Search

Describing ontogenetic morphological change is an important part of integrative taxonomy; still, most taxonomic studies are based only on adult characters. Here, we provide illustrations and a morphological description of the sea stars from Bahia, including ontogenetic and intraspecific variation, and identify taxonomic issues. A total of 293 specimens from different localities along the Bahia State coastline and comparative material from other localities were examined. Eighteen species (11 genera, eight families) of Asteroidea were identified; Astropectinidae was the most representative family. All species identified also occur in subtropical Brazilian waters and most species are from shallow water habitats with soft bottoms. Most observed ontogenetic variation was quantitative in nature, such as the increase in the number of spines in the furrow and of spinelets in the paxillae with specimen growth. Genera that require further taxonomic studies are Astropecten and Othilia, whose specimens are commonly misidentified in local studies. One third of the species from Bahia are currently classified as “Vulnerable” in the Brazilian Red List, but baseline data on the population biology of these species are scarce. An illustrated identification key to the 65 Brazilian sea star species is also provided. This taxonomic study will facilitate the identification of specimens occurring along the Brazilian coast and help scientists and policy makers to establish the conservation status of the Brazilian species.

Genus Othilia Gray, 1840 Type species. Othilia echinophora (Lamarck, 1816) (type by subsequent designation by Fisher, 1911). Remarks. Previously demoted as a subgenus of Echinaster by Clark & Tortonese (1986) because of the lack of exclusive morphological characters (see Clark & Downey [1992] for a thorough revision), the re-erection of the genus Othilia is supported by morphological (Fontanella & Hopkins 2003) and molecular evidence (Lopes et al. 2016). These phylogenies have shown that Othilia is a monophyletic group and that the genus Echinaster is more closely related to Henricia than to Othilia. Fontanella & Hopkins (2003) found five synapomorphies supporting the genus Othilia, but unfortunately these were not listed in their paper. A.M. Clark (1987) distinguished Echinaster from Othilia using two diagnostic characters: the arrangement of the dorsolateral plates, which are organized into an irregular reticulum in Echinaster and a regular reticulum in Othilia; and the presence of actinal plates between the inferomarginal and adambulacral plates in Echinaster. However, Lopes et al. (2016) mentioned that, based on the literature, E. callosus Marenzeller, 1895 and E. luzonicus (Gray, 1840) do not have actinal plates. Three species of Othilia have been reported for the Brazilian coast: O. brasiliensis, O. echinophora and O. guyanensis. Species of the genus Othilia are commercially exploited in Rio de Janeiro and Bahia as souvenirs and for the aquarium trade, and they are also used for medicinal (treatment of asthma and alcoholism) and religious purposes (Nobre & Campos-Creasey 2000; Alves et al. 2009; Alves & Dias 2010; Martins et al. 2012 [as Echinaster guyanensis])., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on page 27, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Gray, J. E. (1840) A synopsis of the genera and species of the class Hypostoma (Asterias Linnaeus). Annals of the Magazine of Natural History, Series 1, 6 (37), 175 - 184 + 275 - 290. https: // doi. org / 10.1080 / 03745484009443296","Lamarck, J. B. (1816) Histoire naturelle des animaux sans vertebres. Meline, Bruxelles, 568 pp.","Fisher, W. K. (1911) Asteroidea of the North Pacific and adjacent waters. 1. Phanerozonia and Spinulosa. Bulletin of the United States National Museum, 76, i - xiii + 1 - 420 pp., 122 pls. https: // doi. org / 10.5479 / si. 03629236.76. i","Clark, A. & Tortonese, E. (1986) Echinaster sepositus madseni n. subsp., from West Africa (Echinodermata: Asteroidea). Atti della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale, Milano, 127, 65 - 71.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Fontanella, F. M. & Hopkins, T. S. (2003) Preliminary phylogeny of Echinaster (Othilia) from the Gulf of Mexico based on morphological characters (Echinodermata: Asteroidea). In: Feral, J. P. & David, B. (Eds.), Proceedings of the Sixth European Conference on Echinoderm Research, Banyuls-sur-Mer, France, 3 - 7 September 2001. Balkema, Rotterdam, pp. 91 - 95.","Clark, A. M. (1987) Notes on Atlantic and other Asteroidea Echinasteridae. Bulletin of the British Museum of Natural History, 53, 65 - 78.","Nobre, C. & Campos-Creasey, L. S. (2000) Efeito da salinidade no desenvolvimento larval de Echinaster echinophorus (Lamarck 1816) (Echinodermata: Asteroidea). Brazilian Journal of Aquatic Science and Technology, 4, 61 - 68. https: // doi. org / 10.14210 / bjast. v 4 n 1. p 61 - 68","Alves, R. N., Neto, N. L., Brooks, S. E. & Albuquerque, U. P. (2009) Commercialization of animal-derived remedies as complementary medicine in the semi-arid region of Northeastern Brazil. Journal of Ethnopharmacology, 124, 600 - 608. https: // doi. org / 10.1016 / j. jep. 2009.04.049","Alves, R. N. & Dias, T. L. P. (2010) Uso de invertebrados na medicina popular no Brasil e suas implicacoes para conservacao. Tropical Conservation Science, 3 (2), 159 - 174. https: // doi. org / 10.1177 / 194008291000300204","Martins, L., Souto, C., Magalhaes, W. F., Alves, O. F. S., Rosa, I. L. & Sampaio, C. L. S. (2012) Echinoderm harvesting in Todosos-Santos Bay, Bahia State, Brazil: the aquarium trade. Sitientibus serie Ciencias Biologicas, 12 (1), 53 - 59. https: // doi. org / 10.13102 / scb 123"]}

Genus Luidia Forbes, 1839 Type species. Luidia ciliaris (Philippi, 1837) (type by monotypy). Remarks. The genus Luidia contains 49 species. Of the 12 Atlantic species, six occur in Brazil — L. alternata (Say, 1825), L. barbadensis Perrier, 1881, L. clathrata (Say, 1825), L. ludwigi Bell, 1917, L. sarsi Düben & Koren, 1845 and L. senegalensis (Lamarck, 1816) — L. patriae Bernasconi, 1941 occurs in Argentina and Uruguay, and five other species are restricted to the North Atlantic and Southeast Atlantic— L. atlantidea Madsen, 1950, L. ciliaris (Philippi, 1837), L. heterozona Fisher, 1940, L. lawrencei Hopkins & Knott, 2010, and L. sagamina Döderlein, 1920., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on page 19, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Forbes, E. (1839) On the Asteriadae of the Irish Sea. Memoirs of the Wernerian Natural History Socierty of Edinburgh, 8, 114 - 129.","Philippi, R. A. (1837) Uber die mit Asterias auranciaca verwanten und verwechselten Asterien der sicilanschen Kuste. Archiv fur Naturgeschichte, 3, 193 - 194.","Say, T. (1825) On the species of the Linnaean genus Asterias inhabiting the coast of the United States. Journal of the Academy of Natural Sciences, Philadelphia, 1 (5), 151 - 154.","Perrier, E. (1881) Description sommaire des especes nouvelles d'Asteries. Museum of Comparative Zoology Bulletin, 9, 1 - 31.","Bell, F. J. (1917) Echinoderma, Part I. Actinogonidiata. Natural History Reports of the British Antarctic \" Terra Nova \" Expedition, Zoology, 4 (1), 1 - 10.","Duben, M. W. V. & Koren, J. (1845) Nya skandinaviska fiskar. Ofversigt af Kongl. Veterskaps-Akademiens Forhandlingar, 2 (1), 9 - 12.","Lamarck, J. B. (1816) Histoire naturelle des animaux sans vertebres. Meline, Bruxelles, 568 pp.","Bernasconi, I. (1941) Los equinodermos de la expedicion del buque oceanografico comodoro rivadavia A. R. A. Physis, 19, 37 - 49.","Madsen, F. J. (1950) The Echinoderms Collected by the Atlantide Expedition 1945 - 46. I. Asteroidea. With Remarks on Other Sea-Stars from Tropical and Northern West Africa. Atlantide Reports, 1, 167 - 222.","Fisher, W. K. (1940) Asteroidea. Discovery Reports, 20, 69 - 306.","Hopkins, T. S. & Knott, K. E. (2010) The establishment of a neotype for Luidia clathrata (Say, 1825) and a new species within the genus Luidia (Asteroidea: Paxillosida: Luidiidae). In: Harris, L. G., Bottger, S. A., Walker, C. W. & Lesser, M. P. (Eds.), Proceedings of the 12 th International Echinoderm Conference, 7 - 11 August 2006, Durham, New Hampshire. Taylor & Francis Group, London, pp. 207 - 212. https: // doi. org / 10.1201 / 9780203869543 - c 32"]}

Othilia brasiliensis (Müller & Troschel, 1842) Figures 13–14 Echinaster brasiliensis Müller & Troschel, 1842: 22, pl. 1, fig. 2. Echinaster brasiliensis — Nunes 1975: 183; Tommasi & Aron 1987: 3; Tommasi et al. 1988: 6; Fernandes et al. 2002: 422; Netto 2006: 30–32, pl. 5a; Alves et al. 2012: 758; Miranda et al. 2012: 144. Echinaster (Othilia) brasiliensis — Clark & Downey 1992: 363–367, fig. 57a, pl. 87A–F; Hopkins et al. 2003: 98–100, figs. 2–6; Brites et al. 2008: 182–183; Lima & Fernandes 2009: 59; Xavier 2010: 75; Benavides-Serrato et al. 2011: 192; Gondim et al. 2014: 37, fig. 11a–e; Alitto et al. 2016: 10, figs. 7g –h; Bumbeer et al. 2016: 8, fig. 4A; Lopes & Ventura 2016; Lopes et al. 2016; Sandino et al. 2017: S294; Bueno et al. 2018: 182, fig. 13; Gurjão & Lotufo 2018: 10; Miranda 2018: 14, fig. 10E; Borrero-Peìrez et al. 2019: 5; Torres & Torres 2019: 412. Echinaster braziliensis — Patrizzi & Dobrovolski 2018: 182. Echinaster guyanensis — Martins et al. 2012, table 1. Material examined (56 specs, 17–65 mm R). BRAZIL. Bahia (12°41’– 13°00’S; 38°29’– 38°53’W)— Salvador: Ribeira beach, intertidal, 1.iii.2006, 36 specs, R 17–29 mm (UFBA 131, 173). Itaparica Island: Medo Island, intertidal, 7.ix.1991, 1 spec, R 29–30 mm (UFBA 31, 37). Cachoeira: Santiago do Iguape, 15 m, 2.x.2012, 1 spec, R 65 mm (UFBA 1710). Madre de Deus: intertidal, 8.iv.2008, 1 spec, R 56 mm (UFBA 625). Itaparica Island: Medo Island, intertidal, 7.ix.1991, 4 specs, R 36–51 mm (UFBA 31, 37); Ponta de Areia beach, 4.vi.2000, 2 specs, R 37–48 mm (UFBA 473). Vera Cruz: Duro beach, 1 m, 20.ix.2009, 2 specs, R 41–42 mm (UFBA 961). Salvador: Plataforma beach, intertidal, 29.vii.2007, 7 specs, R 39–60 mm (UFBA 586); Ribeira beach, intertidal, 15.v.1991, 2 specs, R 31–39 mm (UFBA 38); intertidal, 1.iii.2006, 34 specs, R 36–59 mm (UFBA 131, 173–175); Barra beach, intertidal, 6.vii.2004, 1 spec, R 33 mm (UFBA 33). Comparative material. BRAZIL, 2 specs, R 54–55 mm (ZMB 550–551, syntypes). Description (R 31–65 mm). Disc small, average R/r 4.4 (Fig. 13A–C). Five slender arms, tapering distally, with 9–11 rows of long and sharp spines (ca. 1.0– 1.5 mm) without mammiform base. Small secondary plates between abactinal plates; skin not very thick but obscuring plating. Abactinal region covered by many papulae and glandular cells. Five primary plates on disc, each with a spine forming a pentagon; one central spine (Fig. 13D). Anus near central spine, surrounded by 3–6 spinelets. Madreporite flat, without spinelets. Superomarginal spines same size as abactinal ones. Inferomarginal plates with one spine each, forming a row; inferomarginal spines larger than others (ca. 1.7 mm). Abactinal and marginal spines sharp, straight or slightly curved, tapering. Actinal plates absent. Terminal plate small, curved, with 6–7 small spines. Actinal region with several closed pores (one row plus scattered pores) (Fig. 13E). One curved adambulacral spine. Two subambulacral spines forming a V along the furrow (Fig. 13F), proximal region sometimes with third spine; spines are usually equal, but outer spine may be larger than inner. Interradial region naked, sometimes with a central spine. Oral spines completely covering mouth opening. Tube feet in two rows, sucking disc present. Pedicellariae absent. Ontogenetic variation (R 17–30 mm). Average R/r 3.9 (Fig. 14A–C). Spines proportionally larger and fewer (7 rows) than in large specimen; 4–5 small spines on the terminal plate (Fig. 14D). Two unequal subambulacral spines, outer spine larger than inner (Fig. 14E). Coloration. Specimens in vivo are light to scarlet red. Specimens in ethanol are brown, dark brown or beige. Distribution. Gulf of Mexico, Cuba, Honduras, Nicaragua, Panama, Colombia, Guyana, French Guiana, Uruguay, Argentina, Malvinas Islands (Sandino et al. 2017; Borrero-Peìrez et al. 2019; Mah 2020a). BRAZIL: Pará, Rio Grande do Norte, Paraíba, Pernambuco, Alagoas, Bahia, Espírito Santo, Rio de Janeiro, São Paulo, Paraná, Santa Catarina, Rio Grande do Sul (Rathbun 1879; Verrill 1915; Bernasconi 1956; Brito 1968; Tommasi 1970; Carrera-Rodriguez & Tommasi 1977; Ávila-Pires 1983; Clark & Downey 1992; Hopkins et al. 2003; Miranda et al. 2012; Alvarado & Solís-Marín 2013; Gondim et al. 2014; Alitto et al. 2016; Bumbeer et al. 2016; Bueno et al. 2018; Miranda 2018; Torres & Torres 2019; Mah 2020a). Depth. Intertidal to 60 m (Clark & Downey 1992; present paper). Biological notes. Specimens from Bahia are found in sandy and muddy substrate, commonly on top of rocks. Some populations live in regions of low salinity, such as the delta of the Paraguaçu