriented. Frontal shield 298 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. heavily calcified, rugose, becoming nodular, imperforate except for 4–10 large marginal pores. Primary orifice occupying about one-third of zooidal length, D-shaped, wider than long, sunken, with arcuate and broad distal edge and an almost straight proximal border. Two to four oral spines often seen in young zooids. Condyles and suboral avicularium absent. Secondary orifice transversely elliptical, raised more proximally than distally, heavily calcified, forming a suboral tapering umbo, frequently obscuring the primary orifice. Frontal avicularia small, frequently placed near zooidal margin, often disto-laterally directed, complete crossbar, lanceolate rostrum, proximal edge rounded, calcified palate occupies distal third of rostral length; distal edge forming a more or less acute tip. Large interzooidal avicularia, longer than wide, with complete crossbar, spatulate rostrum, palate calcified in distal part with rounded proximal edge. Ovicells not seen. Remarks. Celleporaria carvalhoi is here characterized using SEM images for the first time since its original description by Marcus (1939). This species has zooids with large marginal pores, a D-shaped primary orifice obscured by a peristome with a suboral tapering umbo, adventitious avicularia with a lanceolate rostrum and large avicularia somewhat spatulate in form. Specimens studied by Marcus (1939) were found encrusting corals. Here we found colonies of C. carvalhoi attached to both rugose and smooth-textured sponges. Distribution. Atlantic: endemic to Brazil (Bahia, Espírito Santo and São Paulo) (Vieira et al. 2008). TABLE 4. Measurements (mm) of Celleporaria carvalhoi (Marcus, 1939) (UFBA 1610). Min: minimum; Max: maximum; N: number; SE: standard error. Structures N Min Max Mean SE Zooid length 15 0.308 0.470 0.376 0.048 Zooid width 15 0.279 0.396 0.334 0.035 Orifice length 15 0.083 0.141 0.104 0.016 Orifice width 15 0.117 0.175 0.141 0.020 Frontal avicularia length 15 0.081 0.132 0.105 0.016 Frontal avicularia width 15 0.042 0.079 0.059 0.012 Family Metrarabdotosidae Vigneaux, 1949 Genus Metrarabdotos Canu, 1914 Metrarabdotos jani Winston, Vieira & Woollacott, 2014 (Figs. 46–47) Metrarabdotos unguiculatum: Souza 1989: 499. Metrarabdotos jani Winston, Vieira & Woollacott, 2014: 198, fig. 38 (cum syn.). Material examined. UFBA 1606, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Timea sp.); UFBA 1607, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Haliclona (Soestella) melana); UFBA 1176, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Petrosia (Petrosia) weinbergi); UFBA 1177, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Halichondria sp.). Remarks. Metrarabdotos jani was previously misidentified as Metrarabdotos unguiculatum (Canu & Bassler, 1928b) (Marcus 1955; Santana et al. 2009), but Winston et al. (2014) distinguished them by the ooecium (centrally imperforate in M. jani and porous in M. unguiculatum). Other distinct characters of M. jani are the encrusting colonies, large elongate zooids, orifice with U-shaped sinus and single to paired latero-oral avicularia placed below the orifice and with rostrum reaching the edge of the peristome (Fig. 47) (Winston et al. 2014). Some Metrarabdotos species were found attached to seagrass rhizomes (Cheetham et al. 2007), but the association between the genus and sponges is here reported for the first time. Metrarabdotos jani was found attached to the rugose-textured sponge Petrosia (Petrosia) weinbergi van Soest, 1980 and to the smooth-textured sponges Halichondria sp. (Fig. 14), Haliclona (Soestella) melana and Timea sp. Distribution. Atlantic: endemic to Brazil (Bahia, Espírito Santo and Rio de Janeiro) (Winston et al. 2014). BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 299 Family Lanceoporidae Harmer, 1957 Genus Calyptotheca Harmer, 1957 Calyptotheca ornatissima (Canu & Bassler, 1928a) n. comb. (Figs. 48–53; Table 5) Gemelliporidra ornatissima Canu & Bassler, 1928a: 79, pl. 5, figs. 1–2; Almeida et al. 2015b: 5. FIGURES 48–53. Calyptotheca ornatissima n. comb. 48–51, UFBA 1178. 48, Overview of encrusting colony; 49, Close-up of primary orifice showing lunula, suboral and marginal avicularium; 50, Group of autozooids and interzooidal avicularium (center); 51, Ovicelled zooids showing orificial dimorphism. 52–53, USNM 8554, holotype. 52, Detail of autozooidal primary orifice and avicularia; 53, Ovicelled zooids (notice the cormidial calcification) and interzooidal avicularium (top right). Scale bars: 48 = 500 µm; 49, 51–52 = 100 µm; 50 = 200 µm; 53 = 250 µm. 300 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. Material examined. UFBA 605, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Timea sp.); UFBA 1178, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Chondrilla nucula); UFBA 1179, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Timea sp.). Comparative material. USNM 8554, Gemelliporidra ornatissima, holotype, F. Canu & R. Bassler det., Bahia, Brazil, 49 m, coll. 1877 by Steamer Norseman. Description. Colony encrusting, orange in life. Zooidal skeleton orange and avicularia white. Autozooids almost quadrangular to rectangular, rarely polygonal, limited by distinct raised walls. Frontal shield heavily calcified, tuberculate, entirely punctured by 20–50 pseudopores. Primary orifice small relative to frontal shield length, dimorphic (shorter in ovicelled zooids than infertile ones), subcircular, sunken, rounded anter separated from widely and shallowly V-shaped poster by two strong rounded condyles placed at the proximal quarter of orifice; lunula restricted to distal edge of the orifice. Proximal border of orifice slightly raised, often surrounded by short tubercles and with a suboral umbo; suboral umbo often carrying a laterally oriented avicularium with an elongated triangular rostrum. Frontal avicularia similar in shape to suboral avicularium, common, scattered throughout the colony, frequently near zooidal margins and also above ovicells. One elongate avicularium often placed at latero-distal zooidal margin, directed distally or proximally, rostrum acute and curved, with complete crossbar; sometimes a curved avicularium seen at proximal zooidal margin. Vicarious