Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 4434 No. 3: 18 Jun. 2018
Type: Article
Published: 2018-06-18
DOI: 10.11646/zootaxa.4434.3.1
Page range: 401–428
Abstract views: 57
PDF downloaded: 1

Bryozoa on disarticulated bivalve shells from Todos os Santos Bay, northeastern Brazil, with the description of two new species

Laboratório de Estudos de Bryozoa—LAEBry, Departamento de Zoologia, Centro de Biociências, Universidade Federal de Pernambuco, Recife, PE, 50670–810, Brazil. Museu de Zoologia da Universidade Federal da Bahia, Universidade Federal da Bahia, Salvador, BA, 40170–290, Brazil. Laboratório de Geoecologia dos Sedimentos Marinhos—GEOECO, Departamento de Zoologia, Instituto de Biologia, Universidade Federal da Bahia, Salvador, BA, 40170–290, Brazil.
Museu de Zoologia da Universidade Federal da Bahia, Universidade Federal da Bahia, Salvador, BA, 40170–290, Brazil. Laboratório de Geoecologia dos Sedimentos Marinhos—GEOECO, Departamento de Zoologia, Instituto de Biologia, Universidade Federal da Bahia, Salvador, BA, 40170–290, Brazil.
Museu de Zoologia da Universidade Federal da Bahia, Universidade Federal da Bahia, Salvador, BA, 40170–290, Brazil. Laboratório de Geoecologia dos Sedimentos Marinhos—GEOECO, Departamento de Zoologia, Instituto de Biologia, Universidade Federal da Bahia, Salvador, BA, 40170–290, Brazil.
Laboratório de Geoecologia dos Sedimentos Marinhos—GEOECO, Departamento de Zoologia, Instituto de Biologia, Universidade Federal da Bahia, Salvador, BA, 40170–290, Brazil.
Laboratório de Estudos de Bryozoa—LAEBry, Departamento de Zoologia, Centro de Biociências, Universidade Federal de Pernambuco, Recife, PE, 50670–810, Brazil.
Bryozoa Cheilostomata Itaparica new combination Phidoloporidae Smittinidae

Abstract

As sessile organisms, bryozoans are among the commonest components of the encrusting communities on hard substrata. On unconsolidated bottoms, benthic animals and their carapaces, including mollusk shells, represent potential surfaces for encrustation. Here we describe the bryozoan fauna growing on disarticulated bivalve shells collected intertidally on Itaparica beach, Todos os Santos Bay, northeast Brazil. We examined a total of 382 valves representing seven species of bivalves, among which we found 82 valves of three bivalve species to be encrusted by bryozoans, with 109 bryozoan colonies detected overall. We identified 11 cheilostome bryozoan species, among which Phidoloporidae and Smittinidae were the most diverse, with four and two species, respectively. We redescribed two bryozoan species, Parasmittina loxoides and Rhynchozoon phrynoglossum, and describe another two, Pleurocodonellina marcusi n. sp. and Rhynchozoon itaparicaensis n. sp., as new. A new combination for Cribella triangulata Canu & Bassler, 1928 is proposed, including Calyptotheca tenuata Harmer, 1957 as a new junior subjective synonym of Calyptotheca triangulata n. comb. All but one of the 11 species have encrusting colonies; Licornia aff. diadema, forms erect branching colonies. For seven of the bryozoans, ours are the first records of these species growing on shells. Although the majority of bryozoans we studied have also been reported on other substrata, Rhynchozoon brasiliensis and Parasmittina loxoides were predominant in the fauna, comprising 79 (72%) of all colonies inventoried. Due to the lack of information on the distribution of these species in northeastern Brazil, further studies are needed to determine whether they show any substratum preference.

