Abstract
In Southeastern Brazil ecoregion, the assessment of calcareous sponges diversity has been particularly focused on Rio de Janeiro State, with 26 species recorded, while São Paulo State remains relatively unexplored, with only 11 species known up to date. São Sebastião, located on the northern coast of São Paulo, encompassing the Alcatrazes Archipelago, is an area of high ecological relevance. This study investigates the diversity of calcareous sponges within two Marine Protected Areas in São Sebastião region, integrating morphological and molecular approaches. A total of 18 species were identified, including two of the subclass Calcinea and 16 of the subclass Calcaronea. Among them, six are new to science: Ascandra arenaria sp. nov., Amphoriscus tenax sp. nov., Leucandra anoducta sp. nov., Leucosolenia sebastianensis sp. nov., Sycon caissarum sp. nov., and Sycon crassapicale sp. nov. Additionally, other eight species represent new records for São Paulo, with Leucandra caribea being reported for the first time in Brazil, extending their distribution ranges. The remaining four species had previously been recorded in the region. Furthermore, as an important additional morphological character, we are originally reporting the presence of a few tetractines, alongside triactines, in the atrial skeleton of the invasive species Paraleucilla magna. Our findings provide a comprehensive baseline inventory of calcareous sponge species from the São Sebastião area (Ilhabela and São Sebastião, including the Alcatrazes Archipelago) and contribute to understanding the biogeographical affinities of the calcarean fauna of Southeastern Brazil with regions within and beyond the Atlantic Ocean.
References
- Alvizu, A., Eilertsen, M.H., Xavier, J.R. & Rapp, H.T. (2018) Increased taxon sampling provides new insights into the phylogeny and evolution of the subclass Calcaronea (Porifera, Calcarea). Organisms Diversity and Evolution, 18, 279–290. https://doi.org/10.1007/s13127-018-0368-4
- Aued, A.W., Smith, F., Quimbayo, J.P., Candido, D.V., Longo, G.O., Ferreira, C.E., Witman, J.D., Floeter, S.R. & Segal, B. (2018) Large-scale patterns of benthic marine communities in the Brazilian Province. PLoS One, 13 (6), e0198452. https://doi.org/10.1371/journal.pone.0198452
- Azevedo, F. & Klautau, M. (2007) Calcareous sponges (Porifera, Calcarea) from Ilha Grande Bay, Brazil, with description of three new species. Zootaxa, 1402 (3), 1–22. https://doi.org/10.11646/zootaxa.1402.1.1
- Azevedo, F., Cóndor-Luján, B., Willenz, P.H., Hajdu, E., Hooker, Y. & Klautau, M. (2015) Integrative taxonomy of calcareous sponges (subclass Calcinea) from the Peruvian coast: morphology, molecules, and biogeography. Zoological Journal of the Linnean Society, 173, 787–817. https://doi.org/10.1111/zoj.12213
- Azevedo, F., Padua, A., Moraes, F., Rossi, A., Muricy, G. & Klautau, M. (2017) Taxonomy and phylogeny of calcareous sponges (Porifera: Calcarea: Calcinea) from Brazilian mid-shelf and oceanic islands. Zootaxa, 4311 (3), 301–344. https://doi.org/10.11646/zootaxa.4311.3.1
- Bachetarzi, R., Dilmi, S., Uriz, M.J., Vázquez-Louis, M., Deudero, S. & Rebzani-Zahaf, C. (2019) The non-indigenous and invasive species Paraleucilla magna Klautau, Monteiro & Borojević, 2004 (Porifera: Calcarea) in the Algerian coast (Southwestern of Mediterranean Sea). Acta Adriatica, 60, 41–46. https://doi.org/10.32582/aa.60.1.3
- Bensari, B., Bahbah, L., Lounaouci, A., Fahci, S.E., Bouda, A. & Bachari, N.E.I. (2020) First records of non-indigenous species in port of Arzew (Algeria: South Western Mediterranean). Mediterranean Marine Science, 21, 393–399. https://doi.org/10.12681/mms.21927
- Bertolino, M., Longo, C., Marra, M.V., Corriero, G. & Pansini, M. (2014) Paraleucilla magna Klautau et al., 2004 (Porifera, Calcarea), specie aliena in continua espansione nel mar mediterraneo/Paraleucilla magna Klautau et al., 2004 (Porifera, Calcarea), an alien species extending its range in the mediterranean sea. Biologia marina mediterranea, 21 (1), 109–110.
- Bidder, G.P. (1898) The Skeleton and Classification of Calcareous Sponges. Proceedings of the Royal Society, 64, 61–76. https://doi.org/10.1098/rspl.1898.0070
- Borojević, R. (1967) Spongiaires d’Afrique du Sud. (2) Calcarea. Transactions of the Royal Society of South Africa, 37 (3), 183–226. https://doi.org/10.1080/00359196709519066
- Borojević, R. (1971) Éponges Calcaires de la Côte Sud-Est du Brésil, Épibiontes sur Laminaria Braziliensis et Sargassum cymosum. Revista Brazileira de Biologia, 31 (4), 525–530.
- Borojević, R. & Boury-Esnault, N. (1987) Revision of the genus Leucilla Haeckel, 1872 with a redescription of the type species – Leucilla amorpha Haeckel, 1872. In: Jones, W.C. (Eds.), European contributions to the taxonomy of sponges. Publications of the Sherkin Island Marine Station, Sherkin Island, pp. 29–14
- Borojević, R., Boury-Esnault, N., Manuel, M. & Vacelet, J. (2002) Order Leucosolenida Hartman, 1958. In: Hooper, J.N.A. & Van Soest, R.W.M. (Eds.), Systema Porifera: A Guide to the Classification of Sponges. Vol. 2. Kluwer Academic/Plenum Publishers, New York, New York, pp. 1157–1184. https://doi.org/10.1007/978-1-4615-0747-5_120
- Borojević, R. & Peixinho, S. (1976) Éponges calcaires du nord-nord-est du Brésil. Bulletin du Muséum National d’histoire Naturelle de Paris, 3 (402), 987–1036. https://doi.org/10.5962/p.281415
- Bouzon, J.L., Brandini, F.P. & Rocha, R.M. (2012) Biodiversity of sessile fauna on rocky shores of coastal islands in Santa Catarina, Southern Brazil. Marine Science, 2 (5), 39–47. https://doi.org/10.5923/j.ms.20120205.01
- Bowerbank, J.S. (1862) On the anatomy and physiology of the Spongiadae. Part III. On the generic characters, the specific characters, and on the method of examination. Philosophical Transactions of the Royal Society, 152 (2), 1087–1135. https://doi.org/10.1098/rstl.1862.0044
- Bowerbank, J.S. (1864) A Monograph of the British Spongiadae. Vol. 1. Ray Society, London, 290 pp. https://doi.org/10.5962/bhl.title.56119
- Bowerbank, J.S. (1873) Contributions to a General History of the Spongiadae. Part IV. Proceedings of the Zoological Society of London, 1873, 319–333.