avicularium large, longer than wide, rostrum somewhat spatulate, oblong with concave lateral edges; palate calcified in distal third with rounded proximal margin, thinning proximally along lateral edges; mandible hinged on two strong square condyles situated in the corners of the straight proximal end of avicularium; cystid surrounding almost the entire avicularium except in the rounded distal edge, frontal shield similar to autozooid. Ovicell prominent, ooecia globose, same tuberculate and porous calcification as autozooids, secondary calcification cormidial (i.e. with Y-shaped suture lines of calcification), closed by zooidal operculum. Orifice dimorphic, wider than in autozooids. TABLE 5. Measurements (mm) of Calyptotheca ornatissima n. comb. (Canu & Bassler, 1928) (UFBA 605). Min: minimum; Max: maximum; N: number; SE: standard error. Structures N Min Max Mean SE Zooid length 15 0.464 0.711 0.561 0.082 Zooid width 15 0.320 0.551 0.418 0.061 Orifice length 15 0.121 0.164 0.142 0.010 Orifice width 15 0.139 0.164 0.149 0.007 Suboral avicularia length 15 0.093 0.120 0.101 0.008 Suboral avicularia width 15 0.043 0.072 0.054 0.008 Frontal avicularia length 15 0.086 0.102 0.138 0.016 Frontal avicularia width 15 0.037 0.066 0.050 0.007 Lateral avicularia length 15 0.113 0.151 0.134 0.009 Lateral avicularia width 15 0.022 0.034 0.027 0.003 Interzooidal avicularia length 4 0.404 0.503 0.476 0.044 Interzooidal avicularia width 4 0.157 0.181 0.167 0.010 Ovicell length 3 0.312 0.338 0.325 0.018 Ovicell width 3 0.374 0.391 0.382 0.011 Remarks. Calyptotheca ornatissima n. comb. has not been figured since its original description (Canu & Bassler 1928a), and only recently it has been reported again from Brazilian coast (Bahia) by Almeida et al. (2015b). Although it was originally assigned to the genus Gemelliporidra Canu & Bassler, 1927, the morphology of orifice (proximal sinus, condyles, lunula and dimorphism; figs. 49, 51, 52 and 53), avicularia (both adventitious and vicarious; figs. 50–53), and ovicell (calcification similar to autozooidal frontal shield; secondary calcification cormidial; figs. 51 and 53) (Cumming & Tilbrook 2014; Cumming 2015) suggest that it belongs to the genus Calypthoteca. The species is characterized by multilayered orange colonies with white spots representing the avicularia and BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 301 ovicells (Canu & Bassler 1928a; figs. 52 and 53). It can be distinguished from other Calyptotheca by its avicularia (suboral, frontal, lateral and vicarious; Figs. 49–50). Other species of the genus were already reported in association with corals, several invertebrates such as other bryozoans and gorgonians, shells, rocks and kelps (e.g., Winston 1986; Florence et al. 2007; Cumming & Tilbrook 2014). Colonies of Calyptotheca ornatissima n. comb. examined here are robust and covered lower and upper portions of the smooth-textured sponges Chondrilla nucula Schmidt, 1862 (Fig. 13) and Timea sp. (Fig. 16), representing the first record of the association of Calyptotheca ornatissima n. comb. with sponges. Distribution. Atlantic: endemic to Brazil (Bahia) (Vieira et al. 2008). FIGURES 54–59. Stylopoma aurantiacum, UFBA 1182. 54, Overview of encrusting colony; 55–57. Close-up of autozooids showing primary orifices and frontal avicularia; 58, Shoe-shaped interzooidal avicularia; 59, Ovicell with crab claw–like extensions. Scale bars: 54 = 500 µm; 55, 57, 59 = 100 µm; 56 = 50 µm; 58 = 200 µm. 302 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. Family Schizoporellidae Jullien, 1883 Genus Stylopoma Levinsen, 1909 Stylopoma aurantiacum Canu & Bassler, 1928a (Figs. 54–59) Stylopoma aurantiacum Canu & Bassler, 1928a: 78, pl. 4, figs. 3, 4; Vieira et al. 2008: 28; Winston et al. 2014: 204, fig. 41. Material examined. UFBA 1587, UFBA 2394, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Haliclona (Soestella) melana); UFBA 1182, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (one colony on sponge Ircinia felix). Remarks. Stylopoma aurantiacum has encrusting colonies (Fig. 54), a keyhole-shaped orifice with distinct condyles, frontal avicularia rounded proximally and subtriangular distally (Fig. 56), large shoe-shaped interzooidal avicularia (Fig. 58) and large ooecia with crab claw-like extensions from either side of the base of the opening (Fig. 59) (Canu & Bassler 1928a; Winston et al. 2014). Winston et al. (2014) first reported S. aurantiacum since its original description, but here we first figure ooecia of S. auratiacum (Fig. 59). We found two colonies attached to the sponges Haliclona (Soestella) melana and Ircinia felix. The widely known Stylopoma spongites (Pallas, 1766) received its name owing to its association with a sponge (Tilbrook 2001), suggesting Stylopoma species might be common on that substratum. Distribution. Atlantic: endemic to Brazil (Pernambuco and Bahia) (Winston et al. 2014). Family Hippaliosinidae Winston, 2005 Genus Hippaliosina Canu, 1918 Hippaliosina imperfecta (Canu & Bassler, 1928a) (Figs. 60–61) Gephyrophora imperfecta Canu & Bassler, 1928a: 86, pl. 7, fig. 1. Hippaliosina imperfecta: Winston et al. 2014: 209, fig. 44 (cum syn.); Almeida et al. 2015b: 5. Material examined. UFBA 2065, UFBA 2395, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Myrmekioderma sp.). Remarks. Hippaliosina imperfecta is characterized by having encrusting colonies (Fig. 60), more or less hexagonal zooids, oblong autozooidal orifices, and small down-curved and elongate avicularia (frequently paired) placed laterally to the orifice (Fig. 61) (Winston et al. 2014). Hippaliosina imperfecta is the only species of the genus recorded from Brazil and is also considered endemic to the area (Winston et al. 2014). Other species of Hippaliosina were already documented in association with organisms such as other bryozoans, molluscs and algae (Osburn 1952; Shier 1961; Hayward 1974; Winston 1982b, 1986). This is the first record, however, of a Hippaliosina species associated with the smooth-textured sponges Myrmekioderma sp. and Desmapsamma anchorata. Distribution. Atlantic: endemic to Brazil (Bahia, Espírito Santo and Rio de Janeiro) (Winston et al. 2014). Family Marcusadoreidae Winston, Vieira & Woollacott, 2014 Genus Marcusadorea Vieira, Migotto & Winston, 2010 Marcusadorea pinheroi n. sp. (Figs. 62–65; Table 6) Material examined. Holotype: UFBA 1186, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 303 sponge Timea sp.). Paratype: UFBA 1946, Salvador, Bahia, Brazil, 12°57’S, 38°21’W, intertidal, coll. April 2012. Type locality. Camamu Bay, Bahia State, NE Brazil. Etymology. Named after Ulisses Pinheiro (Universidade Federal de Pernambuco), in recognition of his contribution to the knowledge of Brazilian biodiversity. FIGURES 60–65. 