River. Othilia brasiliensis usually feeds on sponges of the genera Mycale Gray, 1867, Haliclona Grant, 1841 and Tedania Gray, 1867 (Guerrazi et al. 1999), but they also feed on sea anemones, brine shrimp, true shrimp (Queiroz 2012; Alcantara & Costa 2019). Lopes & Ventura (2016) described the early developmental stages of O. brasiliensis [as Echinaster (Othilia) brasiliensis], which is similar to the development of other Echinasteridae species. Othilia brasiliensis is classified as “Least Concern” by the Ministry of the Environment (MMA 2018) and its harvesting in Brazil is currently prohibited (Gurjão & Lotufo 2018). Syntypes. ZMB 550–552. Type locality. Müller & Troschel (1842) were not specific with regards to the type locality of this species and the locality in the specimen label is Brazil. Walenkamp (1979) cited Ubatuba as the type locality but no justification for this assignment was provided. Remarks. Walenkamp (1976) and Gondim et al. (2014) looked for intraspecific variability in specimens from varied size ranges (R 7–41 mm) but ontogenetic variation was not observed (note that Walenkamp’s specimens were O. guyanensis according to A.M. Clark [1987]). Clark & Downey (1992) reported the presence of spinelets in the madreporite of O. brasiliensis, these were not found in the specimens described here. They also mention the presence of 4–5 ambulacral spines; the specimens described here and those by Hopkins et al. (2003) have 3–4 spines. Seixas et al. (2019) described the mitochondrial genome of this species; its 16,219 base pairs are AT-rich., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 27-28, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Muller, J. & Troschel, F. H. (1842) System der Asteriden. Friedrich Viewig und Sohn, Braunschweig, 134 pp.","Nunes, T. B. (1975) Echinoderma da Baia de Aratu (Bahia, Brasil). Memories of the second Latin American Symposium of Oceanographic Biology, Universidad de Oriente, 1975, 179 - 189.","Fernandes, M. L. B., Tommasi, L. R. & Lima, E. J. B. (2002) Filo Echinodermata de Pernambuco. In: Tabarelli, M. & Silva, J. M. C. (Eds.), Diagnostico da Biodiversidade de Pernambuco. Vol. 2. Massangana, Recife, pp. 405 - 427.","Netto, L. F. (2006) Echinodermata do Canal de Sao Sebastiao, Sao Sebastiao. Unpublished Master Thesis, Universidade de Sao Paulo, Sao Paulo, 201 pp.","Alves, R. R. N., Rosa, I. L., Neto, N. L. & Voeks, R. (2012) Animals for the Gods: Magical and Religious Faunal Use and Trade in Brazil. Human Ecology, 40, 751 - 780. https: // doi. org / 10.1007 / s 10745 - 012 - 9516 - 1","Miranda, A. L. S., Lima, M. L. F., Sovierzoski, H. H. & Correia, M. D. (2012) Inventory of the Echinodermata collection from the Universidade Federal de Alagoas. Biota Neotropica, 12 (2), 135 - 146. https: // doi. org / 10.1590 / S 1676 - 06032012000200014","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Hopkins, T. S., Fontanella, F. M. & Ventura, C. R. R. (2003) Morphological diagnosis of three Brazilian starfishes of the genus Echinaster (Othilia). In: Feral, J. P. & David, B. (Eds.), Proceedings of the Sixth European Conference on Echinoderm Research, Banyuls-Sur-Mer, France, 3 - 7 September 2001. Balkema, Rotterdam, pp. 97 - 103.","Lima, E. J. B. & Fernandes, M. L. B. (2009) Diversidade de equinodermos (Echinodermata) no estado de Pernambuco (Brasil). Revista Brasileira de Zoociencias, 11, 55 - 63.","Xavier, L. A. R. (2010) Inventario dos equinodermos do Estado de Santa Catarina, Brasil. Brazilian Journal of Aquatic Science and Technology, 14, 73 - 78. https: // doi. org / 10.14210 / bjast. v 14 n 2. p 73 - 78","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Alitto, R. A. S., Bueno, M. L., Domenico, M. & Borges, M. (2016) Annotated checklist of Echinoderms from Araca Bay, South- eastern Brazil. Check List, 12 (1), 1 - 15.","Bumbeer, J., Cattani, A. P., Chierigatti, N. B. & Rocha, R. M. (2016) Biodiversity of benthic macroinvertebrates on hard substrates in the Currais Marine Protected Area, in southern Brazil. Biota Neotropica, 16 (4), e 20160246. https: // doi. org / 10.1590 / 1676 - 0611 - BN- 2016 - 0246","Lopes, E. M. & Ventura, C. R. R. (2016) Development of the sea star Echinaster (Othilia) brasiliensis, with inference on the evolution of development and skeletal plates in Asteroidea. Biology Bulletin, 230, 25 - 35. https: // doi. org / 10.1086 / BBLv 230 n 1 p 25","Sandino, O. B., Solis-Marin, F. A., Caballero-Ochoa, A. A., Conejeros-Vargas, C. A., Marquez-Borras, F., Ayala-Aguilera, A. P. & Laguarda-Figueras, A. (2017) Equinodermos de Nicaragua: nuevos registros del Pacifico y Caribe Sur. Revista de Biologia Tropical, 65, 288 - 298. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31696","Bueno, M. L., Alitto, R. A. S., Guilherme P. D. B., Domenico, M. D. & Borges, M. (2018) Guia ilustrado dos Echinodermata da porcao sul do Embaiamento Sul Brasileiro. Pesquisa e Ensino em Ciencias Exatas e da Natureza, 2 (2), 169 - 237. https: // doi. org / 10.29215 / pecen. v 2 i 2.1071","Gurjao, L. M. & Lotufo, T. M. C. (2018) Native species exploited by marine aquarium trade in Brazil. Biota Neotropica, 18 (3), e 20170387. https: // doi. org / 10.1590 / 1676 - 0611 - bn- 2017 - 0387","Miranda, A. P. S. (2018) Diversidade e distribuicao de equinodermos da Plataforma Continental Amazonica. Unpublished Honors Thesis, Universidade Federal do Para, Belem, 62 pp.","Torres, V. S. & Torres, F. S. S. (2019) Inventario preliminar dos invertebrados marinhos observados na praia da Pinheira, Palhoca, SC, Brasil. UNISANTA Bioscience, 8 (4), 408 - 422.","Patrizzi, N. & Dobrovolski, R. (2018) Integrating climate change and human impacts into marine spatial planning: a case study of threatened starfish species in Brazil. Ocean & Coastal Management, 161, 177 - 188. https: // doi. org / 10.1016 / j. ocecoaman. 2018.05.003","Martins, L., Souto, C., Magalhaes, W. F., Alves, O. F. S., Rosa, I. L. & Sampaio, C. L. S. (2012) Echinoderm harvesting in Todosos-Santos Bay, Bahia State, Brazil: the aquarium trade. Sitientibus serie Ciencias Biologicas, 12 (1), 53 - 59. https: // doi. org / 10.13102 / scb 123","Mah, C. L. (2020 a) World Asteroidea Database. Available from: http: // www. marinespecies. org / asteroidea / (accessed 30 April 2020)","Rathbun, R. (1879) A list of the Brazilian echinoderms, with notes on their distribution. Transactions of the Connecticut Academy of Arts and Sciences, 5 (3), 139 - 158. https: // doi. org / 10.5962 / bhl. title. 16126","Verrill, A. E. (1915) Report on the Starfishes of the West Indies, Florida, and Brazil, Including Those Obtained by the Bahama Expedition from the University of Iowa in 1893. Bulletin of the State of University of Iowa, 7, 1 - 232, 29 pls. https: // doi. org / 10.5962 / bhl. title. 12035","Bernasconi, I. (1956) Dos nuevos equinodermos de la costa del Brasil. Neotropica, 2, 33 - 36, 2 figs.","Brito, I. M. (1968) Asteroiides e equinoiides do Estado da Guanabara e adjace ˆ ncias. Boletim do Museu Nacional, 260, 1 - 51.","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Carrera-Rodriguez, C. J. & Tommasi, L. R. (1977) Asteroidea de la plataforma continental de Rio Grande do Sul (Brasil), coleccionados durante los viajes del N / Oc. \" Prof. W. Besnard \" para el proyecto Rio Grande do Sul. Boletim do Instituto Oceanografico da Universidade de Sao Paulo, 26 (1), 51 - 130. https: // doi. org / 10.1590 / S 0373 - 55241977000100004","Avila-Pires, T. C. S. A. (1983) Contribuicao ao estudo do genero Echinaster Muller & Troschel, 1840 (Echinodermata, Asteroidea) no litoral brasileiro. Anais da Academia Brasileira de Ciencias, 55 (4), 431 - 448.","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Vol. 1. Springer, Berlin, 658 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","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, pls. XXVII - XXVIII.","Grant, R. E. (1841) Porifera. In: Bailliere, H. (Ed.), Outlines of comparative anatomy. Vol. 1. Bailliere, London, pp. 5 - 9 + 310 - 313, pls. II - IV. https: // doi. org / 10.5962 / bhl. title. 28330","Queiroz, C. H. (2012) Estrateigias e fisiologia do consumo e digestaTo de esponjas (Porifera) por Echinaster brasiliensis (Echinodermata: Asteroidea). Unpublished Master Thesis, Universidade de SaTo Paulo, SaTo Paulo, 74 pp.","Alcantara, A. A. & Costa, F. M. (2019) Resposta comportamental de Echinaster (Othilia) brasiliensis Muller & Troschel, 1842, (Echinodermata, Asteroidea) com base em preferencia alimentar e observacao em cativeiro. Acta Scientiae et Technicae, 7 (1), 21 - 25. https: // doi. org / 10.17648 / uezo-ast-v 7 i 1.263","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp.","Walenkamp, J. H. C. (1979) Asteroidea (Echinodermata) from the Guyana shelf. Zoologische Verhandelingen, 170, 1 - 97.","Walenkamp. J. H. C. (1976) The asteroids of the coastal waters of Surinam. Zoologische Verhandelingen, 147, 1 - 91, 18 pls.","Clark, A. M. (1987) Notes on Atlantic and other Asteroidea Echinasteridae. Bulletin of the British Museum of Natural History, 53, 65 - 78.","Seixas, V. C., Ventura, C. R. R. & Palva, P. C. (2019) The complete mitochondrial genome of the sea star Echinaster (Othilia) brasiliensis (Asteroidea: Echinasteridae). Conservation Genetics Resources, 11, 151 - 155. https: // doi. org / 10.1007 / s 12686 - 018 - 0986 - 3"]}

Coscinasterias tenuispina (Lamarck, 1816) Asterias tenuispina Lamarck, 1816: 561–562. Coscinasterias tenuispina — Clark & Downey 1992: 427–428, figs. 63h, i, pl. 101, figs. A, B; Alves et al. 2002; Pérez-Ruzafa et al. 2002: 280–281; Netto 2006: 34, fig. 16c, pl. 5b; Koukouras et al. 2007: 70; Ventura et al. 2007: 228; Ventura et al. 2008c: 172; Micael & Costa 2010: 321; Micael et al. 2012: 2–4; Gondim et al. 2014: 43–44, fig. 9i–m; Fonseca 2015; Madeira et al. 2019: 64–66, fig. 12. Distribution. France, Italy, Croatia, Spain, Portugal, Azores, Madeira, Turkey, Greece, Malta, Canary Islands, Bermuda, Cape Hatteras, Cape Verde, North Carolina, Cuba, Gulf of Mexico, Guyana, Ascension Island, Saint Helena (Koukouras et al. 2007; Madeira et al. 2019; GBIF, 2020). BRAZIL: Bahia, Espírito Santo, Rio de Janeiro, São Paulo, Santa Catarina (Rathbun 1879; Verrill 1915; Brito 1962; Tommasi 1970; Clark & & Downey 1992; Ventura et al. 2007; Gondim et al. 2014; Fonseca 2015). Depth. 0–165 m (Clark & Downey, 1992). Biological notes. Coscinasterias tenuispina inhabits rocks, biogenic detritus, sandy and silty sand bottoms, and meadows of seagrass (Koukouras et al. 2007), and typically feeds on mussels and other epifaunal organisms (Ventura et al. 2007). Adults and juveniles are believed to inhabit different depths (Tortonese 1965). This species can reproduce asexually by fission (Alves et al. 2002), and Fonseca (2015) has shown that the contribution of sexual reproduction to maintain four populations in Rio de Janeiro is very low compared to that of asexual reproduction, contrary to the findings by Pazotto (2010) in the same populations. This species is classified as “Vulnerable” (baseline data indicates that the population size has been reduced by at least 30%) by the Ministry of the Environment (MMA 2018). Holotype. The type specimen, most likely deposited at the Muséum National d’Histoire Naturelle, Paris, is currently lost (Marc Eléaume and Christopher Mah, pers. comm. on 30 Sep 20). Type locality. European Ocean (Lamarck, 1816; Clark & Downey, 1992), possibly the Mediterranean Sea. Remarks. Although C. tenuispina has been recorded for Bahia, few specimens have been collected and as Gondim et al. (2014), we could not examine any. We did not have access to the specimens at the EQMN and the single-armed specimen recorded by Rathbun (1879), deposited at the YPM (YPM IZ 1582.EC), is apparently missing. The last reference that we have for this specimen is that it was examined by Addison Emery Verrill, together with three other specimens from the North Atlantic (YPM IZ 1453.EC and YPM IZ 1584.EC); the label from YPM 1582.EC was found in a jar with a single specimen tagged with the label YPM IZ 1453.EC (Daniel Drew and Eric Lazo-Wasem, pers. comm. on 15 Jul 20). Downey (1973) considered the specimen (R 30 mm) she examined from Florida as a juvenile based on morphological differences from larger specimens, for example, the presence of only a few dorsolateral plates (vs. one or two regular rows of dorsolateral plates, armed like the carinal plates), absence of interactinal plates (vs. one row of interactinal plates), small pedicellariae rarely found within the furrow (vs. pedicellariae frequently large and abun- dant on all surfaces). Madeira et al. (2019) also noted differences between juveniles and adults when they analyzed specimens (R 3–110 mm) from the Azores, however, they considered juvenile’s specimens with R Coscinasterias tenuispina differs from C. acutispina by having the adambulacral spines frequently arranged in one row on the inner half of the arm (vs. two rows), from C. muricata by having 6–9 arms (vs. 9–11), and from C. calamaria by having crossed pedicellariae with a well-developed terminal tooth and straight pedicellaria with short stubs at the tips (vs. crossed pedicellariae with little or no development of an enlarged tooth, and straight pedicellariae with little or no modification at the tips). Clark & Downey (1992) pointed out that the taxonomic status of the C. tenuispina specimens from Brazil is uncertain because of morphological differences in relation to specimens from other regions. For example, they noticed that the Brazilian (Cabo Frio, RJ) specimen that they examined has poorly developed straight pedicellariae, never large, and crossed pedicellariae with little or no sign of an enlarged tooth. Clark & Downey (1992) then suggested that C. calamaria could be a subspecies of C. tenuispina based on the morphological similarities observed among Brazilian and South African specimens. However, the Brazilian specimen that they examined had R 33 mm. Using molecular data, Waters & Roy (2003) confirmed the separation between Brazilian populations (n=3) and North Atlantic and Mediterranean populations, but they also found that C. calamaria is more closely related to C. acutispina than to C. tenuispina. An integrated taxonomic revision of juvenile and adult specimens of C. tenuispina along the Brazilian coast is needed to properly characterize its populations., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 55-56, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Lamarck, J. B. (1816) Histoire naturelle des animaux sans vertebres. Meline, Bruxelles, 568 pp.