- Bumbeer, J., Cattani, A.P., Binneck Chierigatti, N. & da Rocha, R.M. (2016) Biodiversity of benthic macroinvertebrates on hard substrates in the Currais Marine Protected Area, in southern Brazil. Biota Neotropica, 16 (4), e20160246. https://doi.org/10.1590/1676-0611-BN-2016-0246
- Burton, M. (1956) The Sponges of West Africa. Atlantide Report: Scientific Results of the Danish Expedition to the Coasts of Tropical West Africa, 1945–1946, 4, 111–147.
- Burton, M. (1963) A Revision of the Classification of the Calcareous Sponges: with a catalogue of the specimens in the British Museum (Natural History). British Museum (Natural History), London, 693 pp.
- Canning-Clode, J., Fofonoff, P., McCann, L., Carlton, J. & Ruiz, G. (2013) Marine invasions on a subtropical island: fouling studies and new records in a recent marina on Madeira Island (Eastern Atlantic Ocean). Aquatic Invasions, 8 (3), 261–270. https://doi.org/10.3391/ai.2013.8.3.02
- Carter, H.J. (1886) Descriptions of sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History, 18 (5), 34–55. https://doi.org/10.1080/00222938509459335
- Carter, H.J. (1890) Porifera. In: Ridley, H.N. (Eds.), Notes on the Zoology of Fernando do Noronha. Journal of the Linnean Society, 20, pp. 564–569.
- Cavalcanti, F.F., Bastos, N. & Lanna, E. (2015) Two new species of the genus Vosmaeropsis Dendy, 1892 (Porifera, Calcarea), with comments on the distribution of V. sericata (Ridley, 1881) along the Southwestern Atlantic Ocean. Zootaxa, 3956 (4), 476–490. https://doi.org/10.11646/zootaxa.3956.4.2
- Cavalcanti, F.F., Chagas, C. & Fonseca, E.S.M. (2019) Taxonomic redescription of Leucandrilla wasinensis (Jenkin, 1908) and L. intermedia (Row, 1909) (Porifera, Calcarea): a first step to improve current knowledge on the typical features of the genus. Zootaxa, 4568 (1), 40–50. https://doi.org/10.11646/zootaxa.4568.1.2
- Cavalcanti, F.F., Menegola, C. & Lanna, E. (2014) Three new species of the genus Paraleucilla Dendy, 1892 (Porifera, Calcarea) from the coast of Bahia State, Northeastern Brazil. Zootaxa, 3764 (5), 537–554. https://doi.org/10.11646/zootaxa.3764.5.3
- Cavalcanti, F.F., Padua, A., Cunha, H., Halasz, M., Nikolić, V., Barreto, S. & Klautau, M. (2020) Population differentiation supports multiple human-mediated introductions of the transatlantic exotic sponge Paraleucilla magna (Porifera, Calcarea). Hydrobiologia, 847, 3571–3590. https://doi.org/10.1007/s10750-020-04368-x
- Cavalcanti, F.F., Rapp, H.T. & Klautau, M. (2013) Taxonomic revision of Leucascus Dendy, 1892 (Porifera: Calcarea) with revalidation of Ascoleucetta Dendy & Frederick, 1924 and description of three new species. Zootaxa, 3619 (3), 275–314. https://doi.org/10.11646/zootaxa.3619.3.3
- Chagas, C. (2021) Esponjas calcareas da família Amphoriscidae Dendy, 1893: revisões taxonômicas, novas espécies e a ocorrência de pentactinas. Programa de Pós-Graduação em Biodiversidade e Evolução. Universidade Federal da Bahia, Salvador, 257 pp.
- Chagas, C. & Cavalcanti, F.F. (2017) Taxonomy of Calcareous Sponges (Porifera, Calcarea) Sampled on Artificial Substrates of a Recreational Marina in the Tropical Northeastern Brazilian Coast. Zootaxa, 4363 (2), 203–224. https://doi.org/10.11646/zootaxa.4363.2.2
- Chagas, C. & Cavalcanti, F.F. (2021) Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea), with description of A. decennis sp. nov. Zootaxa, 5061 (1), 39–68. https://doi.org/10.11646/zootaxa.5061.1.2
- Chebaane, S., Sempere-Valverde, J., Dorai, S., Kacem, A. & Sghaier, Y.R. (2019) A preliminar inventory of alien and cryptogenic species in Monastir Bay, Tunisia: spatial distribution, introduction trends and pathways. Mediterranean Marine Science, 20, 616–626. https://doi.org/10.12681/mms.20229
- Chombard, C., Boury-Esnault, N. & Tillier, S. (1998) Reassessment of homology of morphological characters in tetractinellid sponges based on molecular data. Systematic Biology, 47 (3), 351–366. https://doi.org/10.1080/106351598260761
- Cóndor-Luján, B., Azevedo, F., Hajdu, E., Hooker, Y., Willenz, P. & Klautau, M. (2019) Tropical Eastern Pacific Amphoriscidae Dendy, 1892 (Porifera: Calcarea: Calcaronea: Leucosolenida) from the Peruvian coast. Marine Biodiversity, 49 (4), 1813–1830. https://doi.org/10.1007/s12526-019-00946-y
- Cóndor-Luján, B., Leocorny, P., Padua, A., Azevedo, F., Seixas, V.C., Hooker, Y., Hajdu, E., Willenz, P., Pérez, T. & Klautau, M. (2021) Evolutionary history of the calcareous sponge Clathrina aurea: genetic connectivity in the Western Atlantic and intriguing occurrence in the Eastern Pacific. Marine Biology, 168 (127), 1–23. https://doi.org/10.1007/s00227-021-03934-8
- Cóndor-Luján, B., Louzada, T.S., Hajdu, E. & Klautau, M. (2018) Morphological and molecular taxonomy of calcareous sponges (Porifera: Calcarea) from Curaçao, Caribbean Sea. Zoological Journal of the Linnean Society, 183 (3), 459–525. https://doi.org/10.1093/zoolinnean/zlx082
- Cordeiro, C.A.M.M., Floeter, S.R. & Ferreira, C.E.L. (2024) Biogeographic Patterns of Brazilian Rocky Reef Fauna. In: Coutinho, R. & Christofoletti, R.A. (Eds.), Brazilian Rocky Shores. Brazilian Marine Biodiversity. Springer, Cham, pp. 15–40. https://doi.org/10.1007/978-3-031-67206-4_2
- Cord, I., Nunes, L.T., Barroso, C.X., Freire, A.S., Gadig, O.B.F., Gomes, P.B., Gurgel, C.F.D., Lindner, A., Mantelatto, F.L., Targino, A.K.G. & Floeter, S.R. (2022) Brazilian marine biogeography: a multi-taxa approach for outlining sectorization. Marine Biology, 169 (61), 1–13. https://doi.org/10.1007/s00227-022-04045-8
- Custódio, M.R. & Hajdu, E. (2011) Checklist de Porifera do Estado de São Paulo, Brazil. Biota Neotropica, 11 (1a), 1–17. https://doi.org/10.1590/S1676-06032011000500015
- Cvitković, I., Despalatović, M., Grubelić, I., Nikolić, V., Pleše, B. & Žuljević, A. (2013) Occurrence of Paraleucilla magna (Porifera: Calcarea) in the Eastern Adriatic Sea. Acta Adriatica, 54 (1), 93–99.
