This article was downloaded by: [Ramalho, Lais] On: 24 March 2011 Access details: Access Details: [subscription number 935356722] Publisher Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 3741 Mortimer Street, London W1T 3JH, UK Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713192031 Taxonomic revision of some lepraliomorph cheilostome bryozoans (Bryozoa: Lepraliomorpha) from Rio de Janeiro State, Brazil Laís V. Ramalhoa; Guilherme Muricya; Paul D. Taylorb Departamento de Invertebrados, Laboratório Biologia de Porifera. Quinta da Boa Vista s/n, São Cristóvão, Museu Nacional, Rio de Janeiro, Brazil b Department of Palaeontology, Natural History Museum, London, UK a Online publication date: 24 March 2011 To cite this Article Ramalho, Laís V. , Muricy, Guilherme and Taylor, Paul D.(2011) 'Taxonomic revision of some lepraliomorph cheilostome bryozoans (Bryozoa: Lepraliomorpha) from Rio de Janeiro State, Brazil', Journal of Natural History, 45: 13, 767 — 798 To link to this Article: DOI: 10.1080/00222933.2010.535917 URL: http://dx.doi.org/10.1080/00222933.2010.535917 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf This article may be used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Journal of Natural History Vol. 45, Nos. 13–14, April 2011, 767–798 Taxonomic revision of some lepraliomorph cheilostome bryozoans (Bryozoa: Lepraliomorpha) from Rio de Janeiro State, Brazil Laís V. Ramalhoa* , Guilherme Muricya and Paul D. Taylorb a Museu Nacional, Departamento de Invertebrados, Laboratório Biologia de Porifera. Quinta da Boa Vista s/n, São Cristóvão, Rio de Janeiro, Brazil; b Department of Palaeontology, Natural History Museum, London, UK Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 (Received April 2010; final version received 26 October 2010; printed 22 March 2011) Lepraliomorph-grade ascophorans are the dominant group of bryozoans in most modern faunas. Here we describe five new species (Parasmittina alba sp. nov., Microporella proxima sp. nov., Reteporella antennata sp. nov., Stephanollona robustaspinosa sp. nov. and Turritigera buski sp. nov.) and record an additional five lepraliomorph species [Watersipora subovoidea (d’Orbigny, 1852), Schizoporella errata (Waters, 1878), Arthropoma cecilii (Audouin, 1826), Celleporina diota (Marcus, 1938) and Reteporellina evelinae Marcus, 1955] from the coastal waters off Rio de Janeiro State in Brazil. This increases the biodiversity of the bryozoan fauna from Brazil, and the number of species known from Rio de Janeiro State now totals 79. Keywords: Bryozoa; Lepraliomorpha; taxonomy; Brazil; South Atlantic Introduction Species of bryozoans can be found in great abundance and high diversities in shallow waters all around the globe (e.g. Ryland 1970; Gordon 1984). The Brazilian coastline is 8000 km long but little research has been undertaken on the bryozoans. By 1980, 346 bryozoan species had been recorded from Brazil but records of marine bryozoans existed for only seven of the 17 states bordering the Atlantic Ocean: Pernambuco, Sergipe, Bahia, Espírito Santo, Rio de Janeiro, São Paulo and Paraná states (d’Orbigny 1841–47; Kirkpatrick 1890; Canu and Bassler 1928; Marcus 1937, 1938, 1939, 1941, 1955; Buge 1979). Taxonomic studies of Brazilian bryozoans have recently recommenced. Winston and Migotto (2005) reported 13 species (nine cheilostomes, three ctenostomes and one cyclostome) encrusting or boring into sand grains from the São Paulo coast. Gordon et al. (2006) identified Membraniporopsis tubigera (Osburn, 1940) washed up in great abundance on the shores of some Brazilian states. Vieira et al. (2007) described Catenicellidae from Alagoas State, introducing one new species, and Vieira et al. (2010) revised the shallow-water species of Beania from the Brazilian coast. Ramalho et al. (2005, 2008a,b, 2009) studied parts of the bryozoan fauna from Rio de Janeiro State, describing 18 species, six of which were new, giving a total of 69 species known from this state. Finally, Santana et al. (2009) described two species of Metrarabdotos, one new, from Sergipe, Bahia and Espírito Santo states. A large number of bryozoan families are grouped in the “grade” Lepraliomorpha. They are characterized by a frontal shield, which is cryptocystidean and has an *Corresponding author. Email: laiscanabarro@yahoo.com.br ISSN 0022-2933 print/ISSN 1464-5262 online © 2011 Taylor & Francis DOI: 10.1080/00222933.2010.535917 http://www.informaworld.com 768 L.V. Ramalho et al. outer hypostegal coelom and marginal areolae. The frontal shield may be centrally imperforate or penetrated by pseudopores. Frontal budding, when it occurs, is from interzooidal pore chambers and the hypostegal coelom and areolae (Gordon 1989). The aim of this study is to describe 10 species of lepraliomorphs, five new, collected from Rio de Janeiro State. Most of the established species have not been adequately described previously from Brazil, and few have been illustrated using scanning electron microscopy (SEM), a technique now essential in bryozoan taxonomy. Materials and methods Study area The samples were collected from six sites in Rio de Janeiro State (Figure 1): Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 • Macaé: material was collected from ships’ hulls and oil platforms at Bacia de Campos by SCUBA divers from the Instituto de Estudos do Mar Almirante Paulo Moreira (IEAPM). • Arraial do Cabo: samples were collected between February 2002 and February 2004 by SCUBA divers, from artificial and natural substrata at 0.5–10 m depth in three different areas (Forno Harbour, Pedra Vermelha and Farol Beach). Figure 1. Map of the study area. 1. Macaé; 2. Arraial do Cabo; 3. Guanabara Bay; 4. Sepetiba Harbour; 5. REVIZEE Station 2 (23◦ 47.629′ S, 041◦ 42.514′ W); 6. REVIZEE Station 9 (24◦ 16.353′ S, 043◦ 23.788′ W). Journal of Natural History 769 • Guanabara Bay: samples were collected on artificial substrata (ships’ hulls) dur- Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 ing September 1999 and October 2002. In this bay there are several harbours with large daily influxes of ships on national and international routes. The bay is considered to be a eutrophic area because of industrial and domestic sewage. • Sepetiba Harbour: samples were collected by IEAPM SCUBA divers between November and December 2001 from artificial substrata in several terminals. • REVIZEE Program Campaign Station 2 (23◦ 47.629′ S, 041◦ 42.514′ W) on calcareous substrata collected using a trap from 341 m depth. • REVIZEE Program Campaign Station 9 (24◦ 16.353′ S, 043◦ 23.788′ W) on calcareous substrata collected using a trap from 600 m depth. Both REVIZEE stations are located off the continental margin, near the top of the slope. The area of Station 9 is considered “subtropical” and the water temperature is low. The seabed in this area is composed of fine sand and mud. However, calcareous substrata formed by polychaete tubes and other animals are frequent. Samples were fixed in 70% ethanol. Digital images were mostly obtained of uncoated, dried specimens using a low-vacuum scanning electron microscope (LEO 1455VP) at the Natural History Museum, London, UK. All specimens were deposited in the Bryozoan Collection of the Museu Nacional, Rio de Janeiro, Brazil. Systematics Order CHEILOSTOMATA Busk, 1852 Infraorder ASCOPHORA Levinsen, 1909 ‘Grade’ LEPRALIOMORPHA Gordon, 1989 Superfamily SMITTINOIDEA Levinsen, 1909 Family SMITTINIDAE Levinsen, 1909 Genus Parasmittina Osburn, 1952 Parasmittina alba sp. nov. (Figure 2) Material examined Arraial do Cabo: Holotype: MNRJ-077, 24 May 2002, Pedra Vermelha, Cabo Frio Island, collected by L.V. Ramalho, depth 5 m. Paratype: MNRJ-079, 27 February 2003, Pedra Vermelha, Cabo Frio Island, collected by L.V. Ramalho and R. Melo, depth 5 m. Parasmittina raigioidea: holotype: 95BFS0820-6 1995 (18◦ 11′ N, 109◦ 21′ E), depth 4 m. Etymology The name alba refers to the white colour of the colonies. Diagnosis Colony