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Alves S. L. S., Pereira, A. D. & Ventura, C. R. R. (2002) Sexual and asexual reproduction of Coscinasterias tenuispina (Echinodermata: Asteroidea) from Rio de Janeiro, Brazil. Marine Biology, 140, 95 - 101. https: // doi. org / 10.1007 / s 002270100663","Perez-Ruzafa, A., Entrambasaguas, L., Espejo, C., Marcos, C. & Bacallado, J. J. (2002) Fauna de equinodermos (Echinodermata) de los fondos rocosos infralitorales del archipielago de Salvajes (Oceano Atlantico). Revista de la Academia Canaria de Ciencias, 14 (3 - 4), 277 - 296.","Netto, L. F. (2006) Echinodermata do Canal de Sao Sebastiao, Sao Sebastiao. Unpublished Master Thesis, Universidade de Sao Paulo, Sao Paulo, 201 pp.","Koukouras, A., Sinis, A. I., Bobori, D., Kazantzidis, S. & Kitsos, M. S. (2007) The echinoderm (Deuterostomia) fauna of the Aegean Sea, and comparison with those of the neighboring seas. Journal of Biological Research, 7, 67 - 92.","Ventura, C. R. R., Verissimo, I., Nobre, C. C. & Zama, P. C. (2007) Filo Echinodermata. In: Lavrado, H. P. & Viana, M. S. (Eds.), Atlas de invertebrados marinhos da regiao central da Zona Economica Exclusiva brasileira. Parte 1. Museu Nacional do Rio de Janeiro, Serie Livros 25. Museu Nacional do Rio de Janeiro, Rio de Janeiro, pp. 258.","Ventura, C. R. R., Barcellos, C. F. & Souza, I. V. (2008 c) Coscinasterias tenuispina Lamarck, 1816. In: Machado, A. B. M., Drummond, G. M. & Paglia, A. P. (Eds.), Livro Vermelho da Fauna Brasileira Ameacada de Extincao. Ministerio do Meio Ambiente e Fundacao Biodiversitas, Brasilia, pp. 172 - 173.","Micael, J. & Costa A. C. (2010) Echinodermata. In: Borges, P. A. V., Costa, A., Cunha, R., Gabriel, R., Goncalves, V., Martins, A. F., Melo, I., Parente, M., Raposeiro, P., Rodrigues, P., Santos, R. S., Silva, L., Vieira, P. & Vieira, V. (Eds.), A list of the terrestrial and marine biota from the Azores. Principia, Oeiras, pp. 321 - 323.","Micael, J., Alves, M. J., Jones, M. B. & Costa, A. C. (2012) Diversity of shallow-water asteroids (Echinodermata) in the Azorean Archipelago. Marine Biodiversity Records, 5 (3), 1 - 10. https: // doi. org / 10.1017 / S 1755267211000534","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Fonseca, B. F. (2015) Contribuicao da reproducao sexuada e assexuada e conectividade entre populacoes da estrela-do-mar Coscinasterias tenuispina (Lamarck, 1816) na costa do Estado do Rio de Janeiro. Unpublished Master Thesis, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, 64 pp.","Madeira, P., Kroh, A., Cordeiro, R., Frias, D. E., Martins, A. M. & Avila, S. P. (2019) The Echinoderm Fauna of the Azores (NE Atlantic Ocean). Zootaxa, 4639 (1), 1 - 231. https: // doi. org / 10.11646 / zootaxa. 4639.1","GBIF Secretariat (2020) Coscinasterias tenuispina (Lamarck, 1816) in GBIF Backbone Taxonomy. Available from: https: // www. gbif. org / species / 2273832 (accessed 1 October 2020).","Rathbun, R. (1879) A list of the Brazilian echinoderms, with notes on their distribution. Transactions of the Connecticut Academy of Arts and Sciences, 5 (3), 139 - 158. https: // doi. org / 10.5962 / bhl. title. 16126","Verrill, A. E. (1915) Report on the Starfishes of the West Indies, Florida, and Brazil, Including Those Obtained by the Bahama Expedition from the University of Iowa in 1893. Bulletin of the State of University of Iowa, 7, 1 - 232, 29 pls. https: // doi. org / 10.5962 / bhl. title. 12035","Brito, I. M. (1962) Ensaio de catailogo dos equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zooloigicos, 13, 1 - 10.","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Tortonese, E. (1965) Echinodermata. Fauna d'Italia. Calderini, Bologna, 419 pp.","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp.","Downey, M. E. (1973) Starfishes from the Caribbean and the Gulf of Mexico. Smithsonian Contributions Zoology, 126, 1 - 158. https: // doi. org / 10.5479 / si. 00810282.126","Waters J. M. & Roy, M. S. (2003) Global phylogeography of the fissiparous sea-star genus Coscinasterias. Marine Biology, 142, 185 - 191. https: // doi. org / 10.1007 / s 00227 - 002 - 0915 - 5"]}

Astropecten cingulatus Sladen, 1883 Astropecten cingulatus Sladen, 1883: 266–267. Astropecten cingulatus — Brito 1962: 3; 1968: 9; Clark & Downey 1992: 33, figs. 9d, 11e–g, pl. 6E, F; De Léo & Pires-Vanin 2006: 273; Ventura et al. 2007: 231; Xavier 2010: 75; Benavides-Serrato et al. 2011: 217; Sandino et al. 2017: S293; Lawrence & Cobb 2017; Lawrence et al. 2018a: 130, figs. 2–3, 6, 8–10, 12–14, 17; Lawrence et al. 2018b: figs. 1–2, 4a–b, 6, 9–10, 13a; Cobb et al. 2019: figs. 7, 15c, 16; 17c, f; Cunha et al. 2020: 50. Distribution. U.S.A. (NC, SC, FL), The The Bahamas, Gulf of Mexico, Caribbean, Mexico, Costa Rica, Nicaragua, Panama, Guyana, French Guyana, Uruguay, Argentina (Bernasconi 1941; Downey 1973; Walenkamp 1979; Clark & Downey 1992; Benavidez-Serrato et al. 2011; Sandino et al. 2017; Lawrence et al. 2018a; Mah 2020a). BRAZIL: Pernambuco, Bahia, Vitória-Trindade Seamount Chain, Martin Vaz Archipelago, Trindade Island, Rio de Janeiro, Vitória Island-SP, São Paulo, Santa Catarina (Brito 1962, 1968; Tommasi 1970, 1985; Carrera-Rodríguez & Tommasi 1977; Tommasi & Aron 1987; Manso, 1989; Clark & Downey 1992; De Léo & Pires-Vanin 2006; Ventura et al. 2007; Alvarado et al. 2008; Xavier 2010; Cunha et al. 2020). Depth. 11 to 1350 m (Lawrence et al. 2018a). Biological notes. Lives on soft substrates, feeding on gastropods, bivalves, crustaceans and cirripeds (Ventura et al. 2007), and exhibits an annual reproductive cycle (Ventura et al. 1998). This species is classified as “Least Concern” by the Ministry of the Environment (MMA 2018). Holotype. NHM-UK 1890.5.7.304 (Lawrence et al. 2018a). Type locality. Pernambuco, Brazil. Remarks. To our knowledge, only the EQMN has specimens from Bahia and we were unable to obtain specimens for examination. Records from this region are scarce and often dubious. For example, the specimen of A. cingulatus depicted by Queiroz (2006, fig. 3) is most likely an A. marginatus, and the specimens depicted by Pelaes (2008, fig. 3b) is possibly a smaller specimen of A. brasiliensis brasiliensis because of the presence of superomarginal spines (but note that the quality of the image is poor). The specimen of A. cingulatus depicted by Gondim et al. (2014, fig. 5e–h) also have spines on the superomarginal plates (some of these have fallen but spine scar is visible) and represents another species. Lawrence et al. (2018a) found some specimens of A. cingulatus with small spinelets on the proximal superomarginal plates, but these were never prominent. Ontogenetic variation in A. cingulatus have not been fully studied. Lawrence et al. (2018b) noted that specimens with R A. cingulatus, see Lawrence & Cobb (2017) and Lawrence et al. (2018a; b)., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 14-15, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Sladen, W. P. (1883) The Asteroidea of H. M. S Challenger Expedition. (Preliminary notices). 2. Astropectinidae. Zoological Journal of the Linnean Society, 17, 214 - 269. https: // doi. org / 10.1111 / j. 1096 - 3642.1883. tb 02022. x","Brito, I. M. (1962) Ensaio de catailogo dos equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zooloigicos, 13, 1 - 10.","Brito, I. M. (1968) Asteroiides e equinoiides do Estado da Guanabara e adjace ˆ ncias. Boletim do Museu Nacional, 260, 1 - 51.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","De Leo, F. C. & Pires-Vanin, A. M. S. (2006) Benthic megafauna communities under the influence of the South Atlantic Central Water intrusion onto the Brazilian SE shelf: a comparison between an upwelling and a non-upwelling ecosystem. Journal of Marine Systems, 60, 268 - 284. https: // doi. org / 10.1016 / j. jmarsys. 2006.02.002","Ventura, C. R. R., Verissimo, I., Nobre, C. C. & Zama, P. C. (2007) Filo Echinodermata. In: Lavrado, H. P. & Viana, M. S. (Eds.), Atlas de invertebrados marinhos da regiao central da Zona Economica Exclusiva brasileira. Parte 1. Museu Nacional do Rio de Janeiro, Serie Livros 25. Museu Nacional do Rio de Janeiro, Rio de Janeiro, pp. 258.","Xavier, L. A. R. (2010) Inventario dos equinodermos do Estado de Santa Catarina, Brasil. Brazilian Journal of Aquatic Science and Technology, 14, 73 - 78. https: // doi. org / 10.14210 / bjast. v 14 n 2. p 73 - 78","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Sandino, O. B., Solis-Marin, F. A., Caballero-Ochoa, A. A., Conejeros-Vargas, C. A., Marquez-Borras, F., Ayala-Aguilera, A. P. & Laguarda-Figueras, A. (2017) Equinodermos de Nicaragua: nuevos registros del Pacifico y Caribe Sur. Revista de Biologia Tropical, 65, 288 - 298. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31696","Lawrence, J. M. & Cobb, J. (2017) Validation of Astropecten jarli Madsen, 1950 and implications for A. cingulatus Sladen, 1883 (Paxillosida: Astropectinidae). Zootaxa, 4269 (1), 101 - 114. https: // doi. org / 10.11646 / zootaxa. 4269.1.4","Lawrence, J. M., Cobb, J. C. & Herrera, J. C. (2018 a) Synonymy of Astropecten nitidus Verrill, 1915 with Astropecten cingulatus Sladen, 1883 and Astropecten comptus Verrill, 1915 with Astropecten articulatus (Say, 1825). Bulletin of the Peabody Museum of Natural History, 59 (2), 127 - 157. https: // doi. org / 10.3374 / 014.059.0203","Lawrence, J. M., Cobb, J. C., Herrera, J. C., Duran-Gonzalez, A. & Solis-Marin, F. A. (2018 b) Morphological comparison of Astropecten cingulatus and a new species of Astropecten (Paxillosida, Astropectinidae) from the Gulf of Mexico. Zootaxa, 4407 (1), 86 - 100. https: // doi. org / 10.11646 / zootaxa. 4407.1.5","Cobb, J. C., Lawrence, J. M., Herrera, J. C. & Janies, D. (2019) A new species of Astropecten (Echinodermata: Asteroidea: Paxillosida: Astropectinidae) and a comparison of the Astropecten species from the Gulf of Mexico and the East Florida Shelf. Zootaxa, 4612 (3), 301 - 325. https: // doi. org / 10.11646 / zootaxa. 4612.3.1","Cunha, R., Tavares, M. & Mendonca, J. B. (2020) Asteroidea (Echinodermata) from shallow-waters of the remote oceanic archipelago Trindade and Martin Vaz, southeastern Atlantic, with taxonomic and zoogeographical notes. Zootaxa, 4742 (1), 31 - 56. https: // doi. org / 10.11646 / zootaxa. 4742.1.2","Bernasconi, I. (1941) Los equinodermos de la expedicion del buque oceanografico comodoro rivadavia A. R. A. Physis, 19, 37 - 49.","Downey, M. E. (1973) Starfishes from the Caribbean and the Gulf of Mexico. Smithsonian Contributions Zoology, 126, 1 - 158. https: // doi. org / 10.5479 / si. 00810282.126","Walenkamp, J. H. C. (1979) Asteroidea (Echinodermata) from the Guyana shelf. Zoologische Verhandelingen, 170, 1 - 97.","Mah, C. L. (2020 a) World Asteroidea Database. Available from: http: // www. marinespecies. org / asteroidea / (accessed 30 April 2020)","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Tommasi, L. R. (1985) Equinodermes da regiao da Ilha da Vitoria (SP). Relatorios Internos do Instituto Oceanografico, Universidade de Sao Paulo, 13, 1 - 5.","Carrera-Rodriguez, C. J. & Tommasi, L. R. (1977) Asteroidea de la plataforma continental de Rio Grande do Sul (Brasil), coleccionados durante los viajes del N / Oc. \" Prof. W. Besnard \" para el proyecto Rio Grande do Sul. Boletim do Instituto Oceanografico da Universidade de Sao Paulo, 26 (1), 51 - 130. https: // doi. org / 10.1590 / S 0373 - 55241977000100004","Manso, C. L. C. (1989) Os Echinodermata da plataforma continental interna entre Cabo Frio e Saquarema, Rio de Janeiro, Brasil. Revista Brasileira de Biologia, 49, 355 - 359.","Alvarado, J. J., Solis-Marin, F. A. & Ahearn, C. (2008) Equinodermos (Echinodermata) del Caribe Centroamericano. Revista de Biologia Tropical, 56, 37 - 55.","Ventura, C. R. R., Santos, J. S., Falcao, A. P. C. & Fiori, C. S. (1998) Reproduction and food intake in Astropecten cingulatus (Asteroidea: Paxillosid) in the upwelling environment of Cabo Frio (Brazil). In: Mooi, R. & Telford, M. (Eds.), Proceedings of the Ninth International Echinoderm Conference, San Francisco, California, 5 - 9 August 1996. Balkema, Rotterdam, pp. 313 - 318.","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp.","Queiroz, A. C. M. (2006) Dinamica populacional das estrelas do mar Astropecten brasiliensis Muller & Troschel, 1842, Astropecten cingulatus Sladen, 1889 e Luidia senegalensis (Lamarck, 1816) (Echinodermata: Asteroidea) presentes na fauna acompanhante da pesca do camarao no litoral de Ilheus (Bahia). Unpublished Master Thesis, Universidade Federal da Paraiba, Joao Pessoa, Paraiba, 92 pp.","Pelaes, M. A. L. (2008) Composicao, distribuicao espacial e temporal das especies de Echinodermata na plataforma rasa do sul da Bahia num trecho entre Itacare e Canavieiras. Unpublished Master Thesis, Universidade Estadual de Santa Cruz, Ilheus, Bahia, 97 pp.","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Sladen, W. P. (1889) Asteroidea. Report of the Scientific Results of H. M. S. Challenger, 30, 1 - 893. https: // doi. org / 10.1111 / j. 1096 - 3642.1882. tb 02281. x"]}