60–61. Hippaliosina imperfecta, UFBA 1187. 60, Overview of encrusting colony; 61, Close-up of autozooids showing orifices and avicularia. 62–65. Marcusadorea pinheroi n. sp., UFBA 1186 holotype, Bahia State, Brazil. 62, Overview of encrusting colony; 63, Close-up of autozooid; 64, Close-up of primary orifice showing the condyles; 65, Close-up of ovicelled zooid. Scale bars: 60, 62 = 500 µm; 61, 63, 65 = 200 µm; 64 = 100 µm. Description. Colony encrusting, spot-like, unilaminar. Colony pale yellow. Zooids semi-erect, large, globular, limited by raised distinct lateral walls. Frontal shield heavily calcified, granular, with small nodules of calcification, frontally punctured by 20–24 pseudopores except at the peristomial calcification that remains imperforate; marginally with a distinct row of 12–20 pores. Primary orifice large, hoof-shaped, with a pair of small 304 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. lateral triangular condyles. No oral spines. Secondary orifice raised, forming a well-developed tubular peristome with circular aperture that obscures the primary orifice, with same calcification as frontal shield but without pores; aperture oval in ovicelled zooids. Avicularia absent. Ovicell prominent; ooecia globose, 0.396 mm long and 0.649 mm wide, same granular, nodular and porous calcification as in autozooids, opening into the peristome above zooidal operculum. Remarks. Currently four Marcusadorea species are recognized—Marcusadorea corderoi (Marcus, 1949), Marcusadorea efatensis (Tilbrook, 2006), Marcusadorea jamaicensis Vieira, Migotto & Winston, 2010 and Marcusadorea tubulosa (Canu & Bassler, 1928b). Marcusadorea pinheroi n. sp. most closely resembles M. efatensis in having a frontal shield with numerous pseudopores, a subcircular primary orifice with small condyles, ooecia with tha same calcification as frontal shields, and no avicularia. Both differ, however, in the autozooids (semi-erect in M. pinheroi n. sp. and non-elevate in M. efatensis), development of the peristome (conspicuous in autozooids and ovicelled zooids of M. pinheroi n. sp., and especially well-developed in ovicelled zooids of M. efatensis), and the marginal pores (non-differentiated in M. pinheroi n. sp. and large and distinct in M. efatensis). Other species of Marcusadorea are readily distinguished from M. pinheroi n. sp. in having suboral avicularia (that are absent in M. pinheroi n. sp.), frontal walls with scattered pseudopores (entirely punctured in M. pinheroi n. sp.), and forming a pseudosinus (absent in M. pinheiroi). Marcusadorea pinheroi n. sp. is also the only species of the genus that has semi-erect autozooids. No information regarding substrate type used by Marcusodarea species was given, but M. tubulosa from NE Brazil is often seen on rocks and coral rubble (L.M. Vieira, unpub. data). Here we found colonies of M. pinheroi n. sp. attached to the smooth-textured surface of Timea sp. (Fig. 16). Distribution. Atlantic: Brazil (Bahia). TABLE 6. Measurements (mm) of Marcusadorea pinheroi n. sp. (UFBA 1186). Min: minimum; Max: maximum; N: number; SE: standard error. Structures N Min Max Mean SE Zooid length 6 0.862 1.051 0.972 0.007 Zooid width 6 0.696 0.949 0.840 0.010 Orifice length 6 0.354 0.450 0.378 0.044 Orifice width 6 0.291 0.379 0.335 0.036 Family Microporellidae Hincks, 1879 Genus Microporella Hincks, 1877 Microporella curta n. sp. (Figs. 66–71; Table 7) Material examined. Holotype: UFBA 1580, Todos os Santos Bay, 13°00’S, 38°32’W, 3–8 m, coll. 2013 (on sponge Callyspongia sp.). Paratype: UFBA 1582, Todos os Santos Bay, 13°00’S, 38°32’W, 3–8 m, coll. 2013 (on sponge Callyspongia sp.). Type locality. Todos os Santos Bay, Bahia State, NE Brazil. Etymology. From Latin curtus, short, alluding to the thin row of calcification around the primary orifice of non-ovicelled zooids. Description. Colony encrusting, uni- to multilamellar. Autozooids large, irregularly polygonal to hexagonal, separated by raised walls. Frontal shield slightly convex, with small rounded nodes and uniformly punctured by numerous rounded pseudopores except for the area between ascopore and orifice. Marginal pores more elongate than pseudopores. Primary orifice almost D-shaped, distal edge smooth and proximal border serrated with 12–18 triangular denticles. Autozooids with a thin row of calcification around the primary orifice, not obscuring it. Nonovicelled zooids with three or four oral spines. Ascopore situated proximal to orifice at a distance about half of orifice length, reniform, surrounded by a thin rim, with a rounded median process and a crescentic (C-shaped) BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 305 lumen, margins with regularly spaced sharp denticles. Avicularium single, placed proximolateral and below to the orifice and ascopore, directing distolaterally and slightly upwards, rostrum narrowing distinctly distal to complete crossbar, forming a chute-like strucure in distal half, uncalcified area proximal to crossbar semicircular. Sometimes the avicularium reaches the neighboring zooid. Ovicelled zooids with no visible oral spines; ooecia personate, i.e. with tall and thin granular collar, distally adjacent to ascopore but not obscuring it, raised over orifice and distally joined to thin, smooth, almost straight rim on proximal edge of ooecium forming a complete peristome. Remarkably large marginal pores easily seen around ooecia. Ovicells prominent; ooecium globose, closed by zooidal operculum; endooecial surface similar to autozooidal frontal shield. Aperture in ovicelled zooids transversally oval to quadrangular. FIGURES 66–71. Microporella curta n. sp., UFBA 1580, holotype, Bahia State, Brazil. 