- Dailianis, T., Akyol, O., Babali, N., Bariche, M., Crocetta, F., Gerovasileiou, V., Chanem, R., Gokoglu, M., Hasiotis, T., Izquierdo-Munoz, A., Julian, D., Katsanevakis, S., Lipez, L., Mancini, E., Mytilineou, C., Ounifi, K.B.A., Ozgül, A., Ragkousis, M., Rubio-Portillo, E., Servello, G., Sini, M., Stamouli, C., Sterioti, A., Teker, S., Tiralongo, F. & Trkov, D. (2016) New mediterranean biodiversity records (July 2016). Mediterranean Marine Science, 18 (2), 608–626. https://doi.org/10.12681/mms.1734
- de Laubenfels, M.W. (1936) A discussion of the sponge fauna of the dry Tortugas in particular and the West Indies in general, with material for a revision of the families and orders of the Porifera. Tortugas Laboratory Occasional Papers, 467, 1–225.
- De Voogd, N.J., Alvarez, B., Boury-Esnault, N., Cárdenas, P., Díaz, M.-C., Dohrmann, M., Downey, R., Goodwin, C., Hajdu, E., Hooper, J.N.A., Kelly, M., Klautau, M., Lim, S.C., Manconi, R., Morrow, C., Pinheiro, U., Pisera, A.B., Ríos, P., Rützler, K., Schönberg, C., Turner, T., Vacelet, J., van Soest, R.W.M. & Xavier, J. (2025) World Porifera Database. Available from: https://www.marinespecies.org/porifera (accessed 30 March 2025)
- Dendy, A. (1892) Synopsis of the Australian Calcarea Heterocoela, with a proposed classification of the group and descriptions of some new genera and species. Proceedings of the Royal Society of Victoria, 5, 69–116.
- Dendy, A. (1893) Studies on the Comparative Anatomy of Sponges. V. Observations on the Structure and Classification of the Calcarea Heterocoela. Quarterly Journal of Microscopical Science, 35, 159–257. https://doi.org/10.1242/jcs.s2-35.138.159
- Dendy, A. (1913) Report on the calcareous sponges collected by H.M.S. “Sealark” in the Indian Ocean. Transactions of the Linnean Society of London, Zoology, 16 (1), 1–29. https://doi.org/10.1111/j.1096-3642.1914.tb00121.x
- Dendy, A. (1916) Report on the Calcareous Sponges collected by Mr. James Hornell at Okhamandal in Kattiawar in 1905–6. Report to the Government of Baroda on the Marine Zoology of Okhamandal in Kattiawar, 2, 79–91.
- Dendy, A. & Row, R.W.H. (1913) The classification and phylogeny of the calcareous sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London, 83 (3), 704–813. https://doi.org/10.1111/j.1469-7998.1913.tb06152.x
- Dohrmann, M., Voigt, O., Erpenbeck, D. & Wörheide, G. (2006) Non-monophyly of most supraspecific taxa of calcareous sponges (Porifera, Calcarea) revealed by increased taxon sampling and partitioned Bayesian analysis of ribosomal DNA. Molecular Phylogenetics and Evolution, 40, 830–843. https://doi.org/10.1016/j.ympev.2006.04.016
- Dunnington, D. (2023) ggspatial: Spatial Data Framework for ggplot2. R package 2023. Version 1.1.9. Available from: https://cran.r-project.org/package=ggspatial (accessed 18 July 2025)
- Ellis, J. & Solander, D. (1786) The Natural History of many curious and uncommon Zoophytes, collected from various parts of the Globe. Systematically arranged and described by the late Daniel Solander. 4. Benjamin White & Son, London, 206 pp. https://doi.org/10.5962/bhl.title.2145
- Evcen, A. & Çinar, M.E. (2020) Sponge species from ports of the inner and middle parts of İzmir Bay (Aegean Sea, Eastern Mediterranean). İzmir iç ve orta körfezi limanlarından sünger türleri (Ege Denizi, Doğu Akdeniz). Ege Journal of Fisheries and Aquatic Sciences, 37 (2), 149–155. https://doi.org/10.12714/egejfas.37.2.05
- Fabricius, O. (1780) Fauna Groenlandica. Hafniae et Lipsae, Copenhagen, 452 pp. https://doi.org/10.5962/bhl.title.13489
- Fleming, J. (1828) A history of British animals, exhibiting the descriptive characters and systematical arrangement of the genera and species of quadrupeds, birds, reptiles, fishes, Mollusca, and Radiata of the United Kingdom; including the indigenous, extirpated, and extinct kinds, together with periodical and occasional visitants. Bell & Bradfute, Edinburgh, 565 pp. https://doi.org/10.5962/bhl.title.12859
- Fonseca, E.S.M., Cóndor-Luján, B. & Cavalcanti, F.F. (2023) Calcareous sponges (Porifera, Calcarea) from the Northeastern Brazilian coast: integrative taxonomy and support for the creation of a marine park. Marine Biodiversity, 53, 67. https://doi.org/10.1007/s12526-023-01364-x
- Fontana, T., Cóndor-Luján, B., Azevedo, F., Pérez, T. & Klautau, M. (2018) Diversity and distribution of Calcareous sponges (subclass Calcinea) from Martinique. Zootaxa, 4410 (2), 331–369. https://doi.org/10.11646/zootaxa.4410.2.5
- Fundação para a Conservação e a Produção Florestal do Estado de São Paulo (2022) Plano de Manejo da Área de Proteção Ambiental Marinha do Litoral Norte, São Paulo. Available from: https://sigam.ambiente.sp.gov.br/sigam3/Repositorio/511/Documentos/APAM_LN/APAMLN_Plano_de_manejo_CTBio.pdf (accessed 30 March 2025)
- Gerovasileiou, V., Akel, E.l., Akyol, O., Alongi, G., Azevedo, F., Babali, N., Bakiu, R., Bariche, M., Bennoui, A., Castriota, L., Chintiroglou, C., Crocetta, F., Deidun, A., Galinou-Mitsoudi, S., Giovos, I., Gokoglu, M., Golemaj, A., Hadjioannou, L., Hartingerova, J. & Zenetos, A. (2017) New Mediterranean Biodiversity Records (July 2017). Mediterranean Marine Science, 18, 355–384. https://doi.org/10.12681/mms.13771
- Gray, J.E. (1867) Notes on the Arrangement of Sponges, with the Descriptions of some New Genera. Proceedings of the Zoological Society of London, 1867 (2), 492–558.