Nymphaster arenatus (Perrier, 1881) Figures 19–20 Pentagonaster arenatus Perrier, 1881: 21; 1884: 236, pl. 7, figs. 3–4. Nympluister basilicus — Tommasi, 1970: 12, fig. 35. Nymphaster arenatus — Clark & Downey 1992: 254–255, figs. 38d, 41e, f, pl. 61A–C; Entrambasaguas 2003: 95; Entrambasa- guas 2008: 59; Campos et al. 2010b: 149, fig. 5C; Benavides-Serrato et al. 2011: 165; Costa et al. 2015; Soaréz 2016: 78; Sandino et al. 2017: S294; Rubio-Polania et al. 2018: 190; Borrero-Peìrez et al. 2019: 5; Mah 2020b: 230, fig. 13A–E. Material examined. (4 specs, 45–70 mm R). BRAZIL. Bahia, Canavieiras (15º39’S; 38º32’W)— 1496 m, 26.xii.1887, 4 specs, R 45–70 mm (USNM 18524). Comparative material. BRAZIL. Rio de Janeiro, Cabo Frio (19°43’S; 38°36’W), 29.vi.1999, 1 spec, R 52 mm (EQMN 2325). Description (arms broken: R 70 mm, r 21 mm and R 69 mm and r 27 mm). Body stellate, flat, broad disc, long arms (Fig. 19A–B); R/r 2.9, 19 SM plates, R/SM# 3.68. Abactinal plates low-tabulate, irregularly round, completely covered by regularly spaced, rounded granules; 4–12 central granules. Some plates with small, excavated sugar-tong pedicellariae (Fig. 19C). Papular area includes center of disc; six papular pores surround each plate; single papule in each pore. Madreporite larger than adjacent abactinal plates (Fig. 19D). Subcentral anus, surrounded by small, conical spinelets. Superomarginal plates in contact throughout length of arm, covered by large, regularly-spaced, rounded granules. Terminal plates of specimens broken. Inferomarginal plates about twice as wide as long in interradial arc, square-shaped in middle region of arm and about twice as long as wide in distal region of arm. Granulation of inferomarginal plates similar to that of superomarginal plates. Actinal plates rhombic to polygonal, covered by regularly spaced, large, rounded granules slightly larger than those on marginal plates. Some actinal plates with excavate, sugar-tong pedicellariae almost twice as large as abactinal pedicellariae. Adambulacral plates about 1–1.5x as long as wide, half the size of adjacent actinal plates. Furrow margin strongly angular. Apophyses poorly developed from 3 rd –4 th plate, fully developed from eighth plate (Fig. 19E). Eight adambulacral spines on proximal plates; number of spines increases with apophysis development, up to 11 spines distally (Fig. 19F). Adambulacral spines compressed, moderately long, with rounded tips. 3–4 irregular rows of subambulacral “spines” with 4–5 large, rounded granules slightly taller than actinal granules. Some of first row spines elongated (Fig. 19F). Spatulate pedicellariae with 2–4 valves, until 4 th –5 th adambulacral plate. Oral plates long, triangular, with 9–10 spines each, similar to those of adambulacral plates; median spines slightly taller and more compressed than others (Fig. 19H). Ontogenetic variation (arms broken: R 45 and 55 mm, r 15 mm). Average R/r 3.3. Differ from largest specimen by having 4–8 central granules on abactinal plates (Fig. 20C); six adambulacral spines in the proximal region; 2–3 irregular rows of subambulacral “spines”, with 2–5 granules (Fig. 20F); pedicellariae until the tenth adambulacral plate (Fig. 20F); eight oral spines (Fig. 20H). Coloration. The abactinal surface is orange and the actinal surface is cream-colored (Halpern 1970b). Distribution. Gulf of Mexico, Mexico, Nicaragua, Colombia (Soaréz 2016; Sandino et al. 2017; Rubio-Polania et al. 2018; Borrero-Peìrez et al. 2019; Mah 2020b). BRAZIL: Bahia, Rio de Janeiro (Ventura et al. 2007; Campos et al. 2010b; Costa et al. 2015). Depth. 100–3000 m (Clark & Downey 1992; NMNH 1592525; FSBC I 74877). Biological notes. Nymphaster arenatus lives on soft, unconsolidated sediment and is classified as scavenger and predator (Costa et al. 2015; Wagstaff et al. 2014; Mah 2020b). It feeds on fouling organisms, debris, epifauna and decomposing organisms (Ventura et al. 2007). Campos et al. (2010b) filmed a specimen with an inflated disc in the Campus Basin, between 700–1000 m of depth. A species of polychaete has been found within thestar’s interradius, possibly in a commensal interaction in which the polychaete consumes leftovers from food caught by the sea star (Mah 2020b). Lectotype. MCZ 437 (designated by Halpern [1970b]). Type locality. Off Barbados (Halpern 1970b). Remarks. All specimens examined here have broken arm-tips such that the number of superomarginal plates could not be counted. Some of the ontogenetic variation observed here were also noted by Halpern (1970b). The variation in number of proximal adambulacral spines, however, is not consistent across studies. The specimens R 45–55 mm described here and the specimen from Rio de Janeiro (R 52 mm) have six spines, the specimen described by Clark & Downey (1992) (R 61 mm) has about seven spines; the specimens R 69–70 mm described here have 7–8 spines, and the specimen described by Halpern (1970b) (R 96 mm) has 6–7 spines. The variation in the number of irregular rows of subambulacral “spines” seems to be more consistent. The specimens R 45–55 mm have 2–3 rows, the specimen R 61 mm has 1–4 rows, and the specimens R 69–96 mm have 3–4 rows. The same applies for the number of oral spines as follows: the specimens R 45–55 mm have 6–8 spines, the specimen R 61 mm has 8–12 spines, the specimens R 69–70 have 9–10 spines and the specimen 96 mm has 10–11 spines. Fisher (1913) used this character to separate species of Nymphaster from the Philippines and most likely overestimated the number of species recorded: a total of nine. Genus Plinthaster Verrill, 1899 Type species. Plinthaster dentatus (Perrier, 1884) (type by subsequent designation by Fisher, 1910). Remarks. Plinthaster has four valid species: P. ceramoidea (Fisher, 1906) (Pacific Ocean), P. dentatus (Perrier, 1884) (Amphiatlantic, and Pacific Ocean), P. lenaigae Mah, 2018 and P. untiedtae Mah, 2018 (Indian Ocean). Only P. dentatus is recorded in Brazil., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 36-39, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Perrier, E. (1881) Description sommaire des especes nouvelles d'Asteries. Museum of Comparative Zoology Bulletin, 9, 1 - 31.","Perrier, E. (1884) Memoire sur les etoiles de mer recueillies dans la mer des Antilles et le golfe du Mexique: durant les expeditions de dragace faites sous la direction de M. Alexandre Agassiz. Archives Museum National d'histoire Naturelle Paris, 6 (2), 127 - 276. https: // doi. org / 10.5962 / bhl. title. 82184","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Entrambasaguas, L. (2003) Estudio de la fauna de equinodermos del archipielago de Cabo Verde: escalas de variabilidad espacial y factores que explican su distribucion. Tesis de licenciatura no publicada. Universidad de Murcia, Murcia. [unknown pagination]","Campos, L. S., Moura, R. B., Alcantara, P. F., Vasconcelos, R. F., Curbelo-Fernandez, M. P., Verissimo, I. & Cavalcanti, G. H. (2010 b) On two new records of the Family Brisingidae (Echinodermata: Asteroidea) from the Brazilian continental margin. In: Harris, L. G., Boettger, S. A., Walker, C. W. & Lesser, M. P. (Eds.), Proceedings of the 12 th International Echinoderm Conference, Durham, New Hampshire, 7 - 11 August 2006. CRC Press Boca Raton, Florida and Balkema Taylor & Francis Group, London, pp. 139 - 146. https: // doi. org / 10.1201 / 9780203869543 - c 21","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Costa, P. A. S., Mincarone, M. M., Braga, A. C., Martins, A. S., Lavrado, H. P., Haimovici, M. & Falcao, A. P. C. (2015) Megafaunal communities along a depth gradient on the tropical Brazilian continental margin. Marine Biology Research, 11 (10), 1053 - 1064. https: // doi. org / 10.1080 / 17451000.2015.1062521","Sandino, O. B., Solis-Marin, F. A., Caballero-Ochoa, A. A., Conejeros-Vargas, C. A., Marquez-Borras, F., Ayala-Aguilera, A. P. & Laguarda-Figueras, A. (2017) Equinodermos de Nicaragua: nuevos registros del Pacifico y Caribe Sur. Revista de Biologia Tropical, 65, 288 - 298. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31696","Rubio-Polania, J. C., Torruco-Gomez, D., Gonzalez-Solis, A., Ordaz, J. & Caamal-Jimenez, Y. (2018) Benthic megafauna of outer margins of the continental shelf of Yucatan Peninsula. Regional Studies in Marine Science, 24, 184 - 195. https: // doi. org / 10.1016 / j. rsma. 2018.08.014","Mah, C. L. (2020 b) New species, occurrence records and observations of predation by deep-sea Asteroidea (Echinodermata) from the North Atlantic by NOAA ship Okeanos Explorer. Zootaxa, 4766 (2), 201 - 260. https: // doi. org / 10.11646 / zootaxa. 4766.2.1","Halpern, J. A. (1970 b) Goniasteridae (Echinodermata: Asteroidea) of the straits of Florida. Bulletin of Marine Science, 20 (1), 193 - 286.","Ventura, C. R. R., Verissimo, I., Nobre, C. C. & Zama, P. C. (2007) Filo Echinodermata. In: Lavrado, H. P. & Viana, M. S. (Eds.), Atlas de invertebrados marinhos da regiao central da Zona Economica Exclusiva brasileira. Parte 1. Museu Nacional do Rio de Janeiro, Serie Livros 25. Museu Nacional do Rio de Janeiro, Rio de Janeiro, pp. 258.","Wagstaff, M. C., Howell, K. L., Bett, B. J., Billett, D. S. M., Brault, S., Stuart, C. T. & Rex, M. A. (2014) β-diversity of deep-sea holothurians and asteroids along a bathymetric gradient (NE Atlantic). Marine Ecology Progress Series, 508, 177 - 185. https: // doi. org / 10.3354 / meps 10877","Fisher, W. K. (1913) Four new genera and fifty-eight new species of starfishes from the Philippine Islands, Celebes, and the Moluccas. Proceedings of the US National Museum, 43, 599 - 648. https: // doi. org / 10.5479 / si. 00963801.43 - 1944.599","Verrill, A. E. (1899) Revision of certain genera and species of starfishes, with descriptions of new forms. Transactions of the Connecticut Academy of Arts and Sciences, 10 (1), 145 - 234. https: // doi. org / 10.5962 / bhl. part. 7031","Fisher, W. K. (1910) New genera of starfishes. Annals and Magazine of Natural History, Series 8, 5 (26), 171 - 173. https: // doi. org / 10.1080 / 00222931008692747","Fisher, W. K. (1906) The starfishes of the Hawaiian Islands. Bulletin of the United States Fish Commission, 23, 987 - 1130.","Mah, C. L. (2018) New genera, species and occurrence records of Goniasteridae (Asteroidea; Echinodermata) from the Indian Ocean. Zootaxa, 4539 (1), 1 - 116. https: // doi. org / 10.11646 / zootaxa. 4539.1.1"]}