66, Group of autozooids; 67, Detail of autozooid with four oral spines and a single avicularium; 68, Detail of D-shaped primary orifice and reniform ascopore; 69, Detail of avicularium; 70, Zooids with personate ovicells; 71, Detail of ovicell. Scale bars: 66 = 200 µm; 67, 71 = 100 µm; 68, 69 = 50 µm; 70 = 250 µm. 306 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. Remarks. Among the 92 valid Microporella species (Bock 2016), Microporella curta n. sp. most resembles Microporella browni Harmelin, Ostrovsky, Cáceres-Chamizo & Sanner, 2011, Microporella collaroides Harmelin, Ostrovsky, Cáceres-Chamizo & Sanner, 2011, Microporella dentilingua Tilbrook, 2006, Microporella harmeri Hayward, 1988 and Microporella orientalis (Harmer, 1957) in having personate ovicells, a reniform denticulate ascopore, and a single avicularium placed the ascopore. Differences between Microporella curta n. sp. and M. browni are the number of oral spines (4 in Microporella curta n. sp. and 3–7 in M. browni), and the primary orifice (with smooth distal edge and without condyles in Microporella curta n. sp., and with serrated distal edge and with low condyles in M. browni). Microporella collaroides is distinct in having an orifice with a wavy distal edge and lateral condyles (primary orifice smooth distally and without condyles in M. curta n. sp.). Microporella curta n. sp. differs from M. dentilingua in having polygonal autozooids (roughly hexagonal in M. dentilingua), 4 oral spines (3 in M. dentilingua), and longer orifices (in M. dentilingua it is about 0.07 mm long). Microporella harmeri is distinct in having 2–5 oral spines that disappear in later astogeny (4 conspicuous oral spines in M. curta n. sp.), and a lanceolate avicularium mandible (setiform in M. curta n. sp.). Microporella orientalis can be distinguished from M. curta n. sp. in having 3 oral ephemeral spines (4 conspicuous in M. curta n. sp.). At least 12 species of Microporella are recorded in the Western Atlantic Ocean, three of which are reported from Brazil: Microporella cucullata Canu & Bassler, 1928a, Microporella proxima Ramalho, Muricy & Taylor, 2011 and Microporella umbracula (Audouin, 1826). Microporella curta n. sp. is distinct from M. cucullata in the secondary calcification around the orifice in the autozooids (in M. cucullata the secondary calficiation forms a hood-like structure), avicularia placement and direction (below the ascopore and obliquely directed in M. curta n. sp., and above the ascopore and laterally directed in M. cucullata), and smaller zooids (those from M. cucullata are 0.544 to 0.638 mm long and 0.438 to 0.700 mm wide). Microporella proxima differs in the frontal and ovicell calcification (smooth in M. curta n. sp. and pustulose in M. proxima), avicularia (placed laterally and below the ascopore in M. curta n. sp., and near zooidal margins in M. proxima), and in larger zooidal measurements (autozooids of Mi. proxima are 0.392 to 0.441 mm long and 0.294 to 0.343 mm wide, and orifices are 0.64 to 0.79 mm long and 0.93 to 0.107 mm wide). Microporella umbracula is readily distinguished from M. curta n. sp. by the paired, upward directed avicularia. We found large colonies of M. curta n. sp. firmly attached to the basal surface of the sponge Callyspongia sp. Other Microporella species are commonly found encrusting shells, corals, algae and hydrozoans (e.g., Canu & Bassler 1928a; Marcus 1937; Winston 1984, 2005; Harmelin et al. 2011; Ramalho et al. 2011). Distribution. Atlantic: Brazil (Bahia). TABLE 7. Measurements (mm) of Microporella curta n. sp. (UFBA 1580). Min: minimum; Max: maximum; N: number; SE: standard error. Structures N Min Max Mean SE Zooid length 15 0.433 0.605 0.519 0.053 Zooid width 15 0.335 0.529 0.390 0.046 Orifice length 15 0.074 0.107 0.083 0.007 Orifice width 15 0.098 0.125 0.111 0.007 Ascopore diameter 15 0.041 0.062 0.054 0.004 Avicularia length 15 0.129 0.165 0.146 0.012 Avicularia width 15 0.072 0.102 0.088 0.008 Ovicell length 15 0.192 0.262 0.216 0.018 Ovicell width 15 0.234 0.317 0.272 0.025 Family Cleidochasmatidae Cheetham & Sandberg, 1964 Genus Calyptooecia Winston, 1984 BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 307 Calyptooecia conuma Almeida & Souza, 2014 (Figs. 72–73) Calyptooecia conuma Almeida & Souza, 2014: 285, figs. 2–5; Almeida et al. 2015b: 5. Material examined. UFBA 1183, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Spongosorites sp.). Remarks. Calyptooecia conuma is characterized by small spot-like colonies (Fig. 72), brooding zooids without perioral tubercles, and non-brooding zooids surrounded by 4–5 conical tubercles and a single, asymmetrically placed, suboral avicularium on almost all zooids (Fig. 73) (Almeida & Souza 2014). Only two species of Calyptooecia are known worldwide (Bock 2016). Calyptooecia insidiosa Winston, 1984 is found in the Caribbean where colonies are associated with coral undersurfaces and reef caves (Winston 1984). Calyptooecia conuma commonly encrusts calcareous nodules, shells, coralline algae and sponges (Almeida & Souza 2014). We examined a small colony attached to recesses of the rugose-textured sponge Spongosorites sp. (Fig. 11). Distribution. Atlantic: endemic to Brazil (Bahia) (Almeida & Souza 2014). Family Celleporidae Johnston, 1838 Genus Celleporina Gray, 1848 Celleporina joannae n. sp. (Figs. 74–77; Table 8) Celleporina costazii: Almeida et al. 2015b: 5. Material examined. Holotype: UFBA 1664, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Spongosorites sp.). Paratypes: UFBA 1665, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Spongosorites sp.); UFBA 1671, UFBA 2396–97, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Spongosorites sp.). Type locality. Camamu Bay, Bahia State, NE Brazil. Etymology. Named after JoAnn Sanner (Smithsonian Institution, National Museum of Natural History), for her contribution to bryozoology. Description. Colony encrusting, multilaminar, forming small spherical nodules. Autozooids mound-like, globular to sub-erect. Frontal shield heavily calcified, smooth, marginally punctured by 6–8 large widely-spaced pores. Sometimes additional frontal pseudopores are seen surrounding the peristome, often in young zooids with low peristomes that expose the primary orifice. Primary orifice terminal, circular, sunken, with a broadly U-shaped sinus; condyles rounded, thickened and conspicuous. Secondary orifice formed by a