- Guardiola, M., Frotscher, J. & Uriz, M.J. (2012) Genetic structure and differentiation at a short-time scale of the introduced calcarean sponge Paraleucilla magna to the western Mediterranean. Hydrobiologia, 687, 71–84. https://doi.org/10.1007/s10750-011-0948-1
- Guardiola, M., Frotscher, J. & Uriz, M.J. (2016) High genetic diversity, phenotypic plasticity, and invasive potential of a recently introduced calcareous sponge, fast spreading across the Atlanto-Mediterranean basin. Marine Biology, 163, 123. https://doi.org/10.1007/s00227-016-2862-6
- Haeckel, E. (1870) Prodromus eines Systems der Kalkschwämme. Jenaische Zeitschrift für Medicin und Naturwissenschaft, 5, 236–254.
- Haeckel, E. (1872) Die Kalkschwämme. Eine monographie, Vols 1–3. G. Reimer, Berlin, 512 + 440 + 260 pp. https://doi.org/10.5962/bhl.title.11323
- Hajdu, E., Carvalho, M.S. & Lôbo-Hajdu, G. (2011) Porifera. In: Amaral, A.C.Z., Nallin, S.A.H. (Eds.), Biodiversidade e ecossistemas bentônicos marinhos do litoral norte de São Paulo, Sudeste do Brasil. UNICAMP/IB, Campinas, pp. 50–59. [E-book] Available from: https://www2.unifap.br/alexandresantiago/files/2012/12/E-BOOKBiota_1.pdf (accessed 30 March 2025)
- Hajdu, E., Santos, C.P., Lopes, D.A., Oliveira, M.V., Moreira, M.C.F., Carvalho, M.S. & Klautau, M. (2004) Filo Porifera. In: Amaral, A.C.Z., Rossi-Wongtschowski, C.L.D.B. (Eds.), Biodiversidade bentônica da região sudeste-sul do Brazil - Plataforma Externa e Talude Superior. Série Documentos Revizee - Score Sul. Instituto Oceanográfico da Universidade de São Paulo, São Paulo, pp. 49–56.
- Hartman, W.D. (1958) Re-examination of Bidder’s classification of the Calcarea. Systematic Zoology, 7 (3), 97–110. https://doi.org/10.2307/2411971
- Hoang, D.T., Chernomor, O., Von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution, 35 (2), 518–522. https://doi.org/10.1093/molbev/msx281
- Hoff, N.T. (2015) Integridade biótica dos ecossistemas na região do Arquipélago dos Alcatrazes, São Sebastião - SP. São Paulo State University, São Paulo, 170 pp.
- Hoff, N.T., Takase, L.S., Siegle, E. & Dias, J.F. (2022) Mapeamento da sensibilidade ambiental a derrames de óleo do arquipélago dos Alcatrazes (São Paulo, Brazil). Journal of Integrated Coastal Zone Management, 22 (1), 43–80. https://doi.org/10.5894/rgci-n452
- ICMBIO (2017) Plano de Manejo da Estação Ecológica Tupinambás e Refúgio de Vida Silvestre do Arquipélago de Alcatrazes. Available from: https://www.gov.br/icmbio/pt-br/assuntos/biodiversidade/unidade-de-conservacao/unidades-de-biomas/marinho/lista-de-ucs/refugio-de-alcatrazes/arquivos/plano_de_manejo_esec_tupinambas_revisarquipelogoalcatrazes_vol1.pdf (accessed 30 March 2025)
- ICMBIO (2020) Plano de Uso Público do Refúgio de Vida Silvestre do Arquipélago de Alcatrazes. Available from: https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/planos/plano_de_uso_publico_do_revis_do_arquipelago_de_alcatrazes.pdf (accessed 30 March 2025)
- Ignacio, B.L., Julio, L.M., Junqueira, A.O.R. & Ferreira-Silva, M.A.G. (2010) Bioinvasion in a Brazilian bay: filling gaps in the knowledge of Southwestern Atlantic biota. PLoS One, 5 (9), e13065. https://doi.org/10.1371/journal.pone.0013065
- Jenkin, C.F. (1908) The Marine Fauna of Zanzibar and British East Africa, from Collections made by Cyril Crossland M.A., in the Years 1901 & 1902, The Calcareous Sponges. Proceedings of the Zoological Society of London, 78 (3), 434–456. https://doi.org/10.1111/j.1469-7998.1908.tb07387.x
- Junqueira, A.O.R., Tavares, M.D.S., Schaeffer-Noveli, Y., Radashevsky, V.I., Cirelli, J.O., Julio, L.M., Romagnoli, F.C., Santos, K.C. & Ferreira-Silva, M.A.G. (2009) Capítulo 6. Zoobentos. In: Lopes, R.M., Coradin, L., Pombo, V.B & Cunha, D.R. (Eds.), Informe sobre as espécies invasoras marinhas no Brasil. Ministério do Meio Ambiente, Brasília, pp. 145–371.
- Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods, 14 (6), 587–589. https://doi.org/10.1038/nmeth.4285
- Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software v.7: improvements in performance and usability. Molecular Biology and Evolution, 30 (4), 772–780. https://doi.org/10.1093/molbev/mst010
- Katoh, K. & Toh, H. (2008) Improved accuracy of multiple ncRNA alignment by incorporating structural information into a MAFFT-based framework. BioMedCentral Bioinformatics, 9 (212), 1–13. https://doi.org/10.1186/1471-2105-9-212
- Katsanevakis, S., Poursanidis, D., Hoffman, R., Rizgalla, J., Rothman, S., Levitt-Barmats, Y., Hadjioannou, L., Trkov, D., Garmendia, J. Rizzo, M., Bartolo, A., Bariche, M., Tomas, F., Kleitou, P., Schembri, P., Kletou, D., Tiralongo, F., Pergent-Martini, C., Pergent, G. & Zenetos, A. (2020) Unpublished Mediterranean records of marine alien and cryptogenic species. BioInvasions Records, 9, 165–182. https://doi.org/10.3391/bir.2020.9.2.01
- Klautau, M. & Borojević, R. (2001) Sponges of the genus Clathrina Gray, 1867 from Arraial do Cabo, Brazil. Zoosystema, 23 (3), 395–410.
- Klautau, M., Cavalcanti, F.F. & Borojević, R. (2017) The new sponge species Amphoriscus pedunculatus (Porifera, Calcarea). Zootaxa, 4341 (1), 105–112. https://doi.org/10.11646/zootaxa.4341.1.9
- Klautau, M., Cóndor-Luján, B., Azevedo, F., Leocorny, P., Rocha Brandão, F.D.A. & Cavalcanti, F.F. (2020) Heteropia glomerosa (Bowerbank, 1873) (Porifera, Calcarea, Calcaronea), a new alien species in the Atlantic. Systematics and Biodiversity, 18 (4), 362–376. https://doi.org/10.1080/14772000.2020.1758826
- Klautau, M., Imešek, M., Azevedo, F., Pleše, B., Nikolić, V. & Ćetković, H. (2016) Adriatic calcarean sponges (Porifera, Calcarea), with the description of six new species and a richness analysis. European Journal of Taxonomy, 178, 1–52. https://doi.org/10.5852/ejt.2016.178
- Klautau, M., Lopes, M.V., Tavares, G. & Pérez, T. (2021) Integrative taxonomy of calcareous sponges (Porifera: Calcarea) from Réunion Island, Indian Ocean. Zoological Journal of the Linnean Society, 194, 671–725. https://doi.org/10.1093/zoolinnean/zlab014
- Klautau, M., Monteiro, L. & Borojević, R. (2004) First Occurrence of the Genus Paraleucilla (Calcarea, Porifera) in the Atlantic Ocean: P. magna sp. nov. Zootaxa, 710 (1), 1–8. https://doi.org/10.11646/zootaxa.710.1.1
- Klautau, M., Solé-Cava, A.M. & Borojević, R. (1994) Biochemical systematics of sibling sympatric species of Clathrina (Porifera: Calcarea). Biochemical Systematics and Ecology, 22 (4), 367–375. https://doi.org/10.1016/0305-1978(94)90027-2
- Klautau, M. & Valentine, C. (2003) Revision of the genus Clathrina (Porifera, Calcarea). Zoological Journal of the Linnean Society, 139 (1), 1–62. https://doi.org/10.1046/j.0024-4082.2003.00063.x
- Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33 (7), 1870–1874. https://doi.org/10.1093/molbev/msw054
- Lanna, E., Cavalcanti, F.F., Cardoso, L., Muricy, G. & Klautau, M. (2009) Taxonomy of calcareous sponges (Porifera, Calcarea) from Potiguar Basin, NE Brazil. Zootaxa, 1973 (1), 1–27. https://doi.org/10.11646/zootaxa.1973.1.1
- Lanna, E., Rattis, L. & Cavalcanti, F.F. (2017) The presence of the diagnostic character of the genus Paraleucilla (Amphoriscidae, Calcarea, Porifera) may depend on the volume and body wall thickness of the sponges. Invertebrate Biology, 136 (3), 321–329. https://doi.org/10.1111/ivb.12185
- Lanna, E., Rossi, A.L., Cavalcanti, F.F., Hajdu, E. & Klautau, M. (2007) Calcareous sponges from São Paulo state, Brazil (Porifera: Calcarea: Calcinea) with the description of two new species. Journal of the Marine Biological Association of the United Kingdom, 87 (6), 1553–1561. https://doi.org/10.1017/S0025315407056871
- Lavrov, A., Ekimova, I., Schepetov, D., Koinova, A. & Ereskovsky, A. (2024) The complex case of the calcareous sponge Leucosolenia complicata (Porifera: Calcarea): hidden diversity in Boreal and Arctic regions with description of a new species. Zoological Journal of the Linnean Society, 200 (4), 876–914. https://doi.org/10.1093/zoolinnean/zlad104
- Lendenfeld, R.V. (1885) A monograph of the Australian sponges. Part III. Preliminary description and classification of the Australian Calcispongiae. Proceedings of the Linnean Society of New South Wales, 9, 1083–1150.
- Lerner, C. & Hajdu, E. (2002) Two new Mycale (Naviculina) Gray (Mycalidae, Poecilosclerida, Demospongiae) from the Paulista Biogeographic Province (Southwestern Atlantic). Revista Brazileira de Zoologia, 19 (1), 109–122. https://doi.org/10.1590/S0101-81752002000100009
- Lima, M.V., Brito, D.D. & Milanelli, J.C.C. (2008) Mapeamento da sensibilidade ambiental a derrames de óleo em Ilhabela, São Paulo. Revista Brazileira de Cartografia, 60 (2), 145–154. https://doi.org/10.14393/rbcv60n2-44877
- Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. [The system of nature through the three kingdoms of nature, according to classes, orders, genera, species, with characters, differences, synonyms, places.]. Impensis Direct Laurentii Salvii, Holmiae [Stockholm], 824 pp. https://doi.org/10.5962/bhl.title.542
- Longo, C. (2009) Leucosolenia microspinata sp. nov. (Porifera: Calcarea), a new species from the Mediterranean Sea. Journal of the Marine Biological Association of the United Kingdom, 89 (8), 1663–1669. https://doi.org/10.1017/S0025315409990440
- Longo, C., Liaci, L., Manuel, M. & Correiro, G. (2004) Note sui poriferi del Mar Grande e del Mar Piccolo di Taranto (Mar Ionio). Biologia Marina Mediterranea, 11, 440–443.