encrusting, unilamellar, frontal wall granular with marginal areolar pores; primary orifice rounded with prominent condyles turned downwards; lyrula well Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 770 L.V. Ramalho et al. Figure 2. Parasmittina alba sp. nov. (A) Group of zooids without frontal budding; (B) old part of colony; (C) zooids arrangement and different types of avicularia; (D) autozooids with orifices and lyrulae; (E) detail of two autozooids showing small triangular avicularia and distal spines; (F) autozooids showing small avicularia with rounded distal tips; (G) giant avicularium with elongate spatulate rostrum and small avicularia with round and smooth rostrum; (H) ovicellate autozooid. Scale bars: A, C, E–H: 100 µm; B: 400 µm; D: 200 µm; A: MNRJ-077 and others: MNRJ-079. Journal of Natural History 771 Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 developed; peristome may be elevated proximally, forming two projections; secondary orifice with pseudosinus; oral spines numbering one or two; avicularia comprising one interzooidal type and up to three adventitious types (large with rostrum elongated and spatulate; small with rostrum rounded and smooth; small with rostrum slightly triangular and serrated), only one type occurring on each zooid. Ovicell with ectooecium perforated by pores or fenestra. Description Colony encrusting, variable in shape and size, unilamellar, white (Figure 2A,B). Autozooids rectangular, longer than wide (451 µm long, range 392–490 µm, by 305 µm wide, range 265–383 µm), disposed in linear series on young colonies (Figure 2A), linear pattern obscured on older colonies or colonies growing on irregular substrata (Figure 2B,C); the two daughter zooids after a bifurcation are relatively small and narrow (Figure 2A). Frontal wall granular without pseudopores but having marginal areolar pores (Figure 2C,D). Primary orifice rounded with prominent condyles turned downwards and lyrula. Secondary orifice with pseudosinus, well developed (Figure 2D); peristome slightly elevated laterally and sometimes strongly elevated proximally, hiding the lyrula, sometimes forming two proximal projections (Figure 2C–H). Oral spines numbering two, thin, non-articulated, often broken, leaving only a scar; rarely only one medial oral spine is present (Figure 2D–F). Ovicell prominent, large, wider than long (274 µm long by 323 µm wide). Ectooecium membranous frontally, revealing the perforate, calcified entooecium (Figure 2B,H). Avicularia adventitious and interzooidal. Adventitious avicularia of three types but only one per autozooid: (1) large avicularia generally placed next to orifice, but sometimes randomly, turned proximally and slightly oblique, rostrum elongated (353 µm long by 137 µm wide), spatulate, smooth and more dilated at the tip (166 µm wide), covering almost all the frontal wall of the autozooid (Figure 2C,G); (2) small avicularia adjacent to the orifice or placed more lateromedially, oriented proximally and slightly towards the lateral zooidal margin, with mandible and rostrum rounded, smooth and more dilated at the proximal end (Figure 2C,F–H); (3) small avicularia (109 µm long, range 98–118 µm, by 83 µm wide, range 69–98 µm) with rostrum slightly triangular having small teeth on the border, placed almost randomly, tending to be oriented proximally and slightly towards the lateral zooidal margin (Figure 2C,E). Interzooidal avicularia large, with triangular rostrum, border asymmetrical, the margin of one side curved; cross-bar complete (Figure 2C). Habitat Colonies found growing directly on rocks, 5 m deep. Geographical distribution Colonies collected only at Arraial do Cabo (Pedra Vermelha). 772 L.V. Ramalho et al. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Remarks Among the Brazilian species of Parasmittina, P. evelinae (Marcus, 1937) has five oral spines, large interzooidal avicularia are absent, and the lyrula is wide with acute or bicuspate tips. Barbosa (1964) noted five “varieties” of a species she attributed to the European P. trispinosa: P. trispinosa vars. ligulata, loxa, munita, nitida and spathulata. All of these bear three oral spines, unlike P. alba sp. nov. Several other species of Parasmittina worldwide share some characteristics with P. alba sp. nov. Although P. betamorphaea Winston, 2005 occurs in Brazil, and was previously mentioned as Smittina trispinosa var. nitida by Marcus (1937: 104), it may be distinguished from P. alba sp. nov. in having more than one avicularium per autozooid, lacking a serrated rostrum in the small avicularia, possessing giant avicularia with more rounded rostra, and by the different disposition and shape of the pores on the ovicells. Another species that is close to P. alba sp. nov. is P. raigioidea Liu et al., 2001. However, P. raigioidea lacks avicularia with serrated rostra and the small avicularia have mandibles directed distally. While P. winstonae Liu et al., 2001 has several different types of avicularia, like P. alba, there may be more than one avicularium on each autozooid and none of them have serrated rostra. The oral lateral avicularia are large, directed obliquely distally and have rostra with less dilated tips. The triangular avicularia are small, placed on one side of the peristome or elsewhere on the frontal wall, with oblique and distally directed rostra. Ovicells in P. winstonae have small pores and P. parsevalioidea Liu et al., 2001 has two oral spines, avicularia without serrated rostra and the peristome is laterally elevated. In P. galerita Ryland and Hayward, 1992 there are one or two oral spines and the avicularia vary in shape. However, condyles are rounded and thinly serrated on their free parts, more than one avicularium may be borne by each autozooid, some of them with mandibles obliquely directed distally, and the ovicell has small pores. Family WATERSIPORIDAE Vigneaux, 1949 Genus Watersipora Neviani, 1895 Watersipora subovoidea (d’Orbigny, 1852) (Figure 3) Cellepora subovoidea d’Orbigny 1852: 402. Lepralia cucullata Busk 1854: 81, pl. 96, figs 4–6. Watersipora subovoidea, Ryland et al. 2009: 54, fig. 4C,D,G,H. Material examined Macaé: MNRJ-141, ship hull Valentin Shashin, 14 December 2002, collected by J.E.A. Gonçalves, depth 1.5 m. Arraial do Cabo: MNRJ-012, Forno Harbour, 11 October 2002, collected by L.V. Ramalho and D.C. Savi, depth 0.5 m. MNRJ-138, Forno Harbour, 20 August 2004, collected by L.V. Ramalho and D.C. Savi, depth 0.5 m. MNRJ-139, Forno Harbour, 7 July 2003, collected by L.V. Ramalho, depth 1.5 m. MNRJ-142, Farol beach – Cabo Frio Island, 4 April 2002, collected by L.V. Ramalho, depth 5 m. Rio de Janeiro City: MNRJ-140, Guanabara Bay (ship hull), 23 October 2002, collected by G. Muricy, depth 0.5 m. Sepetiba Harbour: Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Journal of Natural History 773 Figure 3. Watersipora subovoidea (d’Orbigny, 1852). (A) General view of the colony; (B) autozooidal arrangement; (C) detail of autozooids (frontal view) with large pseudopores, slightly developed peristome and orifice with condyles and wide sinus. Scale bars: B: 400 µm; C: 100 µm. MNRJ-012; (D) Detail of operculum. MNRJ-666, Ilha Guaíba, June 2009, collected by J. Silva. Watersipora cucullata: NHM 1973.1.10, Gold Coast, Ghana; NHM 1948.2.16.18, Santos (SP), Brazil, collected by E. Marcus. W. subtorquata: NHM 1980.1.12.3, Townsville Harbour, Queensland, Australia. W. subovoidea: NHM 1986.8.14.3, Florida, USA. NHM 1998.8.4.13, Port Vila Waterfront, Vanuatu. Diagnosis (revised) Colony encrusting, sometimes erect, unilamellar, black or brown, operculum having a median vertical stripe between two pale lateral areas (grey or white); zooids almost rectangular, disposed in quincunx, frontal wall covered by large pseudopores; no ovicells or avicularia. Description Colonies initially circular, from 0.5 to 1.4 cm in diameter, sometimes growing free from substrate forming unilamellar fronds. Central part of the colony dark, varying from brown to black, but colony border is sometimes pale in colour (grey, beige or orange) (Figure 3A). 