 

References

  1. Allman, G.J. (1856) A monograph of the Freshwater Polyzoa, including all the known species, both British and Foreign. The Ray Society, London, 119 pp.
    https://doi.org/10.5962/bhl.title.9143

    Almeida, A.C.S., Alves, O., Peso-Aguiar, M., Dominguez, J. & Souza, F.B.C. (2015a) Gymnolaemata bryozoans of Bahia State, Brazil. Marine Biodiversity Records, 8, e120.
    https://doi.org/10.1017/S1755267215000743

    Almeida, A.C.S., Souza, F.B.C., Sanner, J. & Vieira, L.M. (2015b) Taxonomy of recent Adeonidae (Bryozoa, Cheilostomata) from Brazil, with the description of four new species. Zootaxa, 4013 (3), 348–368.
    https://doi.org/10.11646/zootaxa.4013.3.2

    Almeida, A.C.S., Souza, F.B.C., Gordon, D. & Vieira, L.M. (2015c) The non-indigenous bryozoan Triphyllozoon Canu & Bassler, 1917 in the Atlantic: morphology and dispersion on the Brazilian coast. Zoologia, 32 (6), 476–484.
    https://doi.org/10.1590/S1984-46702015000600007

    Almeida, AC.S., Souza, F.B.C., Menegola, C. & Vieira, L.M. (2017a) Diversity of marine bryozoans inhabiting demosponges in Northeastern Brazil. Zootaxa, 4290 (2), 281–323.
    https://doi.org/10.11646/zootaxa.4290.2.3

    Almeida, A.C.S., Souza, F.B.C. & Vieira, L.M. (2017b) Malacostegine bryozoans (Bryozoa: Cheilostomata) from Bahia State, northeast Brazil: taxonomy and non-indigenous species. Marine Biodiversity, 1–26. [published online]
    https://doi.org/10.1007/s12526-017-0639-x

    Alves, O.F.S., Muehe, D. & Dominguez, J.M.L. (2006) Carbonate contents of bottom sediments of Todos os Santos Bay, Bahia, Brazil: their importance for biodiversity. Journal of Coastal Research, Special Issue 39, 1671–1675.

    Barnes, D.K.A. & Clarke, A. (1995) Epibiotic communities on sublittoral macroinvertebrates at Signy Island, Antarctica. Journal of the Marine Biological Association of the United Kingdom, 75, 689–703.
    https://doi.org/10.1017/S0025315400039102

    Barros, F., Costa, P.C., Cruz, I., Mariano, D.L.S. & Miranda, R.J. (2012) Habitats Bentônicos na Baía de Todos os Santos. Revista Virtual de Química, 4 (5), 551–565.

    Bernadochi, L.C., Alves, J.L. & Marques, H.L.A. (2014) Settlement of juveniles of pearl oyster (Pinctada imbricata) on artificial collectors in Caraguatatuba, São Paulo, Brazil. Aquaculture Research, 47 (2), 1–9.
    https://doi.org/10.1111/are.12503

    Bishop, J.D.D. (1988) Disarticulated bivalve shells as substrates for encrustation by the bryozoan Cribrilina puncturata in the Plio-Pleistocene Red Crag of Eastern England. Palaeontology, 31, 237–253.

    Busk, G. (1852) An account of the Polyzoa, and sertularian zoophytes, collected in the Voyage of the Rattlesnake, on the coasts of Australia and the Louisiade Archipelago. In: MacGillivray, J. (Ed.), Narrative of Voyage of H.M.S. Rattlesnake 1. T.W. Boone, London, pp. 343–402.

    Busk, G. (1856) Zoophytology. Quarterly Journal of Microscopical Science, 4, 176–179.

    Busk, G. (1884) Report on the Polyzoa collected by H.M.S. Challenger during the years 1873–1876. Part 1. The Cheilostomata. Report on the Scientific Results of the Voyage of the H.M.S. Challenger during the years 1873-–76, Zoology, 10 (30), I–XXIV + 1–216, 36 pls.

    Canu, F. & Bassler, R.S. (1927) Bryozoaires des Iles Hawai. Bulletin de la Société des Sciences de Seine-et-Oise, 8 (2), 1–23.

    Canu, F. & Bassler, R.S. (1928) Bryozoaires du Brésil. Bulletin de la Société des Sciences de Seine-et-Oise, 7, 58–110.

    Cumming, R.L. & Tilbrook, K.J. (2014) Six species of Calyptotheca (Bryozoa, Cheilostomata, Lanceoporidae) from the Gulf of Carpentaria and northern Australia, with description of a new species. Zootaxa, 3827 (2), 147–169.
    https://doi.org/10.11646/zootaxa.3827.2.2

    Cumming, R.L. (2015) Two tropical species of Stephanotheca (Bryozoa, Cheilostomata, Lanceoporidae) from the Gulf of Carpentaria, Australia. Zootaxa, 3948 (2), 279–286.
    https://doi.org/10.11646/zootaxa.3948.2.8

    De Blauwe, H. (2006) Bryozoa on shells from the Kwintebank, Southern bight of the North Sea (Belgium). Bulletin Koninklijk Belgisch Instituut voor Natuurwetenschappen, 76, 125–138.