Luidia alternata alternata (Say, 1825) Figure 8 Asterias alternata Say, 1825: 144–145. Luidia alternata — Brito 1968: 12–13, pl. 3, fig. 4; Tommasi & Aron 1987: 5; Tommasi et al. 1988: 6; Ventura et al. 2007: 236; Miranda et al. 2012: 9; Miranda 2018: 14, fig. 10B; Torres & Torres 2019: 413. Luidia alternata alternata — Clark & Downey 1992: 8–9, figs. 4b, c, 5d, 6f, 7a–g, q, 8a, b, pl. 1B; Pérez-Ruzafa et al. 1999: 45, fig. 1A; Entrambasaguas 2003: 68–71, fig. 1; Magalhães et al. 2005: 63; Entrambasaguas 2008: 41–43; Gondim et al. 2014: 7, 9, figs. 3a–d; Alitto et al. 2016: 10, figs. 7c–d; Borrero-Peìrez et al. 2019: 4; Cunha et al. 2020: 47, fig. 4F. Material examined (1 spec, 45 mm R). BRAZIL. Bahia, Porto Seguro (16°20’S; 38°14’W)— 40 m, 30.vi.2001, 1 spec (EQMN 2007). Comparative material. U.S.A. Florida, Dry Tortugas, 1 spec, R 170 mm (NHM-UK 1937.5.9.6, neotype). Description (R 45 mm). Flat disc (Fig. 8A). Five robust arms. Abactinal plates covered by paxillae of varied sizes (Fig. 8C). Larger paxillae with a prominent central spine (~ 1.5 mm) and two rows of peripheral spinelets with denticulate tips: first row with few short spinelets; second row with many long, denticulate spinelets. Smaller paxillae with small and robust central spinelets, and one row of long and denticulate peripheral spinelets. Madreporite inconspicuous, hidden by paxillae. Superomarginal plates paxilliform, inconspicuous. Inferomarginal plates elongated and separated by a gap. Marginal region of plates with two long spines arranged horizontally; actinal surface of inferomarginal plates densely covered by flat spines of varying shapes and sizes and three long located centrally. Three large, curved adambulacral spines arranged in a row along center of plate, perpendicular to furrow. One curved, marginal subambulacral spine larger than surrounding spines, but smaller than spines on center of plate (Fig. 8F). Oral plate prominent, with slender and unequal spines (Fig. 8D). Tube feet in two rows, sucking disc lacking (Fig. 8B). Pedicellariae short, with four hyaline valves, inconspicuous and placed adjacent to ambulacral furrow (likely in development). Coloration. No record of coloration of in vivo specimens from this region but live specimens from Trindade Island (Cunha et al. 2020) display the same coloration pattern as the fixed specimen described here: abactinal region with dark scarlet red to brown transverse bands, actinal region yellow beige, spines white. H.L. Clark (1933) noticed that the coloration of this species is not affected by alcohol or other preservation fluids. Distribution. U.S.A. (NC, FL), Gulf of Mexico, Mexico, The The Bahamas, Caribbean Sea, Cuba, Dominican Republic, Honduras, Panama, Colombia, Venezuela, Guyana, Uruguay, Argentina, Cape Verde (Clark & Downey 1992; Pérez-Ruzafa et al. 1999; Entrambasaguas 2003, 2008; Borrero-Peìrez et al. 2019; Mah 2020a). BRAZIL: Amapá, Pará, Paraíba, Alagoas, Bahia, Espírito Santo, Trindade Island, Rio de Janeiro, São Paulo, Santa Catarina, Rio Grande do Sul (Verrill 1915; Brito 1962, 1968; Tommasi 1970; Carrera-Rodríguez & Tommasi 1977; Walenkamp 1979; Tommasi & Aron 1987; Magalhães et al. 2005; Ventura et al. 2007; Miranda et al. 2012; Gondim et al. 2014; Alitto et al. 2016; Miranda 2018; Torres & Torres 2019; Cunha et al. 2020). Depth. 1–200 m (Clark & Downey 1992). Biological notes. Luidia alternata alternata inhabits sandy and muddy bottoms, and it is also found in coral reefs and mangroves, often associated with shells and calcareous algae (Clark & Downey 1992; Hendler et al. 1995; Cunha et al. 2020; present paper). Although this subspecies occurs in shallow water, it is rarely encountered in shallow habitats from Bahia when compared to L. clathrata and L. senegalensis. Its diet is carnivorous, mostly limited to epifaunal organisms including other sea stars such as A. articulatus and L. clathrata (Schwartz & Porter 1977; Ventura et al. 2007). Luidia alternata alternata is classified as “Least Concern” by the Ministry of the Environment (MMA 2018). According to Gurjão & Lotufo (2018), its harvesting in Brazil is currently prohibited. Neotype. NHM-UK 1937.5.9.6. Type locality. Dry Tortugas, FL, U.S.A. Remarks. The only specimen examined has a total length of 60 mm, with an r of 7 mm. Because all arms are regenerating or damaged, we did not include the R/r ratio. Gondim et al. (2014) found pedicellariae with 2–4 valves in the specimens from Paraíba (north of Bahia). The pedicellariae from the specimen described here are short and seem to be in a formation state, but they all have four valves; and the pedicellariae from the neotype are long, formed by 3–4 valves and placed around the mouth and adjacent to the ambulacral furrow. With regards to the number of inferomarginal spines, the neotype (R 170 mm) has 2–3 spines, the specimen described here has two spines, and Gondim et al. ’s (2014) specimens (R 3–20 mm) have 1–2 spines. This variation is most likely ontogenetic, with the number of inferomarginal spines increasing over time up to three. According to H.L. Clark (1933), adult specimens of L. alternata alternata have R 120–130 mm, and the largest specimen known until that date had R 175 mm. Luidia alternata alternata differs from Luidia alternata numidica Koehler, 1911 by having only single central spinelets on many of the smaller paxillae (vs. more numerous central spinelets on many of the smaller paxillae)., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 19-21, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Say, T. (1825) On the species of the Linnaean genus Asterias inhabiting the coast of the United States. Journal of the Academy of Natural Sciences, Philadelphia, 1 (5), 151 - 154.","Brito, I. M. (1968) Asteroiides e equinoiides do Estado da Guanabara e adjace ˆ ncias. Boletim do Museu Nacional, 260, 1 - 51.","Ventura, C. R. R., Verissimo, I., Nobre, C. C. & Zama, P. C. (2007) Filo Echinodermata. In: Lavrado, H. P. & Viana, M. S. (Eds.), Atlas de invertebrados marinhos da regiao central da Zona Economica Exclusiva brasileira. Parte 1. Museu Nacional do Rio de Janeiro, Serie Livros 25. Museu Nacional do Rio de Janeiro, Rio de Janeiro, pp. 258.","Miranda, A. L. S., Lima, M. L. F., Sovierzoski, H. H. & Correia, M. D. (2012) Inventory of the Echinodermata collection from the Universidade Federal de Alagoas. Biota Neotropica, 12 (2), 135 - 146. https: // doi. org / 10.1590 / S 1676 - 06032012000200014","Miranda, A. P. S. (2018) Diversidade e distribuicao de equinodermos da Plataforma Continental Amazonica. Unpublished Honors Thesis, Universidade Federal do Para, Belem, 62 pp.","Torres, V. S. & Torres, F. S. S. (2019) Inventario preliminar dos invertebrados marinhos observados na praia da Pinheira, Palhoca, SC, Brasil. UNISANTA Bioscience, 8 (4), 408 - 422.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Perez-Ruzafa, A., Entrambasaguas, L. & Bacallado, J. J. (1999) Fauna de equinodermos (Echinodermata) de los fondos rocosos infralitorales del archipielago de Cabo Verde. Revista de la Academia Canaria de Ciencias, 11 (3 - 4), 43 - 62.","Entrambasaguas, L. (2003) Estudio de la fauna de equinodermos del archipielago de Cabo Verde: escalas de variabilidad espacial y factores que explican su distribucion. Tesis de licenciatura no publicada. Universidad de Murcia, Murcia. [unknown pagination]","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 9 (1), 61 - 65. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Entrambasaguas, L. (2008) Estudio fauniistico y ecoloigico de los equinodermos del archipieilago de Cabo Verde. Tesis doctoral no publicada, Universidad de Murcia, Murcia, 315 pp.","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Alitto, R. A. S., Bueno, M. L., Domenico, M. & Borges, M. (2016) Annotated checklist of Echinoderms from Araca Bay, South- eastern Brazil. Check List, 12 (1), 1 - 15.","Cunha, R., Tavares, M. & Mendonca, J. B. (2020) Asteroidea (Echinodermata) from shallow-waters of the remote oceanic archipelago Trindade and Martin Vaz, southeastern Atlantic, with taxonomic and zoogeographical notes. Zootaxa, 4742 (1), 31 - 56. https: // doi. org / 10.11646 / zootaxa. 4742.1.2","Clark, H. L. (1933) A handbook of the littoral echinoderms of Porto Rico and the other West Indian Islands. Scientific Survey of Puerto Rico, 16, 1 - 147.","Mah, C. L. (2020 a) World Asteroidea Database. Available from: http: // www. marinespecies. org / asteroidea / (accessed 30 April 2020)","Verrill, A. E. (1915) Report on the Starfishes of the West Indies, Florida, and Brazil, Including Those Obtained by the Bahama Expedition from the University of Iowa in 1893. Bulletin of the State of University of Iowa, 7, 1 - 232, 29 pls. https: // doi. org / 10.5962 / bhl. title. 12035","Brito, I. M. (1962) Ensaio de catailogo dos equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zooloigicos, 13, 1 - 10.","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Carrera-Rodriguez, C. J. & Tommasi, L. R. (1977) Asteroidea de la plataforma continental de Rio Grande do Sul (Brasil), coleccionados durante los viajes del N / Oc. \" Prof. W. Besnard \" para el proyecto Rio Grande do Sul. Boletim do Instituto Oceanografico da Universidade de Sao Paulo, 26 (1), 51 - 130. https: // doi. org / 10.1590 / S 0373 - 55241977000100004","Walenkamp, J. H. C. (1979) Asteroidea (Echinodermata) from the Guyana shelf. Zoologische Verhandelingen, 170, 1 - 97.","Hendler, G., Muller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea Stars, Sea Urchins and Allies: Echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp.","Schwartz, F. J. & Porter, H. J. 1977. Fishes, macroinvertebrates, and their ecological interrelationships with a calico scallop bed off North Carolina. Fishery Bulletin, 75 (2), 427 - 446.","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp.","Gurjao, L. M. & Lotufo, T. M. C. (2018) Native species exploited by marine aquarium trade in Brazil. Biota Neotropica, 18 (3), e 20170387. https: // doi. org / 10.1590 / 1676 - 0611 - bn- 2017 - 0387"]}

Narcissia trigonaria Sladen, 1889 Figures 24–25 Narcissia trigonaria Sladen, 1889: 414, pl. 65, figs. 5–8. Narcissia trigonaria — Brito 1960: 5, pl. 1, figs. 4–5; 1962: 3; 1968: 5; Tommasi 1966: 244; Tommasi & Aron 1988: 3; Tommasi et al. 1988: 6; Clark & Downey 1992: 278, fig. 43, pl. 68; Pawson 2007: 54, 58, fig. 3; Magris & Deìstro 2010: 59; Benavides-Serrato et al. 2011: 175; Miranda et al. 2012: 144; Gondim et al. 2014: 35–36, figs. 10f–j; Souto & Martins 2017: 305; Gurjão & Lotufo 2018: 11; Miranda 2018: 14, fig. 10D; Rubio-Polania et al. 2018: 190. Material examined (10 specs, 7–122 mm R). BRAZIL. Bahia (12°44’– 13°00’S; 38°05’– 38°31’W)—off Bahia, 1 spec, R 62 mm (NHM-UK 90.5.7.641, holotype). Busca Vida beach, 23 m, vii.2005, 1 spec, R 8 mm (UFBA 1090); 23 m, ii.2008, 1 spec, R 16 mm (UFBA 929); 32 m, vii.2008, 1 spec, R 7 mm (UFBA 1089). Busca Vida beach, 1.v.1993, 1 spec, R 95 mm (UFBA 521). Ponta de Areia beach, Itaparica Island, 2000, 1 spec, R 75 mm (UFBA 469). Salvador: 38 m, 2003, 1 spec, R 122 mm (UFBA 570); Barra beach, intertidal, 4.iv.1994, 2 specs, R 70–82 mm (UFBA 42); Porto da Barra beach, x.2008, 1 spec, R 93 mm (UFBA 962). Comparative material. Narcissia ahearnae: U.S.A. Florida, Cape Canaveral, 137 m, 25.x.1961, 1 spec, R 90 mm (NMNH 9736, paratype). The The Bahamas. Andros Island, Goat Cay, 52 m, S. Abbott coll., 26.ii.1971, 1 spec, R 120 mm (NMNH E12440, paratype). Narcissia canariensis: SPAIN. Canary Islands — Tenerife, 1 spec, R 168 mm (NHMUK 1938.6.23.1, holotype). Narcissia gracilis: MEXICO. Baja California — La Paz Bay, intertidal, 5.v.1976, 1 spec, R 78 mm (CASIZ 35025); Cabo San Lucas, 6.viii.1932, 1 spec, R 43 mm (CASIZ 106137); 57 m, 1.v.1888, 1 spec, R 50 mm (NMNH 38317, holotype). Narcissia trigonaria: GULF OF MEXICO, 53 m, 7.ii.1885, 1 spec, R 50 mm (CASIZ 106143); BRAZIL. São Paulo, 22.ii.1992, 1 spec, R 59 mm (MZUSP 272); 1 spec, R 58 mm (MZUSP 315). Description (R 70–122 mm). Disc small, pyramidal; average R/r 4.3 (Fig. 24A). Arms five, long, distally tapering (Fig. 24B–C), cross-section triangular. Carinal ridge undulating in horizontal and vertical planes, from center of disc to middle of arms (Fig. 24B). Abactinal plates irregular-shaped, of various sizes (Fig. 24D), usually larger along carinal ridge, arranged in irregular series. Abactinal granules round (Fig. 24E). Papulae isolated in groups of 2–3 (Fig. 24E), surrounded by tightly packed granules; papulae absent in distal region of arm and on actinal region. Madreporite small to medium-sized, varied morphology, in interradius (Fig. 24F). Anus evident in center of disc. Superomarginal plates large, mostly confined to lateral surface of arm with only a few proximal plates visible from above. Inferomarginal plates confined to actinal surface. Terminal plates large, squared-shaped, usually bare. Actinal area flat, margins rounded. Actinal plates in four rows, one reaching tip of arm; plates covered by prismatic granules more robust and tightly packed than those of abactinal region (Fig. 24G). Four rows of ambulacral spines (Fig. 24H). Adambulacral row with 4–5 flattened spines with blunt tip; proximal adambulacral spine shortest, twice as wide as others. Three rows of subambulacral spines. First row with four spines shorter than adambulacral spines; proximal spine smallest and prismatic, other spines similar to those of adambulacral row. Second row with four prismatic spines, proximal spine smallest. Third row with 1–3 prismatic spines, smaller than those of second row and similar to actinal granules. Oral plates with six pairs of large and thick triangular spines, constricted at the base (Fig. 24I). Tube feet in two rows, sucking disc present. Pedicellariae absent. Ontogenetic variation (R 7–62 mm). Average R/r 3.0 (Fig. 25A). Compared to the body size, the terminal plate is largest in small specimens, forming the tip of the arm (Fig. 25B). Following are additional differences