- Longo, C., Mastrototaro, F. & Corriero, G. (2007) Occurrence of Paraleucilla magna (Porifera: Calcarea) in the Mediterranean Sea. Journal of the Biological Association of the United Kingdom, 87, 1749–1755. https://doi.org/10.1017/S0025315407057748
- Longo, C., Pontassuglia, C., Corriero, G. & Gaino, E. (2012) Life-cycle traits of Paraleucilla magna, a calcareous sponge invasive in a coastal mediterranean basin. PLoS One, 7, e42392. https://doi.org/10.1371/journal.pone.0042392
- Lopes, M.V., Padua, A., Azevedo, F. & Klautau, M. (2025) Integrative taxonomy of Calcarea (Porifera) from Espírito Santo, Eastern Brazil. Zootaxa, 5618 (2), 151–205. https://doi.org/10.11646/zootaxa.5618.2.1
- Lopes, M.V., Padua, A., Cóndor-Luján, B. & Klautau, M. (2018) Calcareous sponges (Porifera, Calcarea) from Florida: New species, new records and biogeographical affinities. Zootaxa, 4526 (2), 127–150. https://doi.org/10.11646/zootaxa.4526.2.2
- Łukowiak, M., Van Soest, R., Klautau, M., Pérez, T., Pisera, A. & Tabachnick, K. (2022) The terminology of sponge spicules. Journal of Morphology, 283 (12), 1517–1545. https://doi.org/10.1002/jmor.21520
- Mačić, V. & Petović, S. (2016) New data on the distribution of the alien sponge Paraleucilla magna Klautau, Monteiro & Borojević, 2004 in the Adriatic Sea. Studia Marina, 29 (1), 63–68.
- Marra, M.V., Bertolino, M., Pansini, M., Giacobbe, S., Manconi, R. & Pronzato, R. (2016) Long-term turnover of the sponge fauna in Faro Lake (north-East Sicily, Mediterranean Sea). Italian Journal of Zoology, 83, 579–588. https://doi.org/10.1080/11250003.2016.1251981
- Massicotte, P. & South, A. (2023) rnaturalearth: World Map Data from Natural Earth. R package 2023. Version 1.0.1. Available from: https://cran.r-project.org/package=rnaturalearth (accessed 18 July 2025)
- Mattedi, A. & Cavalcanti, F.F. (2024) Sycetta papillata sp. nov. (Porifera, Calcarea): the first record of Sycetta to the Southwestern Atlantic Ocean. Zootaxa, 5481 (3), 353–362. https://doi.org/10.11646/zootaxa.5481.3.4
- Mello-Leitão, A., Pêgo, A.F. & Lopes, W.M. (1961) Poríferos assinalados no Brazil. Avulsos do Centro de Estudos Zoológicos, 10, 1–29.
- Migotto, A.E., Amaral, A.C.Z. & Rocha, C.E.F. (2011) Litoral Norte de São Paulo. In: Amaral, A.C.Z. & Nallin, S.A.H. (Eds.), Biodiversidade e ecossistemas bentônicos marinhos do Litoral Norte de São Paulo, Sudeste do Brazil. UNICAMP/IB, Campinas, pp. 23–26. [E-book] Available from: https://www2.unifap.br/alexandresantiago/files/2012/12/E-BOOKBiota_1.pdf (accessed 30 March 2025)
- Minchin, E.A. (1900) Chapter III. Sponges. In: Lankester, E.R. (Ed.), A Treatise on Zoology. Part II. The Porifera and Coelenterata. 2. Adam & Charles Black, London, pp. 1–178.
- Minchin, E.A. (1904) The characters and synonymy of the British species of sponges of the genus Leucosolenia. Proceedings of the Zoological Society of London, 74, 349–96. https://doi.org/10.1111/j.1469-7998.1905.tb08345.x
- Minh, B.Q., Schmidt, H.A., Chernomor, O., Schrempf, D., Woodhams, M.D., Von Haeseler, A. & Lanfear, R. (2020) IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution, 37 (5), 1530–1534. https://doi.org/10.1093/molbev/msaa015
- Miranda, T.P., Genzano, G.N. & Marques, A.C. (2015) Areas of endemism in the Southwestern Atlantic Ocean based on the distribution of benthic hydroids (Cnidaria: Hydrozoa). Zootaxa, 4033 (4), 484–506. https://doi.org/10.11646/zootaxa.4033.4.2
- Montagu, G. (1814 [1818]) An essay on sponges, with descriptions of all the species that have been discovered on the coast of Great Britain. Memoirs of the Wernerian Natural History Society, 2, 67–122.
- Monteiro, L.C. & Muricy, G. (2004) Patterns of sponge distribution in Cagarras Arquipelago, Rio de Janeiro, Brazil. Journal of the Marine Biological Association of the United Kingdom, 84 (4), 681–687. https://doi.org/10.1017/S0025315404009750h
- Moraes, F.C., Ventura, M., Klautau, M., Hajdu, E. & Muricy, G. (2006) Biodiversidade de esponjas das ilhas oceânicas Brazileiras. In: Alves, R.J.V., Castro, J.W.A. (Eds.), Ilhas Oceânicas Brazileiras – da pesquisa ao manejo. Ministério do Meio Ambiente, Secretaria de Biodiversidade e Florestas, Brasília, pp. 147–178.
- Moreno, T.R., de Faria, S.B. & Rocha, R.M. (2014) Biogeography of Atlantic and Mediterranean ascidians. Marine Biology, 161, 2023–2033. https://doi.org/10.1007/s00227-014-2483-x
- Mothes de Moraes, B. (1985) Sponges collected by the Oxford diving expedition to the Cabo Frio upwelling area (Rio de Janeiro, Brazil). Studies on Neotropical Fauna and Environment, 20 (4), 227–237. https://doi.org/10.1080/01650528509360694
- Mothes, B. & Lerner, C.B. (1994) Esponjas marinhas do infralitoral de Bombinhas (Santa Catarina, Brasil) com descrição de três espécies novas (Porifera: Calcarea e Demospongiae). Biociências, 2 (1), 47–62.