    Denisenko, N.V., Thomsen, E. & Tendal, O.S. (2013) Bryozoan epifauna on brachiopods from the Faroe Islands (NE Atlantic). Fróðskaparrit, 60, 96–113.
    https://doi.org/10.18602/fsj.v60i0.9

    Díaz, O.D. & Liñero-Arana, I. (2003) Poliquetos epibiontes de Pinctada imbricata Röding, 1798 (Bivalvia: Pteriidae) en el Golfo de Cariaco, Venezuela. Interciencia, 28 (5), 298–301.

    Dick, M. & Grischenko, A.V. (2016) Rocky-intertidal cheilostome bryozoans from the vicinity of the Sesoko Biological Station, west-central Okinawa, Japan. Journal of Natural History, 51 (3–4), 141–266.
    https://doi.org/10.1080/00222933.2016.1253797

    Farias, M.C. & Rocha-Barreira, C.A. (2012) Aspectos populacionais de Tagelus plebeius (Mollusca: Bivalvia: Solecurtidae) no estuário do Rio Ceará, Nordeste do Brasil. Arquivos de Ciência do Mar, 45 (2), 32–39.

    Gabb, W.M. & Horn, G.H. (1862) The fossil Polyzoa of the Secondary and Tertiary Formations of North America. Journal of the Academy of Natural Sciences of Philadelphia, 5, 111–179.

    Gautier, Y.-V. (1957) Bryozoaires des Iles Baléares. Résultats des Campagnes Prof. Lacaze-Duthiers. Vie et Milieu, 2 (6), 205–222.

    Gmelin, J.F. (1791) Caroli a Linné: systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus II. Aucta Reformata. Impensis George Emanuel Beer, Lipsiae [Leipzig], 890 pp. [pp. 3021–3910]

    Gmelin, J.F. (1792) Caroli a Linné: systema naturae per regna tria naturae. Vol. 1. Part 6. Vermes. 13th Edition. Impensis George Emanuel Beer, Lipsiae, 890 pp. [pp. 3021–3910]

    Gordon, D.P. (2017) Bryozoa: Cheilostomata. Interim classification for Treatise, version of 8 September 2017. Institute of Water & Atmospheric Research, Wellington, 16 pp. [unpublished but available from the author]

    Gray, J.E. (1848) List of the Specimens of British Animals in the Collections of the British Museum. Part 1. Centroniae or Radiated Animals. Trustees of the British Museum (Natural History), London, XIV + 172 pp.
    https://doi.org/10.5962/bhl.title.132951

    Guimarães, M.S.C. & Rosa, C.N. (1941) Sobre a briozoo fauna duma Santola, Mithrax hispidus (Herbst). Boletim da Faculdade de Filosofia, Ciências e Letras, Universidade de São Paulo, Zoologia, 5, 305–313.

    Harmelin, J.G., Vieira, L.M., Ostrovsky, A.N., Cáceres-Chamizo, J.P. & Sanner, J. (2012) Scorpiodinipora costulata (Canu & Bassler, 1929) (Bryozoa, Cheilostomata), a taxonomic and biogeographic dilemma: complex of cryptic species or human-mediated cosmopolitan colonizer? Zoosystema, 34 (1), 123–138.
    https://doi.org/10.5252/z2012n1a5

    Harmer, S.F. (1957) The Polyzoa of the Siboga Expedition, Part 4. Cheilostomata Ascophora II. Siboga Expedition Reports, 28d, 641–1147.

    Hayward, P.J. (1993) New species of cheilostomate Bryozoa from Antarctica and the Subantarctic southwest Atlantic. Journal of Natural History, 27, 1409–1430.
    https://doi.org/10.1080/00222939300770771

    Hayward, P.J. & Cook, P.L. (1983) The South African Museum's Meiring Naude Cruises. Part 13, Bryozoa II. Annals of the South African Museum, 91, 1–161.

    Hayward, P.J. & Ryland, J.S. (1995) Bryozoa from Heron Island, Great Barrier Reef. 2. Memoirs of the Queensland Museum, 38, 533–573.