deep tubular peristome, margin usually smooth but sometimes somewhat crenulated. Single or paired peristomial avicularia, proximolateral to the orificial border, placed at peristomial lobes; rostrum triangular, with complete cross-bar, margins smooth, oriented upwardly and distolaterally. Additional frontal avicularia absent. Ooecia slightly wider than long, frontal surface in level with peristome; tabula occupying most of the frontal surface, bordered by a single row of 8– 12 distolateral pores; ooecial surface with distinct ribs in later astogeny. Ooecia open into distal part of peristome, just above the operculum; aperture quadrangular, edged by a broad band of smooth ectooecium. Remarks. Celleporina joannae n. sp. can be distinguished from other species of Celleporina by the combination of a tubular peristome with 1–2 avicularia placed at lobes, a widely U-shaped sinus with strong rounded condyles, no frontal avicularia, and ooecia with a quadrangular aperture and a tabula bordered by a single row of 8–12 distolateral pores. At least eight species of Celleporina are reported to the Western Atlantic (Vieira et al. 2008; Bock 2016). Three of them are readily distinguished from C. joannae n. sp. in having ooecia with entirely porous tabula, viz. Celleporina diota (Marcus, 1938), Celleporina langei (Marcus, 1939) and Celleporina lucida (Hincks, 1880). Two other species, Celleporina abtrusa Winston & Vieira, 2013 and Celleporina bicostata Hayward, 1980, have smooth ooecial tabula. Celleporina joannae n. sp. most closely resembles Celleporina lacrimula Hayward, 1992 and 308 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. Celleporinas surcularis (Packard, 1863) in having ooecial pores commonly separated by distinct ribs, and a peristome with lobes. The species are distinct, however, in the primary orifice (drop-shaped in C. lacrimula and almost circular in C. surcularis; with a broadly U-shaped sinus and strong condyles in C. joannae n. sp.) and in their vicarious avicularia (present in C. lacrimula C. lacrimula and C. surcularis; absent in C. joannae n. sp.). FIGURES 72–77. 72–73. Calyptooecia conuma, UFBA 1183. 72, Spot-like colony; 73, Detail of autozooids showing brooding zooid (center) and non-brooding zooids surrounded by conical tubercles. 74–77. Celleporina joannae n. sp., UFBA 1664, holotype, Bahia State, Brazil. 74, Spot-like colony; 75, Semi-erect, non-oriented autozooids with peristomial avicularia and autozooid with primary orifice showing broad median sinus and thickened condyles (center); 76, Close-up of orifice showing deep tubular peristome and peristomial avicularium; 77, Close-up of ovicelled zooid. Scale bars: 72, 74 = 500 µm; 73, 75 = 200 µm; 76–77 = 100 µm. BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 309 Specimens from the Brazilian coast described by Marcus (1937, 1939, 1949) under the name Celleporina costazii (Audouin, 1826) differ from C. joannae n. sp. in having a median peristomial umbo and vicarious avicularia and ooecia becoming entirely porous. Species of Celleporina are commonly found encrusting shells, other bryozoans, corals and hydrozoans, worm tubes and algae (e.g., Cook 1985; Marcus 1938, 1939; Hayward & Ryland 1999; Ramalho et al. 2011; Winston & Vieira 2013). Here we found spot-like colonies with 2–3 mm in diameter encrusting the rugose-textured sponge Spongosorites sp. (Fig. 11). Distribution. Atlantic: Brazil (Bahia). TABLE 8. Measurements (mm) of Celleporina joannae n. sp. (UFBA 1664). Min: minimum; Max: maximum; N: number; SE: standard error. Structures N Min Max Mean SE Zooid length 15 0.232 0.401 0.334 0.051 Zooid width 15 0.245 0.352 0.289 0.033 Orifice length 8 0.108 0.133 0.123 0.008 Orifice width 8 0.103 0.138 0.114 0.011 Avicularia length 15 0.068 0.092 0.078 0.007 Avicularia width 15 0.036 0.058 0.046 0.006 Ovicell length 10 0.130 0.227 0.155 0.028 Ovicell width 10 0.150 0.237 0.182 0.023 Genus Turbicellepora Ryland, 1963 Turbicellepora iarae n. sp. (Figs. 78–80; Table 9) Material examined. Holotype: UFBA 1185, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Haliclona (Soestella) melana). Paratypes: UFBA 1667, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Haliclona (Soestella) melana); UFBA 2398, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Haliclona (Soestella) melana). Type locality. Camamu Bay, Bahia State, NE Brazil. Etymology. Honorific for Iara Coelho Sousa, Ana C.S. Almeida’s mother. Description. Colony encrusting, multilaminar, nodular. Autozooids semi-erect, hexagonal, limited by distinct grooves. Frontal shield heavily calcified, smooth to slightly rugose, convex, raised distally, imperforate centrally, marginally punctured by 5–20 large pseudopores. Primary orifice small relative to frontal shield, sunken, with arcuate distal edge and broad concave proximal sinus, with two triangular condyles placed at proximal third of the orifice. Secondary orifice oval to orbicular, often obscuring primary orifice and forming a smooth tubular peristome, particularly in ovicelled zooids. Avicularia absent. Ooecia globular, partially framed by the tubular peristome, punctured by 10–25 pseudopores, opening into distal part of peristome, just above the operculum. Remarks. Turbicellepora iarae n. sp. is the only species of the genus that lacks any type of avicularium. However, its colony form, frontal calcification, primary and secondary orifices, and distinct porose ooecia fully agrees with the definition of the genus. Other species reported from Brazilian waters are Turbicellepora brasiliensis Winston, Vieira & Woollacott, 2014, Turbicellepora pourtalesi Winston, 2005 and Turbicellepora winstonae Vieira, Gordon, Souza & Haddad, 2010. Besides the absence of avicularia, T. iarae n. sp. can be also distinguished from T. brasiliensis by the peristome (tubular in T. iarae n. sp.; broad and flat-rimmed in T. brasiliensis), and primary orifice (without sinus in T. iarae n. sp.; with shallower U-shaped sinus in T. brasiliensis). Other differences between T. iarae n. sp. and T. pourtalesi include the bluntly pointed frontal mucro, primary orifice with a median U-shaped sinus, sinuate peristome and avicularia of T. pourtalesi, all absent in T. iarae n. sp. Turbicellepora winstonae is distinct from T. iarae n. sp. in having an orificial sinus, oral avicularia and a short peristome (the peristome of T. iarae n. sp. is well-developed and tubular). 