between the smallest (R 7–8 mm), the middle-sized (R 16 mm) and the largest (R 62 mm) specimens examined here: the smallest and the middle-sized (R 7–16 mm) specimens have a flat body, while the largest (R 62 mm) specimen has already a disc and carinal ridge elevated; in R 7–16 mm, all superomarginal plates are visible from above (Fig. 25D), but only the proximal plates are seen from above in R 62 mm; R 7–16 mm have single papulae (Fig. 25C–E), but R 62 mm has 1–2 papulae per group; in the R 7–8 mm, the carinal ridge is not undulating and all granules are rounded, while in R 16–62 mm, the carinal ridge is slightly undulating (Fig. 25B), the abactinal granules are irregular-shaped (Fig. 25C) and the actinal granules are prismatic; the actinal plates form one row in R 7–8 mm, two rows in R 16 mm and three rows in R 62 mm; R 7–8 mm have three adambulacral spines (two large and one small), three spines in the first subambulacral row and four spines in the second subambulacral row (third subambulacral row absent or mixed with second row), while R 16 mm has four adambulacral spines (three large and one small), four spines in the first subambulacral row and 5–7 spines in the second and third subambulacral rows (Fig. 25F) (separation between rows is not clear), and the ambulacral rows in R 62 mm are fully developed. Coloration. Specimens in vivo have a cream color with yellow to red-rust large irregular blotches. Specimens in ethanol are white to light brown. Distribution. U.S.A. (NC, SC, GA, FL, LA, TX), The Bahamas, Gulf of Mexico, Mexico, Honduras, Martinique, Saint Vincent and the Grenadines, Trinidad and Tobago, Panama, Yucatan, Caribbean, Colombia, Venezuela, Guyana, Suriname, French Guiana; Saint Helena Island (Mortensen 1933; Tommasi 1970; Downey 1973; Walenkamp 1976, 1979; Tommasi & Aron 1988; Clark & Downey 1992; Alvarado et al. 2008; Benavides-Serrato et al. 2011; Rubio-Polania et al. 2018). BRAZIL: Pará, Alagoas, Bahia, Rio de Janeiro and São Paulo (Sladen 1889; Verrill 1915; Brito 1960, 1962; Tommasi 1970; Tommasi & Aron 1988; Miranda et al. 2012; Gondim et al. 2014; Souto & Martins 2017; Miranda 2018; present work). Depth. 0–210 m (Pawson 2007b; present work). Biological notes. In Bahia, N. trigonaria lives in habitats with rocks, corals, calcareous algae and coral rubble, but this species has also been found in muddy substrates in Pará (Miranda 2018). The Brazilian populations of N. trigonaria are affected by pollutants and illegal collection for the aquarium trade (Brites et al. 2008). This species is classified as “Least Concern” by the Ministry of the Environment (MMA 2018) and according to Gurjão & Lotufo (2018), its harvesting in Brazil is currently prohibited. Holotype. NHM-UK 90.5.7.641. Type locality. off Bahia, Brazil. Remarks. The original description of N. trigonaria was based on a R 62 mm specimen and lacked much important information, including on the presence of pedicellariae and the shape of its arm carinal ridge. More than a century after its description, the holotype of N. trigonaria is redescribed for the first time herein. Contrary to the conclusion of Gondim et al. (2014), intraspecific variability observed throughout the geographical distribution (~ 20°N to 24°S) of N. trigonaria suggests that this species is not well-established. For example, although Pawson (2007b) mentioned that pedicellariae in N. trigonaria are abundant, we did not find any pedicellaria on its holotype or in other Brazilian specimens. Specimens from Florida, Gulf of Mexico, Caribbean, Guyana and Suriname, on the other hand, have pedicellariae in abundance (Clark 1921; Downey 1973; Walenkamp 1976; present work); these have two long and thin valves ending in 2–3 small teeth (Downey 1973). Also, the literature only describes N. trigo- naria as having a straight carinal ridge, but our observations show that specimens of N. trigonaria from Brazil and Florida have an undulating carinal ridge, while specimens from the Gulf of Mexico have a straight carinal ridge. N. trigonaria differs from N. canariensis by having four rows of ambulacral spines and pedicellariae lacking or with a long, thin stem (vs. three rows of ambulacral spines and abactinal pedicellariae with a short, thick stem) and from N. gracilis by having abactinal plates irregularly distributed and squared terminal plates (vs. abactinal plates forming regular rows and rounded terminal plates). The intraspecific variation in N. trigonaria led Pawson (2007b) to describe N. ahearnae from specimens previously identified as N. trigonaria from Florida and The The Bahamas. The presence of an undulating carinal ridge, of seven popular pores per square mm, and the rarity or absence of pedicellariae were the traits that he used to distinguish N. ahearnae from N. trigonaria. However, specimens of N. trigonaria from Bahia have an undulating carinal ridge, 3–7 papular pores per square mm and lack pedicellariae. Cunha & Tavares (in review) concluded that N. trigonaria differs from N. ahearnae by having rounded abactinal granules and only the proximal superomarginal plates visible abactinally (vs. pointed abactinal granules and all the superomarginal plates visible abactinally), and noted that, in general, specimens with an undulating carinal ridge have a higher density of pores, possibly because the undulation reduces the distance between the pores. Molecular data of specimens of N. trigonaria and N. ahearnae would help to answer questions about this taxon complex, for example, if N. ahearnae is a valid species and in this case, when these species diverged., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 45-48, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Sladen, W. P. (1889) Asteroidea. Report of the Scientific Results of H. M. S. Challenger, 30, 1 - 893. https: // doi. org / 10.1111 / j. 1096 - 3642.1882. tb 02281. x","Brito, I. M. (1960) Asteroides dos estados do Rio de Janeiro e de Sao Paulo. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 5, 1 - 13.","Tommasi, L. R. (1966) Sobre alguns Equinodermas da regiao do Golfo do Mexico e do Mar das Antilhas. Anales del Instituto de Biologia, 37, 155 - 165.","Tommasi, L. R. & Aron, M. A. (1988) Equinodermos da plataforma continental do sudeste do estado da Bahia. Relatos do Instituto Oceanografico da Universidade de Sao Paulo, 19, 1 - 6.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Miranda, A. L. S., Lima, M. L. F., Sovierzoski, H. H. & Correia, M. D. (2012) Inventory of the Echinodermata collection from the Universidade Federal de Alagoas. Biota Neotropica, 12 (2), 135 - 146. https: // doi. org / 10.1590 / S 1676 - 06032012000200014","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Souto, C. & Martins, L. (2017) Os equinodermos. In: Nunes, J. M. C. & Matos, M. R. B. (Eds.), Litoral Norte da Bahia: caracterizacao ambiental, biodiversidade e conservacao. Vol. 1. EDUFBA, Salvador, pp. 303 - 315.","Gurjao, L. M. & Lotufo, T. M. C. (2018) Native species exploited by marine aquarium trade in Brazil. Biota Neotropica, 18 (3), e 20170387. https: // doi. org / 10.1590 / 1676 - 0611 - bn- 2017 - 0387","Miranda, A. P. S. (2018) Diversidade e distribuicao de equinodermos da Plataforma Continental Amazonica. Unpublished Honors Thesis, Universidade Federal do Para, Belem, 62 pp.","Rubio-Polania, J. C., Torruco-Gomez, D., Gonzalez-Solis, A., Ordaz, J. & Caamal-Jimenez, Y. (2018) Benthic megafauna of outer margins of the continental shelf of Yucatan Peninsula. Regional Studies in Marine Science, 24, 184 - 195. https: // doi. org / 10.1016 / j. rsma. 2018.08.014","Mortensen, T. (1933) The Echinoderms of St. Helena (Asteroidea and Ophiuroidea). Videnskabelige Meddelelser fra den Naturhistoriske Forening i Kjobenhavn, 93, 401 - 472.","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Downey, M. E. (1973) Starfishes from the Caribbean and the Gulf of Mexico. Smithsonian Contributions Zoology, 126, 1 - 158. https: // doi. org / 10.5479 / si. 00810282.126","Walenkamp. J. H. C. (1976) The asteroids of the coastal waters of Surinam. Zoologische Verhandelingen, 147, 1 - 91, 18 pls.","Walenkamp, J. H. C. (1979) Asteroidea (Echinodermata) from the Guyana shelf. Zoologische Verhandelingen, 170, 1 - 97.","Alvarado, J. J., Solis-Marin, F. A. & Ahearn, C. (2008) Equinodermos (Echinodermata) del Caribe Centroamericano. Revista de Biologia Tropical, 56, 37 - 55.","Verrill, A. E. (1915) Report on the Starfishes of the West Indies, Florida, and Brazil, Including Those Obtained by the Bahama Expedition from the University of Iowa in 1893. Bulletin of the State of University of Iowa, 7, 1 - 232, 29 pls. https: // doi. org / 10.5962 / bhl. title. 12035","Brito, I. M. (1962) Ensaio de catailogo dos equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zooloigicos, 13, 1 - 10.","Pawson, D. L. (2007 b) Narcissia ahearnae, a new species of sea star from the Western Atlantic (Echinodermata: Asteroidea: Valvatida). Zootaxa, 1386 (1), 53 - 58. https: // doi. org / 10.11646 / zootaxa. 1668.1.31","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp.","Clark, H. L. (1921) The echinoderm fauna of Torres Strait: Its composition and its origin. Carnegie Institution of Washington Publication, 10, 1 - 223. https: // doi. org / 10.5962 / bhl. title. 14613"]}

Luidia senegalensis (Lamark, 1816) Figures 11–12 Asterias senegalensis Lamarck, 1816: 567. Luidia senegalensis — Tommasi 1958: 9, fig. 1, pl. 2; 1985: 3; Brito 1968: 10–11, fig. 1, pl. 3; Lima-Verde 1969: 10; Nomura & Fausto Filho 1966: 19; Nunes 1975: 183, figs. 3–4; Clark & Downey 1992: 21–22, pl. 4A; Hendler et al. 1995: 69–71, fig. 15; Fernandes et al. 2002: 422; Magalhães et al. 2005: 63; Manso et al. 2008: 185, fig. 8c–e; Lima & Fernandes 2009: 58; Magris & Deìstro 2010: 59, 61; Xavier 2010: 75; Benavides-Serrato et al. 2011: 107–108; Gondim et al. 2014: 14–15, figs. 4e–h, 12a; Nisperuza et al. 2016: 115; Alvarado et al. 2017: S276; Alitto et al. 2016: 10, figs. 7e–f; Sandino et al. 2017: S294; Bueno et al. 2018: 180, fig. 10; Gurjão & Lotufo 2018: 11; Miranda 2018: 14, fig. 10A; Rubio-Polania et al. 2018: 190; Borrero-Peìrez et al. 2019: 4; Torres & Torres 2019: 413; Turra et al. 2019; Magris & Giarrizzo 2020: 3. Material examined (2 specs, 19–146 mm R). BRAZIL. Bahia, Todos os Santos Bay (12°46’– 12°51’S; 38°33’– 38°39’W)— 48 m, 29.iv.1997, 1 spec, R 19 mm (UFBA 592); 3 m, 22.v.1997, 1 spec, R 146 mm (UFBA 496). Description (R 146 mm). Nine long, flattened and slender arms (Fig. 11A–C); R/r 6.1. Abactinal surface covered by paxillae. Disc and central region of arm with small paxillae (Fig. 11D). Paxillae covered by 10–15 small, central granules and a fringe of fine peripheral spinelets. Paxillae in disc interradius are larger than others. Arm margins with two rows of rectangular paxillae (Fig. 11F). Madreporite irregularly shaped, often hidden by paxillae (Fig. 11E). Superomarginal plates paxilliform. Inferomarginal plates elongated and separated by a gap. Marginal region of plates with 1–2 short and pointed spines; below these, a fringe of fine, minute spines. Actinal surface of inferomarginal plates densely covered by flat spines, large in central region and minute in marginal region (Fig. 11G). Four adambulacral spines. Two curved spines, innermost spine smallest; two flattened spines of same size, side by side (Fig. 11H). Oral plate narrow, armed with a tuft of strong, long, slender spines (Fig. 11I). Tube feet in two rows, sucking disc lacking. Pedicellariae absent. Ontogenetic variation (R 19 mm). R/r 3.8 (Fig. 12A–B); 1–3 central granules in paxillae (Fig. 12C). Central paxillae spaced out and not tightly packed as in large specimen (Fig. 12E). Inferomarginal plate with only one spine (Fig. 12G). Three adambulacral spines, the innermost curved. Coloration. Specimen in vivo has greyish abactinal region, yellow beige marginal and actinal regions; specimens in ethanol are faded but generally maintain the same pattern. Distribution. U.S.A. (FL), Gulf of Mexico, Mexico, The The Bahamas, the Caribbean, Cuba, Dominican Republic, Jamaica, Haiti, Puerto Rico, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana (Nisperuza et al. 2016; Alvarado et al. 2017; Sandino et al. 2017; Rubio-Polania et al. 2018; Borrero-Peìrez et al. 2019; Mah 2020a). BRAZIL: Amapá, Pará, Rio Grande do Norte, Paraíba, Pernambuco, Bahia, Rio de Janeiro, São Paulo, Paraná, Santa Catarina (Rathbun 1879; Brito 1960, 1968; Walenkamp 1976; 1979; Clark & Downey 1992; Manso et al. 2008; Gondim et al. 2008; 2014; Magris & Deìstro 2010; Alvarado & Solís-Marín 2013; Alitto et al. 2016; Bueno et al. 2018; Miranda 2018; Torres & Torres 2019; Turra et al. 2019). Depth. 