    Hayward, P.J. & Ryland, J.S. (1998) Cheilostomatous Bryozoa. Part 1. Aeteoidea-Cribrilinoidea. In: Barnes, R.S.K & Crothers, J.H. (Eds.), Synopses of the British Fauna. New Series. No. 10. Field Studies Council, Shrewsbury, pp. 1–366.

    Hernández-Ávila, I., Tagliafico, A. & Rago, N. (2013) Composición y estructura de la macrofauna asociada con agregaciones de dos especies de bivalvos en Isla de Cubagua, Venezuela. Revista de Biología Tropical, 61 (2), 669–682.

    Hincks, T. (1882) Polyzoa of the Queen Charlotte Islands: Preliminary notice of new species. Annals and Magazine of Natural History, Series 5, 10 (57), 248–256.
    https://doi.org/10.1080/00222938209459702

    Johnston, G. (1847) A History of British Zoophytes. Vol. 1 & 2. Van Voorst, London, 488 pp., LXXIV pls.
    https://doi.org/10.5962/bhl.title.19627

    Jullien, J. & Calvet, L. (1903) Bryozoaires provenent des campagnes de l’Hirondelle. Résultats des Campagnes Scientifiques Accompli par le Prince Albert I, 23, 1–188.

    Kuklinski, P. & Barnes, D.K.A. (2005) Microhabitat diversity of Svalbard Bryozoa. Journal of Natural History, 39 (7), 539–554.
    https://doi.org/10.1080/00222930400001350

    Kuklinski, P., Gulliksen, B., Lønne, O.J. & Weslawski, J.M. (2005) Composition of bryozoan assemblages related to depth in Svalbard fjords and sounds. Polar Biology, 28, 619–630.
    https://doi.org/10.1007/s00300-005-0726-5

    Kuklinski, P., Gulliksen, B., Lønne, O.J. & Weslawski, J.M. (2006) Substratum as a structuring influence on assemblages of Arctic bryozoans. Polar Biology, 29, 652–661.
    https://doi.org/10.1007/s00300-005-0102-5

    Kuklinski, P., Barnes, D.K.A. & Wlodarska-Kowalczuk, M. (2008) Gastropod shells, hermit crabs and Arctic bryozoan richness. In: Hageman, S.J., Key Jr, M.M.J. & Winston, J.E. (Eds.), Proceedings of the 14th International Bryozoology Association Conference. Special Publication of the Virginia Museum of Natural History. No. 15. Virginia Museum of Natural History, Martinsville, Virginia, pp. 101–110.

    Lamarck, J.B.P.A.M. de (1801) Système des animaux sans vertèbres. L'Auteur, Paris, 432 pp.
    https://doi.org/10.5962/bhl.title.14255

    Lessa, G. & Dias, K. (2009) Distribuição espacial das litofácies de fundo da Baía de Todos os Santos. Quaternary and Environmental Geosciences, 1 (2), 84–97.
    https://doi.org/10.5380/abequa.v1i2.14376

    Levinsen, G.M.R. (1909) Morphological and systematic studies on the cheilostomatous Bryozoa. Nationale Forfatterers Forlag, Copenhagen, 431 pp.

    Lightfoot, J. (1786) A catalogue of the Portland Museum, lately the property of the Duchess Dowager of Portland, deceased: Which will be sold by auction by Mr. Skinner and Co. On Monday the 24th of April, 1786, and the thirty-seven following days (...) at her late dwelling-house, in Privy-Garden, Whitehall, by order of the Acting Executrix. Skinner, London, 214 pp.

    Linnaeus, C. (1758) Systemae naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differetiis, synonymis, locis. 10th Edition. Laurentii Salvii, Holmiae, 795 pp.
    https://doi.org/10.5962/bhl.title.542

    Linnaeus, C. (1767) Systemae naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differetiis, synonymis, locis Regnum Animale. Vol. 1 (2). 12th Edition. Laurentii Salvii, Holmiae, 824 pp.

    Liu, X., Yin, X. & Ma, J. (2001) Biology of Marine-fouling Bryozoans in the Coastal Waters of China. Science Press, Beijing, China, 860 pp.

    Machado, A.J. & Souza, F.B.C. (1994) Principais espécies de foraminíferos e briozoários do Atol das Rocas. Revista Brasileira de Geociências, 24, 247–261.