- Muricy, G., Esteves, E.L., Moraes, F.C., Santos, J.P., Silva, S.M., Almeida, E.V.R., Klautau, M. & Lanna, E. (2008) Biodiversidade Marinha da Bacia Potiguar: Porifera. Museu Nacional, Série Livros 29. Museu Nacional, Rio de Janeiro, 156 pp.
- Muricy, G. & Hajdu, E. (2006) Porifera Brasilis: Guia de Identificação das Esponjas mais comuns do Sudeste do Brazil. Série Livros 17. Museu Nacional, Rio de Janeiro, 104 pp.
- Muricy, G., Hadju, E., Custódio, M., Klautau, M., Russo, C.A.M. & Peixinho, S. (1991) Sponge distribution at Arraial do Cabo, SE Brazil. In: Magoon, O.T., Converse, H., Tippie, V., Tobin, L.T. & Clark, D. (Eds.), Proceedings of the VII Symposium on Coastal and Oceanic Management: Coastal Zone`91. ASCE Publications, Long Beach, pp. 1183–1196.
- Muricy, G., Lopes, D.A., Hajdu, E., Carvalho, M.S., Moraes, F.C., Klautau, M., Menegola, C. & Pinheiro, U. (2011) Catalogue of Brazilian Porifera. Série Livros 46. Museu Nacional, Rio de Janeiro, 300 pp.
- Muricy, G. & Silva, O.C. (1999) Esponjas marinhas do Estado do Rio de Janeiro: um recurso renovável inexplorado. In: Silva, S.H.G. & Lavrado, H.P. (Eds.), Ecologia dos Ambientes Costeiros do Estado do Rio de Janeiro. Série Oecologia Braziliensis. Vol. VII. PPGE-UFRJ, Rio de Janeiro, pp. 155–178. https://doi.org/10.4257/oeco.1999.0701.07
- Nassar, C.A.G. & Silva, S.H.G. (1999) Comunidade incrustante em quatro profundidades na Ilha Guaíba – Rio de Janeiro, Brazil. Oecologia Braziliensis, 7 (1), 195–211. https://doi.org/10.4257/oeco.1999.0701.09
- Nguyen, L.T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution, 32, 268–274. https://doi.org/10.1093/molbev/msu300
- Oricchio, F.T., Marques, A.C., Hajdu, E., Pitombo, F.B., Azevedo, F., Passos, F.D., Vieira, L.M., Stampar, S.N., Rocha, R.M. & Dias, G.M. (2019) Exotic species dominate marinas between the two most populated regions in the southwestern Atlantic Ocean. Marine Pollution Bulletin, 146, 884–892. https://doi.org/10.1016/j.marpolbul.2019.07.013
- Padua, A., Cunha, H.A. & Klautau, M. (2018) Gene flow and differentiation in a native calcareous sponge (Porifera) with unknown dispersal phase: Population structure of C. aurea. Marine Biodiversity, 48 (4), 2125–2135. https://doi.org/10.1007/s12526-017-0742-z
- Palacio, F.J. (1982) Revisión zoogeográfica marina del sur del Brazil. Boletim do Instituto Oceanográfico da Universidade de São Paulo, 31 (1), 69–92. https://doi.org/10.1590/S0373-55241982000100006
- Pebesma, E. & Bivand, R. (2023) Spatial Data Science: With Applications in R. Chapman and Hall/CRC, New York, New York, 314 pp. https://doi.org/10.1201/9780429459016
- Pinheiro, H.T., Bender, M.G., Longo, G.O., Maxwell, M.F., Rocha, L.A. & Floeter, S.R. (2024) Origins, Biogeography and Macroecology of the Southwestern Atlantic Reef Biodiversity. In: Kikuchi, R.K.P., Leão, Z.M.A.N., de Araújo, M.E. & Lotufo, T.M.C. (Eds.), Brazilian Coral Reefs: A Multidisciplinary Approach. Springer, Cham, pp. 59–80. https://doi.org/10.1007/978-3-031-59152-5_3
- Pinheiro, U.S., Sandes, J., Annunziata, B.B., Lopes, M.V. & Muricy, G. (2025) Porifera in Catálogo Taxonômico da Fauna do Brazil. Available from: http://fauna.jbrj.gov.br/fauna/faunadoBrazil/6 (accessed 30 March 2025)
- Poléjaeff, N. (1883) Report on the Calcarea dredged by H.M.S.‘Challenger’, during the years 1873–1876. Report on the Scientific Results of the Voyage of H.M.S. ‘Challenger’, 1873–1876. Zoology, 8 (2), 1–76.
- R Core Team (2023) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Viena. Available from: https://www.R-project.org (accessed 18 July 2025)
- Rapp, H.T., Göcke, C., Tendal, O.S. & Janussen, D. (2013) Two new species of calcareous sponges (Porifera: Calcarea) from the deep Antarctic Eckström Shelf and a revised list of species found in Antarctic waters. Zootaxa, 3692 (1), 149–159. https://doi.org/10.11646/zootaxa.3692.1.9
- Risso, A. (1827) Histoire Naturelle des Principales Productions de l’Europe Méridionale et Particulièrement de Celles des Environs de Nice et des Alpes Maritimes. Tome V. Levrault, Paris, 403 pp. https://doi.org/10.5962/bhl.title.58984
- Rossi, A.L., Farina, M., Borojević, R. & Klautau, M. (2006) Occurrence of five-rayed spicules in a calcareous sponge: Sycon pentactinalis sp. nov. (Porifera: Calcarea). Cahiers de Biologie Marine, 47 (3), 261–270.
- Row, R.W.H. (1909) Reports on the marine biology of the Sudanense Red Sea. XIII. Report on the sponges, collected by Mr. Cyril Crossland in 1904–5. Part I. Calcarea. Journal of the Linnean Society, 31 (206), 182–214. https://doi.org/10.1111/j.1096-3642.1909.tb00983.x
- Santín, A., Wirtz, P., Neves, P. & Ribeiro, C. (2024) Filling gaps in the Webbnesia marine diversity: the Madeiran sponge fauna. Continental Shelf Research, 277, 105230. https://doi.org/10.1016/j.csr.2024.105230
- Sarà, M. (1951) Tre nuove Calcispongie dal Golfo di Napoli. Annuario dell’Instituto e Museo de Zoologia dell’Università di Napoli, 3 (8), 1–16.