1–64 m (Clark & Downey 1992). Biological notes. Specimens from Bahia live in calm waters, on sandy or muddy bottoms (Manso et al. 2008; present paper), sometimes in regions of low salinity (21 ppt), such as the delta of the Paraguaçu River. Astropecten and Luidia species often live in sympatry, partitioning food resources (Ventura 1991; Queiroz 2006; Fernandez et al. 2017); however, populations from Ilhéus, Bahia, live in allopatry, with L. senegalensis mainly at around 15 m of depth and A. marginatus mainly at around 35 m (Pelaes 2008). None of the specimens of Astropecten and Luidia reported in this paper were collected in the same locality. Separation of resources may also occur within populations of L. senegalensis. Turra et al. (2019) noticed that adult individuals seem to migrate to deeper regions, possibly to use different resources. Interestingly, the specimens reported here have a different pattern, in which the smaller specimen was found in a deeper region than that of the large specimen. Luidia senegalensis is not an abundant species. It has a yearly reproductive cycle in Southeastern Brazil, with spawning happening mainly during the rainy season (Turra et al. 2019). This species predates mainly on bivalves, but also feed on other invertebrates such as polychaetes and crustaceans (Monteiro & Pardo 1994), and it is preyed upon by other species of sea stars and sea shorebirds (Brites et al. 2008). With regards to behavior, Tagliafico et al. (2017) observed specimens from Cubagua Island, Venezuela, moving with only four arms in contact with the substrate, while the other five arms were held upright. Luidia senegalensis is used for medicinal purposes in Brazil to treat asthma, cough and metrorrhagia (Costa-Neto & Marques 2000; Alves & Rosa 2007; Alves & Alves 2011). Tangerina et al. (2018) presented a preliminary description of the chemical composition of specimens from São Paulo, but nothing related to pharmaceutical uses so far. Luidia senegalensis is affected by shrimp fisheries, which collects this species as bycatch (Tangerina et al. 2018). In addition to this threat, Magris & Giarrizzo (2020) estimated that L. senegalensis is one of the most vulnerable species impacted by the oil spill recorded in the Brazilian coast in 2019. This species is classified as “Vulnerable” (baseline data indicates that the population size has been reduced by at least 30%) by the Ministry of the Environment (MMA 2018), and its harvesting in Brazil is currently prohibited (Gurjão & Lotufo 2018). Syntypes. MNHN 2014-651, 2014-655, Muséum National d’Histoire Naturelle, Paris. Type locality. Probably West Indies (Clark & Downey 1992). Remarks. Walenkamp (1979) identified a specimen of L. senegalensis from Guyana with only six arms; Clark & Downey (1992) suggested this specimen could belong to L. barbadensis, but because of the presence of a dark stripe along the central region of the arm (vs. stripes in arms) and the depth that the specimen was collected (32 m vs. above 73 m in L. barbadensis), they confirmed Walenkamp’s identification. This specimen should certainly be re-examined since it is the only record of a L. senegalensis with six arms., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 24-26, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Lamarck, J. B. (1816) Histoire naturelle des animaux sans vertebres. Meline, Bruxelles, 568 pp.","Tommasi, L. R. (1958) Os equinodermos do litoral de Sao Paulo. II. Diadematidae, Schizasteridae, Brissidae, Cidaroidea (Echinoidea) e Asteroidea do Bentos Costeiro. Contribuicoes avulsas do Instituto Oceanografico, 2, 1 - 27, 6 pls.","Brito, I. M. (1968) Asteroiides e equinoiides do Estado da Guanabara e adjace ˆ ncias. Boletim do Museu Nacional, 260, 1 - 51.","Lima-Verde, J. S. (1969) Primeira contribuicao ao inventario dos equinodermas do Nordeste brasileiro. Arquivos de Ciencias do Mar, 9 (1), 9 - 13.","Nomura, H. & Fausto Filho, J. (1966) Shrimp survey in coastal and offshore waters of northeastern Brazil. Arquivos da Estacao de Biologia Marinha da Universidade Federal do Ceara, 6, 15 - 29.","Nunes, T. B. (1975) Echinoderma da Baia de Aratu (Bahia, Brasil). Memories of the second Latin American Symposium of Oceanographic Biology, Universidad de Oriente, 1975, 179 - 189.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Hendler, G., Muller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea Stars, Sea Urchins and Allies: Echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp.","Fernandes, M. L. B., Tommasi, L. R. & Lima, E. J. B. (2002) Filo Echinodermata de Pernambuco. In: Tabarelli, M. & Silva, J. M. C. (Eds.), Diagnostico da Biodiversidade de Pernambuco. Vol. 2. Massangana, Recife, pp. 405 - 427.","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 9 (1), 61 - 65. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Manso, C. L. C., Alves, O. F. S. & Martins, L. R. (2008) Echinodermata da Baia de Todos os Santos e da Baia de Aratu (Bahia, Brasil). Biota Neotropica, 8 (3), 179 - 196. https: // doi. org / 10.1590 / S 1676 - 06032008000300017","Lima, E. J. B. & Fernandes, M. L. B. (2009) Diversidade de equinodermos (Echinodermata) no estado de Pernambuco (Brasil). Revista Brasileira de Zoociencias, 11, 55 - 63.","Xavier, L. A. R. (2010) Inventario dos equinodermos do Estado de Santa Catarina, Brasil. Brazilian Journal of Aquatic Science and Technology, 14, 73 - 78. https: // doi. org / 10.14210 / bjast. v 14 n 2. p 73 - 78","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Nisperuza, P. C., Padilla, C. J. & Quiros, R. J. (2016) Current status of the study of the caribbean cordobes echinoderms, Colombia. Revista Colombiana de Ciencia Animal, 8 (1), 112 - 119. https: // doi. org / 10.24188 / recia. v 8. n 1.2016.234","Alvarado, J. J., Chacon-Monge, J. L., Solis-Marin, F. A., Pineda-Enriquez, T., Caballero-Ochoa, A. A., Rivera, S. S. & Chaves, R. R. (2017) Equinodermos del Museo de Zoologia de la Universidad de Costa Rica. Revista de Biologia Tropical, Universidad de Costa Rica, 65 (1), S 272 - S 287. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31695","Alitto, R. A. S., Bueno, M. L., Domenico, M. & Borges, M. (2016) Annotated checklist of Echinoderms from Araca Bay, South- eastern Brazil. Check List, 12 (1), 1 - 15.","Sandino, O. B., Solis-Marin, F. A., Caballero-Ochoa, A. A., Conejeros-Vargas, C. A., Marquez-Borras, F., Ayala-Aguilera, A. P. & Laguarda-Figueras, A. (2017) Equinodermos de Nicaragua: nuevos registros del Pacifico y Caribe Sur. Revista de Biologia Tropical, 65, 288 - 298. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31696","Bueno, M. L., Alitto, R. A. S., Guilherme P. D. B., Domenico, M. D. & Borges, M. (2018) Guia ilustrado dos Echinodermata da porcao sul do Embaiamento Sul Brasileiro. Pesquisa e Ensino em Ciencias Exatas e da Natureza, 2 (2), 169 - 237. https: // doi. org / 10.29215 / pecen. v 2 i 2.1071","Gurjao, L. M. & Lotufo, T. M. C. (2018) Native species exploited by marine aquarium trade in Brazil. Biota Neotropica, 18 (3), e 20170387. https: // doi. org / 10.1590 / 1676 - 0611 - bn- 2017 - 0387","Miranda, A. P. S. (2018) Diversidade e distribuicao de equinodermos da Plataforma Continental Amazonica. Unpublished Honors Thesis, Universidade Federal do Para, Belem, 62 pp.","Rubio-Polania, J. C., Torruco-Gomez, D., Gonzalez-Solis, A., Ordaz, J. & Caamal-Jimenez, Y. (2018) Benthic megafauna of outer margins of the continental shelf of Yucatan Peninsula. Regional Studies in Marine Science, 24, 184 - 195. https: // doi. org / 10.1016 / j. rsma. 2018.08.014","Torres, V. S. & Torres, F. S. S. (2019) Inventario preliminar dos invertebrados marinhos observados na praia da Pinheira, Palhoca, SC, Brasil. UNISANTA Bioscience, 8 (4), 408 - 422.","Turra, A., Checon, H. H., Fernandez, W. S., Majer, A. P., Delboni, C. G., Denadai, M. R. & Dias, G. M. (2019) The importance of embayed coastal areas for the life cycle of two sympatric starfish species (Echinodermata, Asteroidea). Estuarine, Coastal and Shelf Science, 219, 128 - 138. https: // doi. org / 10.1016 / j. ecss. 2019.01.028","Magris, R. A & Giarrizzo, T. (2020) Mysterious oil spill in the Atlantic Ocean threatens marine biodiversity and local people in Brazil. Marine Pollution Bulletin, 153, 1 - 3. https: // doi. org / 10.1016 / j. marpolbul. 2020.110961","Mah, C. L. (2020 a) World Asteroidea Database. Available from: http: // www. marinespecies. org / asteroidea / (accessed 30 April 2020)","Rathbun, R. (1879) A list of the Brazilian echinoderms, with notes on their distribution. Transactions of the Connecticut Academy of Arts and Sciences, 5 (3), 139 - 158. https: // doi. org / 10.5962 / bhl. title. 16126","Brito, I. M. (1960) Asteroides dos estados do Rio de Janeiro e de Sao Paulo. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 5, 1 - 13.","Walenkamp. J. H. C. (1976) The asteroids of the coastal waters of Surinam. Zoologische Verhandelingen, 147, 1 - 91, 18 pls.","Walenkamp, J. H. C. (1979) Asteroidea (Echinodermata) from the Guyana shelf. Zoologische Verhandelingen, 170, 1 - 97.","Gondim, A., Lacouth, P., Alonso, C. & Manso, C. (2008) Echinodermata da praia do Cabo Branco, JoaTo Pessoa, Paraiiba, Brasil. Biota Neotropica, 8, 151 - 159. https: // doi. org / 10.1590 / S 1676 - 06032008000200016","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Vol. 1. Springer, Berlin, 658 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Ventura, C. R. R. (1991) Distribuicao, abundancia e habito alimentar de Asteroidea (Echinodermata) de fundos inconsolidados da plataforma continental do Cabo Frio, RJ. Unpublished Master Thesis, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, 101 pp.","Queiroz, A. C. M. (2006) Dinamica populacional das estrelas do mar Astropecten brasiliensis Muller & Troschel, 1842, Astropecten cingulatus Sladen, 1889 e Luidia senegalensis (Lamarck, 1816) (Echinodermata: Asteroidea) presentes na fauna acompanhante da pesca do camarao no litoral de Ilheus (Bahia). Unpublished Master Thesis, Universidade Federal da Paraiba, Joao Pessoa, Paraiba, 92 pp.","Fernandez, W. S., Dias, G. M., Majer, A., Delboni, C., Denadai, M. R. & Turra, A. (2017) Resource partitioning between sympatric starfish from tropical unconsolidated substrate: implications for coexistence and top-down control on benthic prey. Estuarine, Coastal and Shelf Science, 196, 141 - 149. https: // doi. org / 10.1016 / j. ecss. 2017.06.039","Pelaes, M. A. L. (2008) Composicao, distribuicao espacial e temporal das especies de Echinodermata na plataforma rasa do sul da Bahia num trecho entre Itacare e Canavieiras. Unpublished Master Thesis, Universidade Estadual de Santa Cruz, Ilheus, Bahia, 97 pp.","Tagliafico, A., Matt, J. N. & Rangel, M. S. (2017) Observations of novel locomotion in the sea star Luidia senegalensis (Lamarck, 1816) (Asteroidea: Paxillosida). Marine and Freshwater Behaviour and Physiology, 50 (3), 1 - 5. https: // doi. org / 10.1080 / 10236244.2017.1353883","Costa-Neto, E. M. & Marques, J. G. W. (2000) Faunistic resources used as medicines by artisanal fishermen from Siribinha Beach, State of Bahia, Brazil. Journal of Ethnobiology, 20 (1), 93 - 109.","Alves, R. R. N. & Rosa, I. L. (2007) Biodiversity, traditional medicine and public health: where do they meet? Journal of Ethnobiology and Ethnomedicine, 3, 1 - 9. https: // doi. org / 10.1186 / 1746 - 4269 - 3 - 14","Alves R. R. & Alves H. N. (2011) The faunal drugstore: animal-based remedies used in traditional medicines in Latin America. Journal of Ethnobiology and Ethnomedicine, 10, 1746 - 4269. https: // doi. org / 10.1186 / 1746 - 4269 - 7 - 9","Tangerina, M. M. P., Cesario, J. P., Pereira, G. R. R., Costa, T. M., Valenti, W. C. & Vilegas, W. (2018) Chemical Profile of the Sulphated Saponins from the Starfish Luidia senegalensis Collected as by-Catch Fauna in Brazilian Coast. Natural Products and Bioprospecting, 8 (2), 83 - 89. https: // doi. org / 10.1007 / s 13659 - 018 - 0153 - 2","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp."]}