774 L.V. Ramalho et al. Autozooids disposed in regular quincunx, oblong, almost rectangular, larger than wide (789 µm long, range 456–1038 µm by 390 µm wide, range 262–582 µm), divided from each other by a dark suture (Figure 3B). Frontal wall with scattered large pseudopores (29 µm, range 19–39 µm diameter), decreasing in abundance near the orifice (Figure 3C). Some zooids are wider and bud two narrower zooids to form new bifurcations. Primary orifice rounded (221 µm long, range 155–252 µm, by 234 µm wide, range 194–272 µm), with a sinus wider than deep (52 µm long, range 39– 68 µm, by 116 µm wide, range 58–136 µm), delimited by two prominent condyles (Figure 3C). Peristome variably developed around the primary orifice, often slightly elevated (Figure 3B,C). Operculum very dark (same colour as rest of autozooid), distal half with a vertical, median parallel-sided stripe showing two different colours: a black band and paler peripheral regions (Figure 3D). Avicularia, spines and ovicells absent. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Habitat Colonies collected on harbour piers, ships’ hulls, hydrozoans, bryozoans (Zoobotryon verticillatum) and Sargassum algae; 0–5 m deep. Geographical distribution World: Atlantic and Pacific Oceans (Ryland et al. 2009). Brazil: Study area: Macaé, Arraial do Cabo (Forno Harbour and Farol Beach), Rio de Janeiro City (Guanabara Bay) and Sepetiba Harbour. Remarks The first record of the well-known fouling bryozoans Watersipora from the Rio de Janeiro coast was of W. subtorquata, described by d’Orbigny (1841–47) under the name Escharina torquata. Marcus (1937, 1938) subsequently described specimens from São Paulo and Espírito Santo states as W. cucullata. Winston (1982) included Marcus’s specimens in the synonymy of W. subovoidea, whereas Gordon (1989) included these same specimens in the synonymy of W. subtorquata and suggested that the name W. subovoidea should be discarded because this species cannot be recognized and the type locality is unknown. Taylor and Gordon (2002) figured d’Orbigny’s material from Rio de Janeiro of W. subtorquata, represented by specimen no. 13637 at the MNHN, Paris. Recently, Ryland et al. (2009) reviewed W. subovoidea and W. subtorquata, showing two principal characters that distinguish these two species. Watersipora subovoidea has triangular, tooth-like condyles, located distomedially, and a strongly pigmented operculum with a parallel-sided dark central band; W. subtorquata has condyles in the form of narrow shoulders and an operculum with a dark biconcave band proximally. Ryland et al. (2009) added W. cucullata described by Marcus (1937, 1938) to the synonym of W. subtorquata, agreeing with Gordon (1989). Comparing Rio de Janeiro specimens with the figures shown in Ryland et al. (2009), it is evident that these specimens are very similar to W. subovoidea in having Journal of Natural History 775 triangular condyles and a parallel-sided dark band on the operculum. Therefore, we believe that it is important to review these characters in the species described by Marcus (1937, 1938). Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Superfamily SCHIZOPORELLOIDEA Jullien, 1883 Family SCHIZOPORELLIDAE Jullien, 1883 Genus Schizoporella Hincks, 1877 Schizoporella errata (Waters, 1878) (Figure 4) Figure 4. Schizoporella errata (Waters, 1878). (A) General view of the colony; (B) part of the colony; (C) autozooidal arrangement; (D) detail of the sinus; (E) avicularia; (F) drawing of the ancestrula and the first zooids (Courtesy of Hayward and Ryland 1979). Scale bars: A: 2 cm; B: 600 µm; C: 400 µm; D–F: 100 µm. A–E: MNRJ-010. 776 L.V. Ramalho et al. Lepralia errata Waters 1878: 11, pl. 1, fig. 9. Schizoporella errata, Gautier 1962: 149, fig. 151; Ryland 1965: 64, fig. 31A,B; Hayward and Ryland 1999: 212; Tilbrook et al. 2001: 80, fig. 15B; Tompsett et al. 2009: 2234, figs. 3A–F, 4A–F. Schizoporella violacea Canu and Bassler 1930: 40, pl. 4, figs 1–14. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Material examined Macaé: ship hull Valentin Shashin: MNRJ-212, 1 May 2002, coll. J.E.A. Gonçalves, depth 2 m. MNRJ-213, 14 December 2002, collected by L.V. Ramalho, depth 2 m. Arraial do Cabo city: Forno Harbour: MNRJ-010, 11 October 2002, coll. by L.V. Ramalho, depth 0.5 m; MNRJ-218, MNRJ-219, 31 July 2003, collected by L.V. Ramalho and D.C. Savi, depths 3 and 0.5 m, respectively. MNRJ-215, Pedra Vermelha, 27 February 2003, collected by L.V. Ramalho, depth 5 m. MNRJ-220, Farol beach, 22 January 2003, coll. L.V. Ramalho, depth 0.5 m. Guanabara Bay: MNRJ-211, ship hull Leo Segerius, 15 September 1999, coll. J. Quintanilha, depth 2 m. Sepetiba Harbor: Guaíba Pier: MNRJ-216, MNRJ-206, MNRJ-208, 14 November 2001, collected by IEAPM, depths 0.5, 3 and 7 m, respectively. Tebig Oil Terminal 2: MNRJ-207, MNRJ-209, 12 November 2001, coll. IEAPM, depth 3 and 7 m, respectively. MNRJ-210, Iron Ore Terminal, 10 November 2001, collected by IEAPM, depth 7 m, MNRJ-217, Container Terminal 2, 9 November 2001, collected by IEAPM, depth 7 m. Schizoporella errata. NHM 26.9.6.226, Suez Channel. NHM 1969.11.4.29, Charkor, Gold Coast (on wood and net), collected by Bassindale, depth 14 m. NHM 1969.11.4.28, Gorée Harbour, Senegal, collected by M. Marchad. Description Colony encrusting, uni- or multilamellar, sometimes developing erect tubes up to 10 cm high (Figure 4A); older parts brown, orange at the growing edges, sometimes a shade paler when alive; when fixed in alcohol orange colour lost, becoming brown and very dark, almost black, (Figure 4A,B). Size variable, initial growth circular, becoming irregular, up to 30 cm high; forming extensive growths on ship hulls in which boundaries of individual colonies are difficult to define. Autozooids rectangular to hexagonal, longer than wide (598 µm long, range 456–776 µm, by 395 µm wide, range 272–534 µm), regularly disposed in basal layer of young colonies, irregularly disposed in older colonies with extensive frontal budding (Figure 4C). Autozooids separated by furrows, more visible in young colonies. Frontal wall evenly porous (Figure 4C–F), a small umbo sometimes present below orifice. Orifice circular (163 µm long, range 136–184 µm, by 156 µm wide, range 116–175 µm), with U-shaped sinus, wider than deep (40 µm long, range 29–58 µm by 68 µm wide, range 49–107 µm), and two small condyles on each side (Figure 4E). Avicularia present or absent, only one per autozooid, triangular with acute mandible, located proximolaterally and close to orifice, directed distolaterally (Figure 4F). Ovicell globose, surface wrinkled, with some pores. Journal of Natural History 777 Habitat Colonies directly encrusting harbour piers, nylon ropes or other organisms (ascidians and cirripeds), 0–6 m depth. Geographical distribution This is generally considered to be a warm temperate and subtropical species recorded from the eastern and western Atlantic, western Pacific, Mediterranean and Red Sea (Hayward and Ryland 1999; Tilbrook et al. 2001; Tompsett et al. 2009). In Brazil S. errata has been previously recorded in an ecological study (São Paulo State) (Morgado and Tanaka 2001). In the study area it has been found at Macaé, Arraial do Cabo, Guanabara Bay and Sepetiba Harbour. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Remarks Schizoporella pungens (Canu and Bassler, 1928) resembles S. errata, their measurements overlapping and colony shape and colour being similar. However, Winston (2004, 2005) mentioned distinct differences, with S. pungens having a wider and shorter orifice, narrower sinus, smaller and blunter avicularia, and an ancestrula with an ovoidal membranous area. Despite the great confusion in the identification of Schizoporella species worldwide (Hayward and Ryland 1995; Soule et al. 1995; Winston 2005; Tompsett et al. 2009), the Brazilian material agrees well with S. errata because of characteristics of the orifice, including the sinus, the avicularia and the presence of large pseudopores in shallow depressions on the frontal wall. Previous taxonomic works (d’Orbigny 1841–47; Kirkpatrick 1890; Marcus 1937; Hayward and Ryland 1979) have mentioned the occurrence of several Schizoporella species from the Brazilian coast: S. trimorpha Canu and Bassler, 1928, S. carvalhoi Marcus, 1937, S. horsti (Osburn, 1927), S. pungens (Canu and Bassler, 1928), S. isabelleana (d’Orbigny, 1839), S. errata (Waters, 1878) and S. unicornis (Johnston, 1847). Schizoporella unicornis has been recorded several times from Brazil, but almost certainly in error. Some authors have placed this species in synonymy with other species of Schizoporella because it occurs with S. pungens, mentioned by Marcus (1937) from São Paulo and Bahia. A detailed review of the Schizoporella species recorded from Brazil must be done to confirm the correct identities of the species which occur along the Brazilian coast. Family LACERNIDAE Jullien, 1888 Genus Arthropoma Levinsen, 1909 Arthropoma cecilii (Audouin, 1826) (Figure 5) Flustra cecilii Audouin 1826: 239, pl. 8, fig. 3. Arthropoma cecilii, Canu and Bassler 1930: 316; Marcus 1937: 93, fig. 49; 1955: 297; Harmer 1957: 1001, pl. 72, fig. 23, text-fig. 108; Gautier 1962: 127; Ryland and Hayward 1992: 263, fig. 19C, D. ?Arthropoma cecilii, Canu and Bassler 1929: 296, pl. 32, fig. 1, text-fig. 126; Osburn 1952: 333, pl. 3, figs 1–3; Liu et al. 2001: 607, pl. 51, fig. 4. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 778 L.V. Ramalho et al. Figure 5. Arthropoma cecilii (Audouin, 1826). (A) Part of the colony; (B) detail of the zooids; (C) detail of an elongate autozooid; (D) fertile zooids. Scale bars: A: 600 µm; B–D: 200 µm. MNRJ-069. Material examined Arraial do Cabo, Rio de Janeiro State: MNRJ-069, 24 May 2002, Pedra Vermelha (Cabo Frio Island), collected by L.V. Ramalho, depth 5 m. Arthropoma cecilii. NHM 1948.2.16.57, Santos, Brazil, collected by E. Marcus; NHM 1975.7.1.34, Chios, Expedition to Chios, 1967; NHM 11.10.1.1257, British Barleei; NHM 2000.4.11.1042, 14 October 1978, Swa Barrier Reef, Fiji, collected by J.S. Ryland; NHM 1931.12.30.99, Jolo, Philippines, Bassler Collection, 38 m. Revised diagnosis Colony encrusting, unilamellar; autozooids hexagonal to rectangular with frontal wall perforated by circular pseudopores except centrally; orifice with proximal border almost straight; sinus moderate to deep; ovicell globose, imperforate. Description Colony encrusting, unilamellar (Figure 5A), varying in size and shape, white with orange embryos. Autozooids hexagonal to rectangular, longer than wide (746 µm long, range 657–951 µm, by 437 µm wide, range 333–490 µm), disposed in quincunx or irregularly (Figure 5A,B); occasionally very irregular shaped zooids. Frontal wall with large circular pseudopores, except in a central area beneath the orifice, which is Journal of Natural History 779 smooth and imperforate. Umbo often present immediately below the orifice. Orifice D-shaped, wider than high (Figure 5C), with a straight proximal border and narrow; U-shaped sinus, moderate to deep (168 µm long, range 157–196 µm by 213 µm wide, 196–245 µm), constricted and almost closed distally (Figure 5B–D). Ovicell globose, higher than wide (437 µm long, range 392–470 µm long by 384 µm wide, range 323–412 µm, without pores, surface rough, seemingly cryptocystal; ectooecium reduced to a smooth, narrow band bordering orifice; closed by zooidal operculum. No oral spines observed. Avicularia absent. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Habitat Colonies were found on rock at 5 m depth. Geographical distribution This species is widely distributed in temperate-circumtropical regions of the southeast Atlantic (Brazil), Western Atlantic, Mediterranean Sea, Indian and Pacific Oceans (Canu and Bassler 1929, 1930; Marcus 1937; Osburn 1952; Ryland and Hayward 1992). In Brazil it has been recorded from São Paulo (Santos) and Espírito Santo (south of Vitória) (Marcus 1937, 1955), and now Arraial do Cabo (Pedra Vermelha, Cabo Frio Island) in Rio de Janeiro State. Remarks The Brazilian material is very similar to specimens collected in the North Atlantic (NHM 1975.7.1.34, Chios; and NHM 11.10.1.1257, British Barleei) and the Pacific (NHM 2000.4.11.1042, Fiji), sharing with them a sinus of the same shape, not very deep and narrowing distally, as well as lacking pseudopores on the median region below the sinus. However, material from Chios (NHM 1975.7.1.34) has two oral spines. A specimen from Santos (NHM 1948.2.16.57) is also very similar to those described here. It shows variability in sinus depth, which is sometimes deep and narrow and sometimes shallow and wide, but always the sinus narrows at its opening. The specimens described by Osburn (1952) and Canu and Bassler (1929) are distinguished from Brazilian species because they have a sinus that is deep and straight, without the extension of the orificial proximal border, pseudopores over the entire frontal surface, and the umbo may be absent. The specimen described by Liu et al. (2001) also has pseudopores over the entire frontal wall, the sinus is shallower than in Osburn’s material, and the extension of the orificial proximal border does not occur. Comparing the measurements of Osburn’s material to the Brazilian specimens, substantial differences are observed in zooid size, which is smaller and narrower in Osburn’s material; the operculum is also smaller. Similar differences are observed in the Chinese specimens described by Liu et al. (2001). These specimens are in need of a detailed review because of several differences with A. cecilii sensu Audouin (1826). 780 L.V. Ramalho et al. Marcus (1937) and Harmer (1957) regarded A. perugiana as a synonym of A. cecilii. However, SEM study of A. perugiana shows differences between these two species in the size and shape of the sinus, which is narrower, deeper and straight in A. perugiana, the proximal border of the orifice not extending over the sinus entrance. These differences suggest that A. cecilii and A. perugiana are separate species. New studies should be carried out on material previously identified as A. cecilii to correct the identification and geographical distributions of these two species. The Brazilian material studied here is considered to be A. cecilii. This is the first record of A. cecilii from the Rio de Janeiro coast. Its distribution in Rio de Janeiro State seems to be limited to a small area in Arraial do Cabo on the basis of sampling undertaken to date. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Family MICROPORELLIDAE Hincks, 1880 Genus Microporella Hincks, 1877 Microporella proxima sp. nov. (Figure 6) Examined material Holotype: MNRJ-180, Macaé, Rio de Janeiro State, Brazil, collected by J.E.A. Gonçalves, 14 December 2002, on Sargassum from the hull of the Valentin Shashin, depth 3 m. Figure 6. Microporella proxima sp. nov. (A) General view of the colony on an algal frond; (B) zooidal arrangement; (C) autozooids, avicularia and ovicell; (D) detail of autozooid showing oral spines, avicularia with setiform mandible and ascopore. Scale bars: A: 400 µm; B: 300 µm; C,D: 100 µm. A–D: MNRJ-180 Journal of Natural History 781 Etymology The name proxima refers to the position of avicularium, near to the zooidal margin and more proximal than most other species of Microporella. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Diagnosis Microporella with encrusting colony; autozooids small, averaging 416 µm long by 322 µm wide; frontal wall pustolose, evenly perforated by pseudopores and with narrow, inconspicuous areolar pores around the edges; orifice D-shaped, slightly flattened, the proximal edge beaded, four oral spines, one pair distally and the other pair mediolaterally; ascopore reniform with short, tooth-like spines, situated on a raised area of frontal shield; avicularium single, located near the margin, about mid-length on the autozooid, distant from ascopore, oriented laterally to distolaterally, mandible elongated, setose, irregular. Description Colony encrusting, unilamellar (Figure 6A), up to 1 cm in diameter. Early astogeny unknown. Autozooids disposed in quincunx, rounded rhombic in frontal outline, separated by deep grooves, small, higher than wide (416 µm long, range 392–441 µm, by 322 µm wide, range 294–343 µm). Frontal wall convex with granulations and rounded pseudopores scattered across entire surface except for area between ascopore and orifice (Figure 6B); areolar pores inconspicuous, more elongate than pseudopores. Orifice D-shaped (70 µm long, range 64–79 µm, by 105 µm wide, range 93–107 µm), slightly flattened, proximal border straight, beaded; oral spines delicate (Figure 6C), four in number, one pair distal and the other pair mediolateral, not close to the proximal corners of orifice; only two spines generally visible in ovicellate zooids. Ascopore reniform, located on a raised area of frontal shield, margin with minute teeth (Figure 6D). Ovicell prominent, closed by zooidal operculum, almost round, slightly higher than wide; surface similar to an autozooidal frontal shield, porous and granular (Figure 6D). Avicularium single, located laterally roughly midway along autozooid, rarely close to the orifice, directed laterally or distolaterally; mandible elongate, (209 µm long, range 196–216 µm), pointed, thin and irregular (not straight), turned distolaterally (Figure 6C,D). Habitat Colonies found on algae (Sargassum) growing on the hull of the Valentin Shashin. Geographical distribution Macaé. Colonies were collected from the hull of the drilling ship Valentin Shashin but as this has remained in this area for a least 5 years, it seems likely that the new species is established here and did not colonize when the ship was elsewhere. 782 L.V. Ramalho et al. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Remarks In the past, this species would probably have been determined as Microporella ciliata (Pallas, 1766). However, many identifications of M. ciliata are now considered incorrect, having been made before the concept of the species was stabilized by Kuklinski and Taylor (2008) through the selection of a neotype, NHM 2008.02.06.1, from the Bay of Naples. Comparing Microporella proxima sp. nov. with the similar M. ciliata, the following differences are evident: (1) M. ciliata has smaller pseudopores; (2) autozooids are larger in M. ciliata than in M. proxima, average lengths being 790 and 420 µm respectively; (3) the avicularium is located more proximally and further away from the ascopore in M. proxima; and (4) the most proximal pair of oral spines is just above the proximolateral corners of the orifice in M. ciliata but more distally located in M. proxima. Marcus (1937, 1955) described putative M. ciliata from São Paulo and Espírito Santo states but his material represents neither M. ciliata nor M. proxima and is in need of revision. Canu and Bassler’s (1928) material from Brazil identified as M. ciliata differs from M. proxima because it has a transverse avicularium located between the orifice and ascopore, and very large autozooids. Another species very similar to Microporella proxima sp. nov. is M. monilifera Liu and Liu, 2003, the two species having avicularia and ascopores of the same shape and similar-sized autozooids. However, the Chinese species has up to five oral spines, a smooth proximal border to the orifice, smaller pseudopores, and the avicularium is located nearer to the ascopore. Winston (2005) described a new species of Microporella, M. protea, from the Atlantic with a distribution stretching south from Cape Hatteras to Florida, and possibly into the Caribbean. This species is similar to the new Brazilian species in its orifice shape, beaded proximal border, number and positions of oral spines. However, it differs in having wider zooids, in the shape and size of the avicularia, which are located more distally, and in the less elongate ascopore. The species Microporella gibbosula Canu and Bassler, 1930, originally described from the Galapagos Islands, is similar to M. proxima sp. nov. in both avicularium shape and position (located near the margin and the zooidal lateral angle) and the setose mandible. However, it differs in its larger autozooids (500–550 µm long and 450 µm wide), five distal spines, a small ascopore and smooth ovicells. Approximately 100 species of Microporella have been recognized, the number increasing as more use is made of SEM (Taylor and Mawatari 2005). Many established species remain to be characterized using SEM and some supposedly cosmopolitan species, notably M. ciliata, are undoubtedly complexes of more narrowly distributed species such as M. proxima sp. nov. Superfamily CELLEPOROIDEA Johnston, 1838 Family CELLEPORIDAE Busk, 1852 Genus Celleporina Gray, 1848 Celleporina diota (Marcus, 1938) (Figure 7) Siniopelta diota Marcus 1938: 48, pl. 11, fig. 27a–c. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Journal of Natural History 783 Figure 7. Celleporina diota (Marcus, 1938). (A) General view of the colony; (B) frontal view of the colony showing lateral avicularia, sinus, and ovicells; (C) ovicell detail. Scale bars: A: 300 µm; B: 100 µm; C: 50 µm. MNRJ-066. 784 L.V. Ramalho et al. Material examined Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Arraial do Cabo: MNRJ-066, Farol beach, Cabo Frio Island, 1 February 2000, collected by L.V. Ramalho, depth 5 m. Description Colony massive or nodular, small, white, with irregularly disposed autozooids (Figure 7A). Autozooids semi-erect, elongated with convex frontal walls, imperforate, except for large marginal areolar pores, varying from three to five along each margin, located in furrows (Figure 7A,B). Primary orifice longer than wide (121 µm long, range 116–126 µm, by 124 µm, range 116–136 µm), distally rounded and with a shallow U-shaped sinus on proximal border that is wider than deep (29 µm long, range 19–39 µm, by 56 µm, range 48–68 µm); condyles rounded (Figure 7B). Peristome becoming elevated in older zooids. Ovicell frontally flattened, closed by zooidal operculum, with pores of various shapes and sizes located between 13 and 16 longitudinal ribs crossed by one or two transverse ribs (Figure 7A–C). Avicularia paired, lateral to orifice, small 65 µm long (range 58–68 µm), located at tip of erect process, directed proximally outwards and upwards; rostrum oval, slightly pointed, indented at distal border, cross-bar complete (Figure 7A,B). Additional avicularium sometimes developed distally or proximally of the orifice (Figure 7A). Interzooidal avicularia rare (only one observed), large, 204 µm long by 87 µm wide, slipper-shaped, mandible elongate, rounded. Habitat Colonies found on Sargassum algae at depths of up to 5 m. Geographical distribution Apparently endemic to Brazil: Santos (SP) (Marcus 1938) and Arraial do Cabo (Farol beach), Rio de Janeiro State (present study). Remarks Three Celleporina species are known from Brazil: C. diota, C. costazii (Audouin, 1826) and C. langei (Marcus, 1939) (see Marcus 1937, 1938, 1939, 1941; Guimarães and Rosa 1941). Celleporina langei differs in having a weak peristome and orificial avicularia with smooth mandibles. Celleporina diota differs from C. costazii as the latter has more flattened autozooids, smaller frontal and marginal pores, no furrows at the zooidal borders, a sinus of different shape and narrow slits among the ribs of the ovicells. The species C. souleae Morris, 1979 has a narrower and deeper sinus and an ovicell with up to eight ribs. Celleporina robertsoniae (Canu and Bassler, 1923) has a very much deeper V-shaped sinus, more interzooidal avicularia, smaller marginal pores, and the ovicell is smaller, sometimes triangular and has ribs arising from the centre of the frontal area in rays. The species most