    Marcus, E. (1937a) Bryozoarios marinhos brasileiros I. Boletim da Faculdade de Filosofia, Ciências e Letras da Universidade de São Paulo, Zoologia, 1, 5–224.
    https://doi.org/10.11606/issn.2526-4877.bsffclzoologia.1937.113912

    Marcus, E. (1937b) Bryozoen von St. Helena. In Papers from Doktor Th. Mortensen’s Pacific Expedition 1914–16, LXIX. Videskabelige Meddelelser fra Dansk Naturhistorisk Forening I Kjøbenhavn, 101, 183–252.

    Marcus, E. (1938) Bryozoarios marinhos brasileiros II. Boletim da Faculdade de Filosofia, Ciências e Letras, Universidade de São Paulo, Zoologia, 2, 1–137.
    https://doi.org/10.11606/issn.2526-4877.bsffclzoologia.1938.113913

    Marcus, E. (1939) Bryozoarios Marinhos Brasileiros, 3. Boletim da Faculdade de Filosofia, Ciências e Letras da Universidade de São Paulo, Zoologia, 3, 111–353.

    Marcus, E. (1955) Notas sobre briozoos marinhos brasilieros. Arquivos do Museu Nacional do Rio de Janeiro, 42, 273–342.

    Marques, R.C. & Barbieri, E. (2015) Anatomical differences among specimens of Pinctada imbricata Röding, 1798 from different South American localities. Boletim do Instituto de Pesca, 4, 751–761.
    https://doi.org/10.20950/1678-2305.2015v41n.ep751

    Marshall, D.J. & Keough, M.J. (2003) Variation in the dispersal potential of non-feeding invertebrate larvae: the desperate larva hypothesis and larval size. Marine Ecology Progress Series, 255, 145–153.
    https://doi.org/10.3354/meps255145

    Miranda, A.A., Almeida, A.C.S. & Vieira, L.M. (2018)Non-native marine bryozoans (Bryozoa: Gymnolaemata) in Brazilian waters: Assessment, dispersal and impacts. Marine Pollution Bulletin, 130, 184–191.
    https://doi.org/10.1016/j.marpolbul.2018.03.023

    McKinney, F.K. (1996) Encrusting organisms on co-occurring disarticulated valves of two marine bivalves: comparison of living assemblages and skeletal residues. Paleobiology, 22 (4), 543–567.
    https://doi.org/10.1017/S0094837300016523

    Milne Edwards, H. (1836) Histoire des polypes. In: Deshayes, G.P. & Milne Edwards, H. (Eds) Histoire naturelle des animaux sans vertebres par J.- B.P.A. Lamarck, 2e edition revue et aug- mentee. J.B. Bailliere, London, 684 pp.

    Norman, A. (1903) Notes on the natural history of East Finmark. Polyzoa. Annals and Magazine of Natural History, Series 7, 11 (66), 567–598.
    https://doi.org/10.1080/00222930308678818

    Osburn, R.C. (1927) The Bryozoa of Curacao. Bijdr Dierkunde, Amsterdam, 25, 123–132.

    Orbigny, A. d’ (1851) Récherches zoologiques sur la classe des Molluscques Bryozoaires. Annales des Sciences naturelles, Zoologie & Biologie Animale, 3, 292–339.

    Röding, P.F. (1798) Museum Boltenianum sive catalogus cimeliorum e tribus regnis naturæ quæ olim collegerat Joa. Fried Bolten, M. D. p. d. per XL annos proto physicus Hamburgensis. Pars secunda continens conchylia sive testacea univalvia, bivalvia & multivalvia. Typis Johan Christi Trappi, Hamburg, 199 pp.

    Ryland, J.S. & Hayward, P.J. (1992) Bryozoa from Heron Island, Great Barrier Reef.
    Memoirs of the Queensland Museum, 32, 223–301.

    Simone, L.R.L. (2017) Bivalvia in Catálogo Taxonômico da Fauna do Brasil. PNUD. Available from: http://fauna.jbrj.gov.br/fauna/faunadobrasil/31 (accessed 7 May 2017)

    Smitt, F.A. (1873) Floridan Bryozoa collected by Count L.F. de Pourtales, Part 2. Kongliga Svenska Vetenskaps–Akademiens Handlingar, 11, 1–83.