310 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. Turbicellepora species are frequently found on hard substrata such as calcareous nodules but also encrusting algae, worm tubes, bryozoans and hydrozoans (e.g., Hayward 1978; Winston 2005; Vieira et al. 2010). Colonies studied here were attached to the smooth-textured sponge Haliclona (Soestella) melana (Fig. 15). Distribution. Atlantic: Brazil (Bahia). FIGURES 78–83. 78–80. Turbicellepora iarae n. sp., UFBA 1185, holotype, Bahia State, Brazil. 78, Overview of encrusting colony; 79, Close-up of autozooids showing marginal pores, primary orifice with condyles, and tubular secondary orifice; 80, Close-up of zooids showing immersed porous ovicell. 81–83. Triphyllozoon arcuatum, UFBA 1184. 81, Overview of erect lace-like colony; 82, Detail of primary orifice and peristomial avicularia; 83, Detail of ovicelled zooid. Scale bars: 78 = 500 µm; 79–80, 81 = 200 µm; 82 = 50 µm; 83 = 100 µm. BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 311 TABLE 9. Measurements (mm) of Turbicellepora iarae n. sp. (UFBA 1185). Min: minimum; Max: maximum; N: number; SE: standard error. Structures N Min Max Mean SE Zooid length 15 0.582 0.944 0.737 0.013 Zooid width 15 0.459 0.645 0.551 0.064 Orifice length 15 0.196 0.272 0.228 0.017 Orifice width 15 0.161 0.224 0.191 0.017 Ovicell length 5 0.270 0.358 0.343 0.036 Ovicell width 5 0.283 0.405 0.355 0.044 Family Phidoloporidae Gabb & Horn, 1862 Genus Triphyllozoon Canu & Bassler, 1917 Triphyllozoon arcuatum (MacGillivray, 1889) (Figs. 81–83) Retepora monilifera form arcuata MacGillivray, 1889: 29. Triphyllozoon arcuatum: Almeida et al., 2015a: 2, figs. 2–17 (cum syn.). Material examined. UFBA 1612, UFBA 2399, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Tedania ignis). Remarks. Triphyllozoon arcuatum has erect robust, thickly calcified, lace-like colonies (Fig. 81), autozooids with low peristome and peristomial avicularia, frontal avicularia either small and round and/or large and oval (Fig. 82), large columnar avicularia, and a globose ovicell with sutures varying from arcuate to trifoliate with a short distal limb (Fig. 83) (Almeida et al. 2015a). Triphyllozoon arcuatum was recently recorded in the Western Atlantic for the first time, being recognized as a non-indigenous species to the area (Almeida et al. 2015a). Specimens of T. arcuatum here examined were found associated with the rugose-textured sponges Tedania ignis (Fig. 12). Triphyllozoon species are frequently found associated with other invertebrates, including the demosponges Tedania ignis and Dysidea etheria, and the calcareous sponge Clathrina sp. in Bahia State (for more information see Almeida et al. 2015a). Distribution. Indo-Pacific: South Australia, Northern Territory and Singapore. (Hayward 1999; Tilbrook & Gordon 2015). Atlantic: Brazil (Pernambuco and Bahia) (Almeida et al. 2015a). Genus Rhynchozoon Hincks, 1895 Rhynchozoon brasiliensis n. sp. (Figs. 84–89; Table 10) Rhynchozoon rostratum: Souza 1989: 502; Machado & Souza 1994: 259. Rhynchozoon verruculatum: Almeida et al. 2015b: 5 (in part). Material examined. Holotype: UFBA 1579, Todos os Santos Bay, 13°00’S, 38°32’W, 3–8 m, coll. 2013 (on sponge Callyspongia sp.). Paratypes: UFBA 1584, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Haliclona (Soestella) melana); UFBA 1189, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Bubaris sp.); UFBA 2340, Camamu Bay, 13°53’S, 38°59’W, 18–20 m, coll. October 2012 (on sponge Bubaris sp.). Type locality. Todos os Santos Bay, Bahia State, NE Brazil. Etymology. Alluding to the type locality in Brazil. 312 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. FIGURES 84–89. Rhynchozoon brasiliensis n. sp., UFBA 1579, holotype, Bahia State, Brazil. 84, Zooids at the growing edge with frontal and suboral avicularia; 85, Autozooids increasing calcification obscuring the primary orifice and suboral avicularium; 86, Group of older autozooids showing verrugate-like frontal process and frontal diamond-shaped avicularia; 87, Detail of primary orifice showing denticles and condyles; 88, Detail of diamond-shaped avicularium; 89, Ovicelled zooids. Scale bars: 84 = 250 µm; 85, 89 = 100 µm; 86 = 200 µm; 87, 88 = 50 µm. Description. Colony encrusting, uni- to multilaminar. Zooids at the growing edge polygonal, rectangular to hexagonal; primary orifices often with a single prominent proximo-lateral tubercle, and with a rhombic suboral avicularium directing distolaterally, with complete crossbar. In later astogeny the autozooidal frontal shield thickens and the peristome develops 4–6 tubercles, generally two distal and three proximal ones, obscuring the primary orifice and suboral avicularium. Autozooids almost indistinct, limited by slightly raised lateral walls. Frontal shield with tubercular processes, imperforate except by a single row of 10–16 large marginal pores. BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 313 Primary orifice small relative to zooidal length, the circular distal edge with 12–20 rounded denticles, the proximal edge with a deep and broadly V-shaped sinus; condyles small at proximal corners of the orifice. No oral spines. Peristome well-developed and frequently obscuring the primary orifice, formed by solid tubercles that become indistinct in later astogeny. Frontal avicularia sometimes absent in young zooids, but numerous and irregularly scattered throughout the colony, placed near zooidal margins; outline rhombic, small, with complete crossbar, rostrum elongate triangular. Ovicells prominent in young zooids, becoming immersed with increasing calcification; ooecia subglobular and frontally flat, endooecium completely calcified, ectooecium frontally uncalcified, with almost circular tabula, completely bordered by endooecium along the proximal margin; ooecia often covered by tubercular secondary calcification of the frontal shields of surrounding zooids along the lateral and distal margins. Remarks. Rhynchozoon brasiliensis n. sp. resembles Rhynchozoon fistulosum Hayward, 1993, Rhynchozoon incrassatum (Hincks, 1882) and Rhynchozoon papuliferum Souto, Kaufmann & Canning-Clode, 2015 in having a primary orifice with a