similar to C. diota seems to be C. costata (MacGillivray, 1869). They differ as C. costata has rounded avicularia, less elevated peristomes, smaller Journal of Natural History 785 Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 marginal pores and a different sculpture on the ovicells. Marcus (1938) mentioned in his description of C. diota that the avicularia are turned distally and obliquely. In the specimen from Rio de Janeiro, the avicularia have the same shape, but they are directed slightly proximally. The other characteristics are very similar to those mentioned in Marcus’s description. As the only difference between the two species is in the position of the avicularia, the specimen from Rio de Janeiro can be considered to be C. diota. Celleporina diota was previously recorded from Santos Bay on the São Paulo coast by Marcus (1938). This is the first record of the species from the Rio de Janeiro coast. Like Marcus’s (1938) material, the Rio de Janeiro colonies occur on algae. The range of C. diota along the Brazilian coast may be greater, but because of its apparent preference for algae, it may be overlooked during sampling. Family PHILODOPORIDAE Gabb and Horn, 1862 Genus Reteporellina Harmer, 1933 Reteporellina evelinae Marcus, 1955 (Figure 8) Reteporellina evelinae Marcus 1955: 302, figs 66–69; Barbosa 1964: 32. Examined material Arraial do Cabo: MNRJ-174, Forno Harbor (Cais do Anel), 7 September 2003, collected by L.V. Ramalho, depth 5 m. Reteporellina delicatula: NHM 1975.1.12.311, Chios, Greece; NHM 87.12.9.465, Sandwich Island. Reteporellina idmoneoides: NHM 1862.2.4.22, eastern Australia. Description Colony erect, arborescent, branched, non-fenestrate, thickly calcified, reaching 1–1.5 cm high and 2 cm wide; coloured pale pink when alive but white when in alcohol (Figure 8A). Branches with autozooids disposed in quincunx, forming three to four alternate series on the frontal sides of the branches (Figure 8B). Autozooids longer than wide (405 µm long, range 353–470 µm, by 170 µm wide, range 167–176 µm), frontally convex, delineated by salient walls; frontal wall with one or two asymmetric pseudopores located below median line or absent altogether (Figure 8B–D). Peristome approximately tubular, tall, frontally and laterally well developed (Figure 8C,D), a median fissure occurring on one side and one or two protuberances similar to thick teeth on the other side (Figure 8E,F). Ovicell globose (169 µm long, range 167–176 µm, by 137 µm, range 127–147 µm), smooth, sometimes embedded in frontal wall; a labellum and median fissure present, the fissure ontogenetically variable in length and shape (Figure 8E,F). Labellum and flanges of equal length, sometimes hidden by development of peristome. Ovicell often covered by secondary calcification, but ectooecium in frontal region left exposed (Figure 8F). Avicularium inserted into peristome (Figure 8E,F), mandible triangular, bicuspidate; frontal avicularium absent. Dorsal avicularium located on some outer Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 786 L.V. Ramalho et al. Figure 8. Reteporellina evelinae Marcus, 1955. (A) General view of the colony; (B) branch bifurcation; (C) autozooids with well-developed peristomes; (D) frontal view of the branch showing autozooidal arrangement; (E) fertile autozooids and tooth-like structures on on the peristomes; (F) peristomial avicularium. Scale bars: A: 5 mm; B: 500 µm; C, E: 100 µm; D: 200 µm; F: 50 µm. MNRJ-174. autozooids; chamber elevated, well-developed, teeth bending to form a hook; mandible triangular, unicuspidate. Habitat Colony found on artificial substratum (concrete) in a sheltered environment, 5 m deep. Journal of Natural History 787 Geographical distribution Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 In Brazil, this species is recorded south of Vitoria, Espírito Santo State (Marcus 1955), and at Arraial do Cabo-Forno Harbour, Rio de Janeiro State (present study). Remarks Most species of Reteporellina are known from the Indo-Pacific, except for R. delicatula Hayward, 1974 (Mediterranean), R. marsupiata (Smitt, 1873) (West Atlantic) and R. evelinae Marcus, 1955 (Brazil). The Brazilian species differs from the others in the following features: R. delicatula has frontal avicularia and the internal border of the peristome is toothed; although R. marsupiata has a similar colour to R. evelinae, the peristome is less developed, frontal avicularia are elevated and the colony has a different shape. In R. idmoneoides Harmer, 1934 the peristomial sinus is invisible externally, the ovicells are small, as are the dorsal avicularia, and the peristome does not form the protuberances found in R. evelinae. Reteporellina capistrata Harmer, 1934 has labial pores; R. laxipes (Canu and Bassler, 1929) lacks avicularia; R. denticulata (Busk, 1884) has frontal and infra-fenestrae avicularia, flanges longer than the labellum, a chaliceshape colony, and less developed toothed peristome; and in R. bilabiata Osburn, 1952 the labial avicularium is absent, frontal avicularia are present, and frontal pseudopores more numerous. In comparison with the material of R. evelinae described by Marcus (1955) from Espírito Santo State, the Rio de Janeiro specimen has a pale pink colour (vitreous in the Espírito Santo specimens). However, other characteristics agree with Marcus’s description, and the material is therefore considered to belong to R. evelinae. This is the first record of R. evelinae from the Rio de Janeiro coast, extending its distribution along the Brazilian coast. Genus Reteporella Busk, 1884. Reteporella antennata sp. nov. (Figure 9) Material examined Holotype: MNRJ-175. Paratypes: MNRJ-236, MNRJ-237. REVIZEE Station 2 (23◦ 47.629′ S, 041◦ 42.514′ W), all collected by D.C. Savi from the naval ship Diadorim, 20 July 2004, using a trap, depth 341 m. Reteporella beaniana: NHM 1963.3.30.248, Lervik, Hardanger Fiord, Norway, Harmer Collection. Etymology The name antennata refers to the antenna shape of the distal spines. Diagnosis Colony fenestrate, chalice-like; frontal walls smooth, containing few marginal areolar pores; oral spines antenniform, six in number; ovicell with indistinct fissure and short Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 788 L.V. Ramalho et al. Figure 9. Reteporella antennata sp. nov. (A,B) Views of the cup-shaped colony; (C) distal branch showing autozooidal arrangement, antenna-like spines and labial pore; (D) autozooidal arrangement at bifurcation region; (E) ovicellate autozooids and frontal avicularia. F. Avicularia. Scale bars: A,B: 0.5 cm; C–E: 200 µm; F: 50 µm. MNRJ-175. and wide labellum; frontal avicularia monomorphic, oval with rounded mandible, proximally directed; dorsal avicularia of two kinds, one the same as the frontal avicularia, the second a giant avicularium with rounded mandible, attached below the fenestrae or on internal borders. Journal of Natural History 789 Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Description Colony erect, fenestrate, large (20 mm high by 30 mm diameter), four to five branches originating from a narrow base, bifurcating and anastomosing to form a circular, chalice-shaped net; white in colour (Figure 9A,B). Autozooids longer than wide (635 µm long, range 588–735 µm, by 257 µm wide, range 206–294 µm), convex, narrower distally, arranged in two series alternately on frontal side of branches; frontal wall smooth with a few marginal pores; secondary calcification present (Figure 9B,C). Peristome variably developed, hiding the primary orifice; a long fissure partially closed forming the labial pore which is large and round (Figure 9D). Six long peristomial spines (three external and three internal), articulated, antenniform; thicker base, narrowing towards the tip; complete in the distal zooids and often broken in other zooids (Figure 9B,C). Ovicell almost globose, hyperstomial, imperforate, wider than long, 196 µm long by 235 µm wide (range 216–255 µm), sometimes