- Schmidt, O. (1862) Die Spongien des adriatischen Meeres. Wilhelm Engelmann, Leipzig, 88 pp.
- Schuffner, O. (1877) Beschreibung einiger neuer Kalkschwämme. Jenaische Zeitschrift für Naturwissenschaft, 11, 403–433.
- Sghaier, Y.R., Zakhama-Sraieb, R., Ben, Hmida A. & Charfi, F. (2019) An inventory of nonindigenous species (NIS) inside and outside three tourist marinas from the southern Mediterranean coast. Journal of Black Sea/Mediterranean Environment, 25 (1), 29–48.
- Silva, D.A.M. & Bícego, M.C. (2010) Polycyclic aromatic hydrocarbons and petroleum biomarkers in São Sebastião Channel, Brazil: Assessment of petroleum contamination. Marine Environmental Research, 69 (5), 277–286. https://doi.org/10.1016/j.marenvres.2009.11.007
- Soares, M.O., Lotufo, T.M.C., Vieira, L.M., Salani, S., Hajdu, E., Matthews-Cascon, H., Leão, Z.M.A.N. & Kikuchi, R.K.P. (2017) Brazilian Marine Animal Forests: A New World to Discover in the Southwestern Atlantic. In: Rossi, S., Bramanti, L., Gori, A. & Orejas, C. (Eds.), Marine Animal Forests. Springer, Cham, pp. 73–110. https://doi.org/10.1007/978-3-319-21012-4_51
- Solé-Cava, A.M., Klautau, M., Boury-Esnault, N., Borojević, R. & Thorpe, J.P. (1991) Genetic evidence for cryptic speciation in allopatric populations of two cosmopolitan species of the calcareous sponge genus Clathrina. Marine Biology, 111 (3), 381–386. https://doi.org/10.1007/BF01319410
- Spalding, M.D., Fox, H.E., Allen, G.R., Davidson, N., Ferdaña, Z.A., Finlayson, M., Halpern, B.S., Jorge, M.A., Lombana, A., Lourie, S.A., Martin, K.D., Mcmanus, D., Molnar, J., Recchia, C.A. & Robertson, J. (2007) Marine Ecoregions of the World: A Bioregionalization of Coastal and Shelf Areas. BioScience, 57 (7), 573–583. https://doi.org/10.1641/B570707
- Tamburini, M., Keppel, E., Marchini, A., Repetto, M.F., Ruiz, G.M., Ferrario, J. & Occhipinti-Ambrogi, A. (2021) Monitoring non-indigenous species in port habitats: First application of a standardized North American protocol in the Mediterranean Sea. Frontiers in Marine Science, 8, 700730. https://doi.org/10.3389/fmars.2021.700730
- Tanita, S. (1942) Report on the Calcareous sponges obtained by the Zoological Institute and Museum of Hamburg. Part II. Science Reports of the Tôhoku University, 4 (17), 105–135.
- Targino, A.K.G. & Gomes, P.B. (2020) Distribution of sea anemones in the Southwest Atlantic: biogeographical patterns and environmental drivers. Marine Biodiversity, 50, 80. https://doi.org/10.1007/s12526-020-01099-z
- Tennekes, M. (2018) tmap: Thematic Maps in R. Journal of Statistical Software, 84 (6), 1–39. https://doi.org/10.18637/jss.v084.i06
- Thacker, A.G. (1908) On collections of the Cape Verde Islands fauna made by Cyril Crossland, M.A. The calcareous sponges. Proceedings of the Zoological Society of London, 49, 757–782. https://doi.org/10.1111/j.1469-7998.1908.tb07405.x
- Topaloğlu, B., Evcen, A. & Cinar, M.E. (2016) Sponge fauna in the Sea of Marmara. Turkish Journal of Fisheries and Aquatic Sciences, 16 (1), 51–59. https://doi.org/10.4194/1303-2712-v16_1_06
- Topsent, E. (1891) Voyage de la Goëlette ‘Melita’ aux Canaries et au Sénégal, 1889-1890. Spongiaires. Mémoires de la Société Zoologique de France, 4, 11–15.
- Trifinopoulos, J., Nguyen, L.T., von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research, 44 (1), 232–235. https://doi.org/10.1093/nar/gkw256
- Ulman, A., Ferrario, J., Occhpinti-Ambrogi, A., Arvanitidis, C., Bandi, A., Bertolino, M., Bogi, C., Chatzigeorgiou, G., Çiçek, B.A., Deidun, A., Ramos-Esplá, A., Koçak, C., Lorenti, M., Martinez-Laiz, G., Merlo, G., Princisgh, E., Scribano, G. & Marchini, A. (2017) A massive update of non-indigenous species records in Mediterranean marinas. PeerJ, 5, e3954. https://doi.org/10.7717/peerj.3954
- Urban, F. (1902) Rhabdodermella nuttingi, nov. gen. et nov. spec. Zeitschrift für Wissenschaftliche Zoologie, 71, 268–275.
- Van Soest, R.W. (2017) Sponges of the Guyana Shelf. Zootaxa, 4217 (1), 1–225. https://doi.org/10.11646/zootaxa.4217.1.1
- Van Soest, R.W.M. & De Voogd, N.J. (2018) Calcareous sponges of the Western Indian Ocean and Red Sea. Zootaxa, 4426 (1), 1–160. https://doi.org/10.11646/zootaxa.4426.1.1
- Voigt, O. & Wörheide, G. (2016) A short LSU rRNA fragment as a standard marker for integrative taxonomy in calcareous sponges (Porifera: Calcarea). Organisms, Diversity and Evolution, 16 (1), 53–64. https://doi.org/10.1007/s13127-015-0247-1
- Voigt, O., Wülfing, E. & Wörheide, G. (2012) Molecular phylogenetic evaluation of classification and scenarios of character evolution in calcareous sponges (Porifera, class Calcarea). PLoS One, 7 (3), e33417. https://doi.org/10.1371/journal.pone.0033417
- Wickham, H. (2016) ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag, New York, New York, 213 pp. https://doi.org/10.1007/978-0-387-98141-3
- Zammit, P., Longo, C. & Schembri, P. (2009) Occurrence of Paraleucilla magna Klautau et al., 2004 (Porifera: Calcarea) in Malta. Mediterranean Marine Science, 10, 135–138. https://doi.org/10.12681/mms.114