distinct sinus, no oral spines and vicarious avicularia, frontal and zooidal avicularia with similar shape and size, and immsersed ovicells (in later astogeny) with exposed ooecial tabula. Rhynchozoon fistulosum is distinguished in having a shallow sinus (deep in R. brasiliensis n. sp.), a primary orifice that is wider than long (often longer than wide in R. brasiliensis n. sp.), smooth frontal calcification (with wart-like processes in R. brasiliensis n. sp.) and a single frontal avicularium (numerous in R. brasiliensis n. sp.). Rhynchozoon incrassatum is distinct in having a suboral avicularium placed on a well-developed calcified camara (absent in R. brasiliensis n. sp.) and a single frontal avicularium placed at the center of the frontal shield (numerous and marginal in R. brasiliensis n. sp.). Rhynchozoon papuliferum is distinguished from R. brasiliensis n. sp. in having many marginal pores (few in R. brasiliensis n. sp.), a primary orifice that is not obscured by the peristome (in R. brasiliensis n. sp. the peristome embeds the orifice), and the ectooecium is somewhat triangular in shape (almost circular in R. brasiliensis n. sp.). Although Souza (1989) and Machado & Souza (1994) recognized this species as Rhynchozoon rostratum (Busk, 1856), we analysed specimens studied by them (uncatalogued specimens deposited at UFBA) and conclude that they truly belong to R. bransiliensis n. sp. Differences between R. rostratum and R. brasiliensis n. sp. include the primary orifice (with a distinct rounded sinus in R. rostratum and almost V-shaped in R. brasiliensis n. sp.), frontal avicularia (distolaterally directed in R. rostratum and with no defined orientation in R. brasiliensis n. sp.), and a subtriangular to oval ooecial tabula (in R. brasiliensis n. sp. the tabula is almost circular). Almeida et al. (2015) also misidentified some specimens of R. brasiliensis n. sp. as Rhynchozoon verruculatum (Smitt, 1873), but R. verruculatum has a semicircular primary orifice with a shallow broad sinus (orifice circular and sinus almost Vshaped in R. brasiliensis n. sp.), and large diamond-shaped avicularia below and beside the peristome (absent in R. brasiliensis n. sp.). Since Souza (1989) and Machado & Souza (1994) were the only ones to report R. rostratum from Brazil, we conclude that this species does not occur in Brazilian waters, as already suggested by Vieira et al. (2010). TABLE 10. Measurements (mm) of Rhynchozoon brasiliensis n. sp. (UFBA 1579). Min: minimum; Max: maximum; N: number; SE: standard error. Structures N Min Max Mean SE Zooid length 15 0.280 0.475 0.351 0.052 Zooid width 15 0.250 0.404 0.303 0.049 Orifice length 8 0.095 0.118 0.103 0.007 Orifice width 8 0.079 0.113 0.098 0.012 Suboral avicularia length 10 0.056 0.113 0.093 0.014 Suboral avicularia width 10 0.037 0.062 0.050 0.007 Frontal avicularia length 15 0.081 0.111 0.094 0.008 Frontal avicularia width 15 0.042 0.057 0.048 0.004 Ovicell length 8 0.124 0.210 0.159 0.032 Ovicell width 8 0.175 0.262 0.214 0.033 314 · Zootaxa 4290 (2) © 2017 Magnolia Press ALMEIDA ET AL. Species of Rhynchoozoon are commonly found in coral reefs, on shells and other hard substrata (e.g., Marcus 1938, 1939; Osburn 1952; Winston 1986). Large colonies of R. brasiliensis n. sp. were found attached to the rugose-textured sponge Callyspongia sp. Distribution. Atlantic: Brazil (Bahia). Discussion The large number of bryozoan colonies (105 colonies) found associated with sponges in this study indicates that sponges may provide a good substratum for bryozoan settlement and survivorship in the study area. Environments with hard substrata such as rocky seafloor constitute in ideal habitats for larval settlement and colonial development for many bryozoans (Hughes 2001; Dick et al. 2005). Sponges can be considered one of the most common hard substrata available in both bays here studied. At Todos os Santos Bay, rocks are commonly found at the intertidal zone whereas in the subtidal zone (i.e. the studied area), sponges are frequently the dominant component of the seabed (Hadju et al. 2011). At Camamu Bay, sponges are often found in intertidal and subtidal area, near mangroves or with sandy bottom, being one common substrata for other invertebrates. At Todos os Santos and Camamu bays, a greater number of encrusting bryozoan species (67%; 14 species) were found on sponges when compared with erect species (33%; 7 species). This result follows the general faunal composition of bryozoans from Bahia State, which is represented by more than 80% by encrusting patches species (Almeida et al. 2015a,b,c; Almeida et al. 2017). At least 48 bryozoan species were already reported from Todos os Santos Bay and 36 from Camamu Bay (Almeida et al. 2015a,b,c; Almeida et al. 2017), from those most are encrusting species (40 species in Todos os Santos Bay and 30 species in Camamu Bay) that grows on several types of substrata, including rocks, shells, other bryozoans and artificial structures (Almeida et al. 2015a,b,c; Almeida et al. 2017). On the other hand, two studies on bryozoan/sponge-associations reported higher numbers of erect species than encrusting taxa (Klitgaard 1995; Padua et al. 2013). Klitgaard (1995) studied the fauna associated with 11 species of Demospongiae from the North Atlantic and found 31 bryozoan species, 20 of which (65%) were erect and 11 (35%) were encrusting. Since it is likely that there are more erect bryozoans living in greater depths and in environments lacking (or with less) hard substrata (e.g., Hayward 1981; Vieira et al. 2010), it could be possible that the colony growth forms found by Klitgaard (1995) are also dependent on the available bryozoan species in the surrounding environment. Thus, it may not be surprising that the most common bryozoan species found in her study was Bicellarina alderi (Busk, 1860), that has erect delicate branching colonies and is found on a wide range of substrata (Hayward & Ryland 1998: 242). Padua et al. (2013) studied the fauna associated with a single shallow water calcaerous sponge (Paraleucilla magna Klautau, Monteiro & Borojevic, 2004), reporting only three species widely recorded in Brazil– Bugula neritina (Linnaeus, 1758) (10 colonies), Scrupocellaria aff. reptans (Linnaeus, 1758) (20 colonies) and Hippoporina sp. (one