embedded in autozooidal calcification; labellum short and wide; fissures normally indistinct; secondary calcification present, usually not covering the entire frontal region (Figure 9D). Frontal avicularia small (94 µm long, range 78–118 µm, by 57 µm wide, range 49–69 µm), oval with short mandible rounded at the tip, cross-bar weak; located below zooid midline, directed proximally; one to two avicularia on frontal wall; avicularium chamber sometimes slightly elevated forming projection on frontal wall (Figure 9B–E). Dorsal avicularia of two kinds: one similar to frontal avicularium, small, with a mandible rounded at tip (Figure 9F); the other larger (245 µm long, range 216–274 µm, by 122 µm wide, range 98–147 µm), very rare, with mandible rounded at the tip, located immediately below fenestrae or between the dorsal and frontal sides near bifurcations. Habitat Found on rock and coralline substrates in deep water (341 m). Geographical distribution REVIZEE station 2 (23◦ 47.629′ S, 041◦ 42.514′ W), Rio de Janeiro State, Brazil. Remarks In the presence of antenniform spines and an oval frontal avicularium, this new species is similar to R. flabellata Busk, 1884. However, R. flabellata has a dichotomously branched colony which is not fenestrate, dorsal avicularia are absent, and branches are formed by one to three series of zooids. Hitherto, no species of Reteporella has been recorded from Brazil. Hayward (1993, 1995) mentioned nine Reteporella species in the Antarctic region, but all of these have large to giant avicularia and oral spines are absent or less frequent than in the new species. Gordon (1989) described three new species of Reteporella from New Zealand, but most of these are not fenestrate and differ in other characters from the Brazilian species. Hayward and Ryland (1996) described five Reteporella species from North Atlantic waters. These species differ from the new Brazilian species principally in having fewer spines, avicularia lateral or proximal of the orifice, and triangular avicularian mandibles. Gordon and d’Hondt (1997) described five species from New 790 L.V. Ramalho et al. Caledonia which also differ from the new Brazilian species as giant avicularia and oral spines are absent. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Genus Stephanollona Duvergier, 1920 (=Brodiella Uttley and Bullivant, 1972; Paracleidochasma Soule et al., 1991) Stephanollona robustaspinosa sp. nov. (Figure 10) Figure 10. Stephanollona robustaspinosa sp. nov. (A) General view of the colony; (B) part of the colony showing arrangement and shape of autozooids, marginal pores, oral spines, ovicells and dimorphic avicularia; (C) detail of distal region of an autozooid with small avicularia spines and an ovicell; (D) dimorphic avicularia and oral spines of different shapes; (E) detail of distal region of an autozooid with elongate avicularia. Scale bars: A: 1 mm; B: 200 µm; C–E: 100 µm. MNRJ-178. Journal of Natural History 791 Material examined Holotype: MNRJ-178; Paratype: MNRJ-241, REVIZEE Station 2 (23◦ 47.629′ S, 041◦ 42.514′ W), collected by D.C. Savi from the naval ship Diadorim using a trap, 20 July 2004, depth 341 m. Stephanollona longispinosa: holotype: NHM 99.5.1.1086, Hincks Collection. NHM 1990.11.20.19, 25 March 1927, depth 79 m. NHM 1991.4.18.5–6, Discovery station WS84, depth 75 m. Stephanollona armata: NHM 1936.12.30.215, Providence, Thornely Collection, depth 228 m. NHM 1966.1.12.16, Kullana Point, Malta. Etymology The name robustaspinosa refers to the strong, thick spines. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Diagnosis Colonies small, almost circular; frontal wall granular, with marginal areolar pores only; orificial sinus long, narrow; oral spines numbering six, either spear-shaped or tapering towards tips; ovicell imperforate, ectooecium with a broad, membranous frontal area, exposing the entooecium; a short, broad labellum present, defined by short lateral fissures; avicularia lateral to orifice dimorphic, one small, rounded with dented rostrum, the other setose, directed distally. Description Colony either small, roughly circular, semi-erect, generally with free borders, or larger, irregular in shape, closely encrusting substratum, up to 15 mm in diameter; translucent (Figure 10A). Autozooids almost hexagonal, a little longer than wide (782 µm long, range 637–862 µm, by 729 µm wide, range 441–902 µm wide (Figure 10B). Frontal wall granular, slightly convex, pseudopores absent, areolar pores around margin rounded; zooids separated by slender salient walls (Figure 10B,C). Orifice circular, slight dilated proximally; sinus long, slightly wider in some zooids; condyles present; lyrula absent; denticles developed around distal and lateral edges of orifice (Figure 10C); oral spines numbering six, basally articulated, irregularly calcified, dilated and flattened at distal ends of some, followed by narrowing towards tip, resembling a spear; other spines resembling needles; sometimes all of the oral spines become very elongated (Figure 10B–E). Ovicell slightly wider than long (232 µm long, range 225–235 µm, by 219 µm wide, range 205–235 µm), imperforate, not closed by operculum (Figure 10C), frontally slightly flattened, sometimes covered only by the entooecium (Figure 10B); labellum wide with fissure (Figure 10C). Avicularia oral, dimorphic, one on each side above proximal line of orifice (Figure 10B–E). Smaller avicularia measuring 98 µm long (range 88–108 µm) by 82 µm wide (range 78–88 µm); teeth well developed (Figure 10B–D). Large avicularia measuring 252 µm long (range 98–302 µm) by 101 µm wide (range 59–118 µm); mandible thin and elongated (265 µm long, range 157–294 µm), sometimes slightly bent at the tips, directed distally; rostrum with same shape as mandible (Figure 10B,D,E). The two types of avicularium seemingly occur randomly on autozooids: one small plus one large (Figure 10B), two small (Figure 10C), or two large (Figure 10E). 792 L.V. Ramalho et al. Habitat Colonies found on rock substrata and tubes of polychaetes, 6–10 m deep. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Geographical distribution REVIZEE Station 2 (23◦ 47.629′ S, 041◦ 42.514′ W), Rio de Janeiro State, Brazil. Remarks Two fossil species and a few Recent species distributed in the Atlantic and Pacific are known of this genus. Only one species has been recorded from Brazil: S. asper (Canu and Bassler, 1923), which was mentioned by Marcus (1937: 98) as Perigastrella contracta (Waters, 1899). Canu and Bassler’s species differs from S. robustaspinosa sp. nov. in having a shorter sinus, a large spatulate avicularium with rounded rostrum, small avicularia with rough rostra, unserrated, and in the position of the avicularia, shape of the distal spines, and colonies that may be multilamellar. The new species is more similar to S. longispinata (Busk, 1884) but differs mainly in having a median frontal avicularium. Other differences are the thinner oral spines, which are generally longer, large avicularium directed to the sides and/or distally, and the much narrower labellum. The species S. orbicularis (Hincks, 1881) has a wider sinus with an avicularium just below it. Differences between S. robustaspinosa sp. nov., S. armata (Hincks, 1861) and S. ignota Hayward and Cook, 1983 are in avicularium shape and position, and oral spine number and shape. Although S. serrata (Osburn, 1912) has small avicularia with serrated rostra, this species differs from S. robustaspinosa sp. nov. in having a shorter and wider sinus, only four distal spines, and avicularia oriented in several directions. Superfamily CONESCHARELLINOIDEA Levinsen, 1909 Family LEKYTHOPORIDAE Busk, 1884 Genus Turritigera Busk, 1884 Turritigera buski sp. nov. (Figures 11, 12) Material examined Holotype: MNRJ-182; Paratype: MNRJ-240, REVIZEE Station 9 (24◦ 16.353′ S, 043◦ 23.788′ W), collected by D.C. Savi from naval ship Diadorim with a trap, 24 July 2004, depth 600 m. Turritigera stellata: NHM 1887.12.9.520, 1944.1.8.240, Challenger, Station 320, depths 1100 m and 600 m, respectively. NHM 1934.2.16.13, Challenger, Cape Hope, station 142, depth 275 m. Turritigera cribrata: Holotype: NHM 1992.10.19.7, Discovery Expedition station 42, depth 120–204 m. Paratype: NHM 1992.10.19.8, British Antarctic Expedition, station 355, Terra Nova. NHM 1997.8.1.9, Discovery Expedition. NHM 1997.8.1.10, McMurdo Sound, Ross Sea. Turritigera fenestella: Holotype: NHM 1890.4.16.2B, Marion Island (38◦ S, 046◦ 40′ W). Turritigera reticulate: Holotype: NHM 1890.4.16.2A, Marion Island (38◦ S, 046◦ 40′ W). Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 Journal of Natural History 793 Figure 11. Turritigera buski sp. nov. (frontal view). (A) General frontal view of the colony; (B) frontal view showing autozooids; (C) form and disposition of the autozooids, peristomial avicularia and marginal pores (larger) and small pseudopores spread on the frontal wall; (D) detail of the primary orifice, showing U-shaped sinus; (E) detail of peristome showing form and disposition of avicularia; (F) peristome with one avicularium. Scale bars: A: 3 mm; B: 900 µm; C, F: 100 µm; D, E: 60 µm. MNRJ-182. Etymology The name buski is in honour of George Busk who first described species of Turritigera. Diagnosis Colony arborescent, white, with two or three series of zooids opening on frontal sides of branches. Autozooids with large sinus, slightly wider than deep; peristome well Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 794 L.V. Ramalho et al. Figure 12. Turritigera buski sp. nov. (dorsal view). (A) General view; (B) pseudopores and small avicularia; (C) bifurcation showing a large, spatulate avicularium; (D) detail of large avicularium; (E) ovicell (frontal side). Scale bar: A: 400 µm; B, E: 100 µm; C: 300 µm; D: 60 µm. MNRJ-182. developed. Ovicell proximal, globose, opening inside peristome. Peristomial avicularia numbering one to five, the most distal being largest; small avicularia, similar to peristomial avicularia, located on frontal and dorsal sides; large avicularia only on dorsal side, spatulate. Description Colony arborescent, dichotomously branched, strongly calcified, up to 30 mm high, white in colour (Figure 11A). Autozooids opening on one side of branch (frontal), in series of up to three (Figure 11B), elongate, longer than wide, not separated by sutures, delimited by large Journal of Natural History 795 Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 pores (Figure 11C); peristome variably developed, sometimes very elevated, normally covering primary orifice (Figure 11C) which has a narrow to moderately wide sinus (38 µm long by 54 µm wide), well delimited (Figure 11D). A few small pseudopores scattered on frontal surface (Figure 11C–F). In some zooids, a large fissure occurs immediately below the peristome (Figure 11F). Peristomial avicularia numbering one to five, depending on ontogenetic stage; distal avicularium sometimes a little larger than the others, all avicularia turned outwards, sometimes on a process elevated above surface of peristome (Figure 11C,E,F). Other small avicularia, similar to peristomial avicularia, scattered across dorsal and frontal branch surface (Figure 11B, 12A,B), mandible triangular. Giant avicularia occur on dorsal sides (300 µm long), the sole example observed located at a bifurcation (Figure 12C,D); mandible dilated and rounded at tip, turned to the side, facing slightly downwards; cross-bar complete. Ovicell prominent, located proximally of orifice, surface similar to the zooidal frontal wall, opening inside the peristome; small pores present (Figure 12E). Habitat Colonies found on rock at 600 m deep, fouled by small colonies of Patinella sp. Geographical distribution REVIZEE Station 9 (24◦ 16.353′ S, 043◦ 23.788′ W), Rio de Janeiro State, Brazil. Remarks Five species of the genus Turritigera have been described previously: T. cribrata Hayward, 1993, T. spectabilis d’Hondt, 1981, T. reticulata Cook and Hayward, 1983, T. fenestella Cook and Hayward, 1983 and T. stellata Busk, 1884. The four first species are clearly different from T. buski (see d’Hondt 1981a and b; Cook and Hayward 1983; Hayward 1993). Busk’s species T. stellata, as redescribed by Cook and Hayward (1983) and Hayward (1993) from two different Challenger stations (Stn 320 from Uruguay, and Stn 142 from South Africa), is variable. According to Hayward (1993), the specimen from Uruguay has a more delicate colony, narrower and U-shaped sinus, peristomial avicularia of equal size and a dorsal avicularium that is large and spatulate, whereas the specimen from South Africa has a more robust colony, large and shallow sinus, the distal peristomial avicularium is larger than the others, and large dorsal avicularia are lacking. If it were not for the different shape of the sinus, specimens from the two stations could be assigned to the same species as there may be variation among the types of peristomial avicularia in the same colony and the dorsal avicularia may be rare. However, the shape of the sinus is clearly important taxonomically (see Hayward 1993: 1427), and this feature argues against the specimens from Uruguay and South Africa being conspecific. Turritigera buski sp. nov. is close to putative T. stellata from Uruguay but T. stellata has more (up to seven) peristomial avicularia which are all very similar in size, lacks small triangular avicularia on the dorsal side, has interzooidal avicularia on the frontal shield, and, importantly, a primary orifice with a narrow, U-shaped median sinus (40 µm long by 30 µm wide; Hayward 1993, table 1, fig. 7D). 796 L.V. Ramalho et al. Downloaded By: [Ramalho, Lais] At: 22:43 24 March 2011 This is the first record of Turritigera from the Brazilian coast. The lack of previous records may reflect the paucity of studies of bryozoan faunas from deep water where Turritigera is typically found (d’Hondt 1981a,b; Cook and Hayward 1983; Gordon 1989; Hayward 1995). Almost all species from the southern hemisphere have been recorded in deep (up to 4300 m) and cool (−0.3 to +17◦ C) water; hitherto tropical or subtropical species were unknown (d’Hondt 1981a; Cook and Hayward 1983; Hayward 1993). Conclusion This study of lepraliomorph cheilostomes from Rio de Janeiro State describes 10 species, five of which are new: Parasmittina alba, Microporella proxima, Reteporella antennata, Stephanollona robustaspinosa and Turritigera buski. The genera Reteporella and Turritigera are newly recorded from Brazilian waters. Of the species previously recorded from Brazil, Arthropoma cecilii, Celleporina diota and Reteporellina evelinae are new occurrences for Rio de Janeiro State. The diversity of bryozoans known from Brazil and Rio de Janeiro State is increased: before this study, 69 species were recorded from Rio de Janeiro State (Barbosa 1964; Buge 1979; Ramalho et al. 2005, 2008a,b, 2009); this number now rising to 79 species. It is likely that far more species await discovery in this poorly sampled region. Arraial do Cabo was found to have the greatest local diversity of lepraliomorph species, containing seven of the 10 species described here. Most of these species were found in small areas, except for S. errata and W. subovoidea, which were more widespread and were collected in four areas: Macaé, Arraial do Cabo, Guanabara Bay and Sepetiba Harbor. Both species were in reproductive condition and were normally found in polluted harbour environments, as they are elsewhere in the world. Acknowledgements Thanks are due to: the Instituto de Estudos do Mar Almirante Paulo Moreira and David C. Savi for collection of samples during the REVIZEE Program Pesca; Mary Spencer-Jones for help with the recent bryozoan collections at the NHM, London; Prof. John Ryland and Dr Joshua Mackie for information about Watersipora species; and Drs Peter Hayward and John Ryland for figure 4F. 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