    Soares, L.S.H., Salles, A.C.R., Lopez, J.P., Muto, E.Y. & Giannini, R. (2009) Pesca e produção pesqueira. In: Hatje, V. & Andrade, J.B. (Eds.), Baía de Todos os Santos, Aspectos oceanográficos. EDUFBA, Salvador da Bahia, pp. 158–206.

    Soule, D.F. & Soule, J.D. (1973) Morphology and speciation of Hawaiian and eastern Pacific Smittinidae (Bryozoa, Ectoprocta). Bulletin of the American Museum of Natural History, 152 (6), 365–440.

    Soule, D.F., Soule, J.D. & Chaney, H.W. (1995) The Bryozoa. Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel, 13, 1–344.

    Soule, D.F. & Soule, J.D. (2002) The eastern Pacific Parasmittina trispinosa complex (Bryozoa, Cheilostomatida): new and previously described species. Irene McCulloch Foundation Monograph Series, 5, 1–40.

    Souza, F.B.C. (1989) Espécies de briozoários da Bahia. Anais do XI Congresso Brasileiro de Paleontologia, 1989, 493–507.

    Taylor, J.D. & Glover, E.A. (2016) Lucinid bivalves of Guadeloupe: diversity and systematics in the context of the tropical Western Atlantic (Mollusca: Bivalvia: Lucinidae). Zootaxa, 4196 (3), 301–380.
    https://doi.org/10.11646/zootaxa.4196.3.1

    Tilbrook, K.J., Hayward, P.J. & Gordon, D.P. (2001) Cheilostomatous Bryozoa from Vanuatu. Zoological Journal of the Linnean Society, 131 (1), 35–109.
    https://doi.org/10.1006/zjls.2000.0249

    Tilbrook, K.J. (2006) Cheilostomatous Bryozoa from the Solomon Islands. Santa Barbara Museum of Natural History, Monographs 4 (Studies in Biodiversity Number 3), 4, 1-–386.

    Tilbrook, K.J. & Vieira, L.M. (2012) Scrupocellaria (Bryozoa: Cheilostomata) from the Queensland coast, with the description of three new species. Zootaxa, 3528, 29–48.

    Vieira, L.M., Migotto, A.E. & Winston, J.E. (2008) Synopsis and annotated checklist of Recent marine Bryozoa from Brazil. Zootaxa, 1810, 1–39.

    Vieira, L.M., Nascimento, K.B. & Almeida, A.C.S. (2018) Smittinidae in Catálogo Taxonômico da Fauna do Brasil. PNUD. Available from: http://fauna.jbrj.gov.br/fauna/faunadobrasil/157991 (accessed 20 March 2018)

    Vigneaux, M. (1949) Révision des Bryozoaires néogènes du Bassin d'Aquitaine et essai de classification. Mémoires de la Société Géologique de France, 28, 1–153.

    Ward, M.A. & Thorpe, J.P. (1989) Assessment of space utilisation in a subtidal temperate bryozoan community. Marine Biology, 103, 215–224.
    https://doi.org/10.1007/BF00543350

    Ward, M.A. & Thorpe, J.P. (1991) Distribution of encrusting bryozoans and other epifauna on the subtidal bivalve Chlamys opercularis. Marine Biology, 110, 253–259.
    https://doi.org/10.1007/BF01313711

    Waters, A.W. (1889) Bryozoa from New South Wales, North Australia, etc. Part 4. Annals and Magazine of Natural History, Series 6, 4 (19), 1–24.
    https://doi.org/10.1080/00222938909460465

    Winston, J.E. & Håkansson, E. (1986) The Interstitial bryozoan fauna from Capron Shoal, Florida. American Museum Novitates, 2865, 1–50.

    Winston, J.E. & Vieira, L.M. (2013) Systematics of interstitial encrusting bryozoans from southeastern Brazil. Zootaxa, 3710 (2), 101–146.
    https://doi.org/10.11646/zootaxa.3710.2.1

    Winston, J.E., Vieira, L.M. & Woollacott, R.M. (2014) Scientific results of the Hassler expedition. Bryozoa. No. 2. Brazil. Bulletin of the Museum of Comparative Zoology, 161 (5), 139–239.
    https://doi.org/10.3099/MCZ14.1