colony). Thus, it is not possible to infer on the preference of bryozoans to colonize that sponge. Although we found there are distinctly more encrusting species growing on the sponges than erect ones, reflecting the pattern of the surrounding area, the absolute number of erect colonies found on the sponges at the studied bays is higher than the number of encrusting colonies (64 erect versus 41 encrusting colonies). This may reflect that encrusting patches species need a larger surface to colonial attachment/development than erect ones. Besides encrusting patches, encrusting bryozoans were also represented by small spot-like colonies (Celleporina joannae n. sp., Marcusadorea pinheroi n. sp. and Calyptooecia conuma), but they were only found at the recesses, where cryptic habitats for colonial attachment were available. Some encrusting patches colonies covered a large portion of the sponge surface, perhaps preventing the attachment of other colonies (e.g., Calyptotheca ornatissima n. comb. on the sponge Chondrilla nucula and Celleporaria carvalhoi on Topsentia sp.; Figs. 13 and 17, respectively), whereas most erect species cover partially the substrata using rhizoids (five species; 54 colonies). The fixation of the colony through rhizoids requires minute points of attachment, possibly allowing the establishment of a greater number of bryozoan colonies either at the lower, lateral and upper surfaces of the sponges. Most of erect species were attached to the sponge surface using these anchoring rhizoids (articulated and delicate branching species; five species, comprising 54 colonies) and few used rigid bases (the fenestrate Triphylozoon arcuatum with two colonies) and stolons (the stoloniferan Amathia distans with eight colonies). It is not unlikely that the most common bryozoan on the sponges was Licornia aff. diadema, with erect delicate BRYOZOANS INHABITING SPONGES IN NE BRAZIL Zootaxa 4290 (2) © 2017 Magnolia Press · 315 branching colonies, since it is a common and widespread species that occurs along the Brazilian coast on variable substrata (L.M. Vieira, unpublished data). Fenestrate colonies usually require broader space for the attachment of rigid bases and here were found attached to the lower portion of the sponges. Sometimes the colony grows embracing the substratum, becoming enmeshed with it (Fig. 12). Bryozoans preferentially colonized the undersurface of the sponges, which may be a measure to avoid being smothered by sediment. A similar condition is reported in other organisms that serve as substratum for bryozoan attachment, such as foraminiferal tests and brachiopods (Berning et al. 2009; Denisenko et al. 2013). As also known in other localities in Brazilian coast (e.g., Winston & Vieira 2013, who reported bryozoans on sand grains), bryozoans may be present on some unusual substrata in high energy localities with terrigenous/carbonate sediments, as those of Todos os Santos Bay and Camamu Bay. The availability of hard, protective substrata is perhaps the major factor acting upon the composition and distribution of the bryozoan fauna in environments that are usually regarded as inapt for bryozoan settlement, such as in areas with high irradiance and/or a high sedimentation rate (Hughes 2001; Berning et al. 2009; Figuerola et al. 2012; Wisshak et al. 2015). Moreover, as the sponge preferentially grows away from the substratum, settlement of bryozoan larvae on its lower side reduces the risk of being overgrown by the sponge. Despite absence of data on sponges without epizoic bryozoans, most bryozoan colonies studied here were attached to the surface of rugose-textured sponges, providing evidence that these animals are more likely to attach to irregular and rough surfaces. The preference of both adults and larvae of bryozoan to colonize rough surfaces was already demonstrated by Ward & Thorpe (1989), Marshall & Keough (2003) and Kuklinski & Barnes (2005) dealing with bryozoan association with several types of substrata, including rocks, algae, molluscs shells, hydrozoans and other bryozoans. It is possible that the nature of the substratum surface rather than the type of substratum determines the diversity and distribution of bryozoan species (Ward & Thorpe 1989; Kuklinski & Barnes 2005). The bryozoan fauna associated with sponges from Bahia State include genera and species often reported from NE Brazil (Vieira et al. 2008; Almeida et al. 2015b). The two new records of bryozoans from Todos os Santos Bay are represented by two new species here described—Microporella curta n. sp. and Rhynchozoon brasiliensis n. sp. —whereas the great number of new records from Camamu Bay can be attributed to the paucity of studies on bryozoan fauna from that area. Most of the species recorded here were already found on other substrata such as rocks, algae, other invertebrates and artificial structures. Thus, these species seem to be generalists (following Barnes & Clarke 1995) in terms of larval settlement requirements. A commensal relationship (inquilinism) between bryozoans and sponges is considered to be the case in the associations documented here, benefiting the bryozoan and with no apparent advantages or disadvantage to the sponges. Acknowledgements This study is part of A.C.S. Almeida’s PhD thesis supported by PROTAX-CNPq (440620/2015-5) through the Graduate Program in Animal Biology (Programa de Pós-Graduação em Biologia Animal) of the Departamento de Zoologia, UFPE. We are grateful to the Smithsonian’s National Museum of Natural History (USA) and JoAnn Sanner for sending SEM images of comparative species. Thanks also to Karoline Rebello, Jamile Farias, Anaíra Lage (Museu Nacional da Universidade Federal do Rio de Janeiro, UFRJ), Cristiana Castello-Branco and Julio Fernandez (Laboratório de Taxonomia de Porifera, UFRJ) for specimens collection and sponges procedures during their performance with LABPOR-UFBA team. Logistical support was provided for A.C.S. Almeida by the Centro de Pesquisa Gonçalo Moniz (FIOCRUZ/BA), Orane Alves (Laboratório de Geoecologia de Sedimentos Marinhos, UFBA) and Ulisses Pinheiro (Laboratório de Porifera, UFPE). Expeditions to Camamu Bay (2012) and Todos os Santos Bay (2012, 2013) were funded by FAPESB, CNPq and PETROBRAS to C. 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