Othilia echinophora (Lamarck, 1816) Figures 15–16 Asterias echinophora Lamarck, 1816: 560. Echinaster echinophorus — Tommasi & Aron 1988: 3; Fernandes et al. 2002: 422; Gondim et al. 2008: 155, fig. 3a; Alves et al. 2012: 758; Miranda et al. 2012: 144; Patrizzi & Dobrovolski 2018: 182. Echinaster (Othilia) echinophorus — Clark & Downey 1992: 367–371, pls. 89A, B, F, 90F–H; Hendler et al. 1995: 84–85, fig. 27; Hopkins et al. 2003: 100–101, figs. 7–9; Magalhães et al. 2005: 63; Brites et al. 2008: 182–183; Lima & Fernandes 2009: 59; Magris & Deìstro 2010: 59; Xavier 2010: 75; Gondim et al. 2011: 6, fig. 3e; Gondim et al. 2014: 40, figs. 11e–j, 12e–f; Lopes et al. 2016; Sandino et al. 2017: S294. Othilia echinophoru s— Souto & Martins 2017: 304, fig. 1B. Material examined (40 specs, 25–55 mm R). BRAZIL. Bahia (12°45’– 13°00’S; 38°21’– 38°45’W)— Salvador: Barra beach, 6.viii.2004, 1 spec, R 25 mm (UFBA 30); Ribeira beach, intertidal, 20.vi.2005, 2 specs, R 28–30 mm (UFBA 35). Cairu de Salinas, Salinas das Margaridas, intertidal, 26.viii.2007, 3 specs, R 34–40 mm (UFBA 588). Itaparica: Medo Island, intertidal, 29.v.1994, 1 spec, R 45 mm (UFBA 190). Salvador: Frade Island, Nossa Senhora beach, 3 m, 17.x.2008, 6 specs, R 43–55 mm (UFBA 673); Itapuã beach, intertidal, 11.ii.1993, 1 spec, R 40 mm (UFBA 472); Penha beach, 3 m, 10.iv.2008, 24 specs, R 33–54 mm (UFBA 667); Ribeira beach, intertidal, 5.vi.2004, 1 spec, R 38 mm (UFBA 36); intertidal, 5.xii.2004, 2 specs, R 38–43 mm (UFBA 29). Description (R 34–55 mm). Disc small, average R/r 4.0. Five short and robust arms, tapering distally, with 7–9 rows of long and robust spines (ca. 1.7 mm) with or without a mammiform base (Fig. 15A–B). Abactinal region covered by many papulae and glandular cells; skin not very thick but obscuring plating. Five primary plates on disc, each with a large spine forming a pentagon; one central spine (Fig. 15C). Anus near central spine, surrounded by 4–5 robust spinelets. Madreporite flat, peripherally armed with spinelets. Superomarginal spines larger than abactinal ones (Fig. 15E). Inferomarginal plates with one (sometimes two) spine each, forming a row. Abactinal and marginal spines sharp, straight or slightly curved, tapering. Actinal plates absent. Terminal plate robust, slightly curved at arm tip, and with five large, unequal spines. Actinal region with several open pores (one row plus scattered pores). Two unequal, curved adambulacral spines. Two equal subambulacral spines forming a V along the furrow (Fig. 15F). Interradial region naked, with a prominent, central spine. Oral spines completely covering mouth opening (Fig. 15D). Tube feet in two rows, sucking disc present. Pedicellariae absent. Ontogenetic variation (R 25–30 mm). Average R/r 3.6 (Fig. 16A–B). Spines proportionally larger and fewer (7 rows) than in large specimen, arms constricted at the base. Two unequal subambulacral spines, outer spine larger than inner (Fig. 16E). Coloration. Specimens in vivo are dark red. Specimens in ethanol are brown, dark brown or beige. Distribution. U.S.A. (FL), Gulf of Mexico, Mexico, The The Bahamas, Caribbean Sea, Cuba, Jamaica, Dominican Republic, Puerto Rico, Nicaragua, Venezuela, Guyana, French Guiana (Sandino et al. 2017; Mah 2020a). BRAZIL: Amapá, Pará, Ceará, Rio Grande do Norte, Paraíba, Pernambuco, Alagoas, Bahia, Espírito Santo, Rio de Janeiro (Lamarck 1816; Rathbun 1879; Verrill 1915; Bernasconi 1958; Walenkamp 1979; Ávila-Pires 1983; Tommasi & Aron 1988; Clark & Downey 1992; Magris & Deìstro 2010; Miranda et al. 2012; Alvarado & Solís-Marín 2013; Gondim et al. 2014; Souto & Martins 2017). Depth. Intertidal to 55(65?) m (Clark & Downey 1992). Biological notes. Othilia echinophora is a relatively abundant species found often in hard substrate, sometimes in sympatry with O. brasiliensis. In the southeast of the state eight specimens were found in fine calcareous sand with calcareous rocks (Tommasi & Aron 1988). Some populations live in regions of low salinity, such as the delta of the Paraguaçu River. Specimens from Bahia are often found associated with corals (Souto & Martins 2017), above the sediment, and in rock crevices (Alves & Cerqueira 2000). Spongivory by Caribbean specimens have been reported (Waddell & Pawlik 2000). Othilia echinophora has been used in the treatment of asthma (Costa-Neto 1999; Alves et al. 2009; Alves & Dias 2010) and its extracts contain low toxicity compounds that can be used to treat cutaneous leishmaniosis (Parra et al. 2010). In northeastern Brazil, this species is collected for the aquarium trade (Martins et al. 2012). Othilia echinophora is classified as “Least Concern” by the Ministry of the Environment (MMA 2018). According to Gurjão & Lotufo (2018), its harvesting in Brazil is currently prohibited. Lectotype. MNHN-IE-2014072 (Ec As 1976), Muséum National d’Histoire Naturelle, Paris. Type Location. The type locality cited by Lamarck (1816, p. 560) is “ Virginie ”, which would refer to the state of Virginia, U.S.A.; however, O. echinophora has only been found up to Florida. Clark & Downey (1992) deemed unlikely that Lamarck’s assignment was correct and suggested that the type locality is most likely in northeastern Brazil. We are not aware of any coastal Brazilian locality with this name. Another possibility is that the specimens at the MNHN do not belong to the type series, as the label indicates uncertainty on their status (i.e. “ types supposés”) (but see Perrier [1875, p. 101]. Taxonomic remarks of the genus Othilia Here we used the characteristic suggested by Hopkins et al. (2003), i.e. presence of a row of closed pores between the adambulacral and the inferomarginal plates in O. brasiliensis versus the presence of open pores in O. echinophora, to distinguish these species. A.M. Clark (1987) distinguished O. brasiliensis from O. echinophora and O. guyanensis by the blunt tip of the abactinal spines (vs. pointed tip) and the presence of intermarginal plates beyond half of the arm length (vs. plates restricted to less than half of the arm length). In our specimens, the tip of the spines in O. brasiliensis are pointier than the tip of the spines in O. echinophora. Hopkins et al. (2003) used three characteristics to diagnose the Brazilian O. guyanensis, but molecular data did not support its separation from O. brasiliensis. As a result, Lopes et al. (2016) suggested that these two species should be synonymized. Their conclusion, however, is premature because their analysis did not include specimens from the type locality of O. guyanensis and given the similarity among the species of the genus Othilia, the specimens included could have been misidentified. In the diagnostic table presented by A.M. Clark (1987), the best character used to distinguish O. guyanensis from O. brasiliensis and O. echinophora is the flat to slightly concave madreporite (vs. convex). Discussing the status of O. guyanensis is beyond the scope of this study, but similar to Gondim et al. (2014), we have not found specimens unambiguously identified as O. guyanensis in northeastern Brazil. Order Valvatida Perrier, 1884, Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 30-33, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Lamarck, J. B. (1816) Histoire naturelle des animaux sans vertebres. Meline, Bruxelles, 568 pp.","Tommasi, L. R. & Aron, M. A. (1988) Equinodermos da plataforma continental do sudeste do estado da Bahia. Relatos do Instituto Oceanografico da Universidade de Sao Paulo, 19, 1 - 6.","Fernandes, M. L. B., Tommasi, L. R. & Lima, E. J. B. (2002) Filo Echinodermata de Pernambuco. In: Tabarelli, M. & Silva, J. M. C. (Eds.), Diagnostico da Biodiversidade de Pernambuco. Vol. 2. Massangana, Recife, pp. 405 - 427.","Gondim, A., Lacouth, P., Alonso, C. & Manso, C. (2008) Echinodermata da praia do Cabo Branco, JoaTo Pessoa, Paraiiba, Brasil. Biota Neotropica, 8, 151 - 159. https: // doi. org / 10.1590 / S 1676 - 06032008000200016","Alves, R. R. N., Rosa, I. L., Neto, N. L. & Voeks, R. (2012) Animals for the Gods: Magical and Religious Faunal Use and Trade in Brazil. Human Ecology, 40, 751 - 780. https: // doi. org / 10.1007 / s 10745 - 012 - 9516 - 1","Miranda, A. L. S., Lima, M. L. F., Sovierzoski, H. H. & Correia, M. D. (2012) Inventory of the Echinodermata collection from the Universidade Federal de Alagoas. Biota Neotropica, 12 (2), 135 - 146. https: // doi. org / 10.1590 / S 1676 - 06032012000200014","Patrizzi, N. & Dobrovolski, R. (2018) Integrating climate change and human impacts into marine spatial planning: a case study of threatened starfish species in Brazil. Ocean & Coastal Management, 161, 177 - 188. https: // doi. org / 10.1016 / j. ocecoaman. 2018.05.003","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Hendler, G., Muller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea Stars, Sea Urchins and Allies: Echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp.","Hopkins, T. S., Fontanella, F. M. & Ventura, C. R. R. (2003) Morphological diagnosis of three Brazilian starfishes of the genus Echinaster (Othilia). In: Feral, J. P. & David, B. (Eds.), Proceedings of the Sixth European Conference on Echinoderm Research, Banyuls-Sur-Mer, France, 3 - 7 September 2001. Balkema, Rotterdam, pp. 97 - 103.","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 9 (1), 61 - 65. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Lima, E. J. B. & Fernandes, M. L. B. (2009) Diversidade de equinodermos (Echinodermata) no estado de Pernambuco (Brasil). Revista Brasileira de Zoociencias, 11, 55 - 63.","Xavier, L. A. R. (2010) Inventario dos equinodermos do Estado de Santa Catarina, Brasil. Brazilian Journal of Aquatic Science and Technology, 14, 73 - 78. https: // doi. org / 10.14210 / bjast. v 14 n 2. p 73 - 78","Gondim, A. I., Dias, T. L. P., Campos, F. F., Alonso, C. & Christoffersen, M. L. (2011) Macrofauna bentica do Parque Estadual Marinho de Areia Vermelha, Cabedelo, Paraiba, Brasil. Biota Neotropica, 11 (2), 1 - 12. https: // doi. org / 10.1590 / S 1676 - 06032011000200009","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Sandino, O. B., Solis-Marin, F. A., Caballero-Ochoa, A. A., Conejeros-Vargas, C. A., Marquez-Borras, F., Ayala-Aguilera, A. P. & Laguarda-Figueras, A. (2017) Equinodermos de Nicaragua: nuevos registros del Pacifico y Caribe Sur. Revista de Biologia Tropical, 65, 288 - 298. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31696","Souto, C. & Martins, L. (2017) Os equinodermos. In: Nunes, J. M. C. & Matos, M. R. B. (Eds.), Litoral Norte da Bahia: caracterizacao ambiental, biodiversidade e conservacao. Vol. 1. EDUFBA, Salvador, pp. 303 - 315.","Mah, C. L. (2020 a) World Asteroidea Database. Available from: http: // www. marinespecies. org / asteroidea / (accessed 30 April 2020)","Rathbun, R. (1879) A list of the Brazilian echinoderms, with notes on their distribution. Transactions of the Connecticut Academy of Arts and Sciences, 5 (3), 139 - 158. https: // doi. org / 10.5962 / bhl. title. 16126","Verrill, A. E. (1915) Report on the Starfishes of the West Indies, Florida, and Brazil, Including Those Obtained by the Bahama Expedition from the University of Iowa in 1893. Bulletin of the State of University of Iowa, 7, 1 - 232, 29 pls. https: // doi. org / 10.5962 / bhl. title. 12035","Bernasconi, I. (1958) Asteroideos de la coleccion del Instituto Oceanografico de la Universidad de San Pablo. Boletim do Instituto Oceanografico, 9, 13 - 19. https: // doi. org / 10.1590 / S 0373 - 55241958000100002","Walenkamp, J. H. C. (1979) Asteroidea (Echinodermata) from the Guyana shelf. Zoologische Verhandelingen, 170, 1 - 97.","Avila-Pires, T. C. S. A. (1983) Contribuicao ao estudo do genero Echinaster Muller & Troschel, 1840 (Echinodermata, Asteroidea) no litoral brasileiro. Anais da Academia Brasileira de Ciencias, 55 (4), 431 - 448.","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Vol. 1. Springer, Berlin, 658 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Alves, O. F. S. & Cerqueira, W. R. P. (2000) Echinodermata das praias de Salvador (Bahia, Brasil). Revista Brasileira de Zoologia, 17 (2), 543 - 553. https: // doi. org / 10.1590 / S 0101 - 81752000000200024","Waddell, B. & Pawlik, J. (2000) Defenses of Caribbean sponges against invertebrate predators: II. Assays with sea stars. Marine Ecology Progress Series, 195, 133 - 144. https: // doi. org / 10.3354 / meps 195133","Costa-Neto, E. M. (1999) Healing with animals in Feira de Santana City, Bahia, Brazil. Journal of Ethnopharmacology, 65, 225 - 230. https: // doi. org / 10.1016 / S 0378 - 8741 (98) 00158 - 5","Alves, R. N., Neto, N. L., Brooks, S. E. & Albuquerque, U. P. (2009) Commercialization of animal-derived remedies as complementary medicine in the semi-arid region of Northeastern Brazil. Journal of Ethnopharmacology, 124, 600 - 608. https: // doi. org / 10.1016 / j. jep. 2009.04.049","Alves, R. N. & Dias, T. L. P. (2010) Uso de invertebrados na medicina popular no Brasil e suas implicacoes para conservacao. Tropical Conservation Science, 3 (2), 159 - 174. https: // doi. org / 10.1177 / 194008291000300204","Parra, M. G., Fidalgo, L. M., Martinez, J. M., Alvarez, A. M. & Iglesias, O. V. (2010) Leishmanicidal activity of Echinaster (Othilia) echinophorus crude extract. Revista do Instituto de Medicina Tropical de Sao Paulo, 52, 89 - 93. https: // doi. org / 10.1590 / S 0036 - 46652010000200005","Martins, L., Souto, C., Magalhaes, W. F., Alves, O. F. S., Rosa, I. L. & Sampaio, C. L. S. (2012) Echinoderm harvesting in Todosos-Santos Bay, Bahia State, Brazil: the aquarium trade. Sitientibus serie Ciencias Biologicas, 12 (1), 53 - 59. https: // doi. org / 10.13102 / scb 123","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp.","Gurjao, L. M. & Lotufo, T. M. C. (2018) Native species exploited by marine aquarium trade in Brazil. Biota Neotropica, 18 (3), e 20170387. https: // doi. org / 10.1590 / 1676 - 0611 - bn- 2017 - 0387","Perrier, E. (1875) Revision de la collection de stellerides du Museum d'Histoire Naturelle de Paris. Archives de Zoologie Experimentale et Generate, 4, 265 - 450.","Clark, A. M. (1987) Notes on Atlantic and other Asteroidea Echinasteridae. Bulletin of the British Museum of Natural History, 53, 65 - 78.","Perrier, E. (1884) Memoire sur les etoiles de mer recueillies dans la mer des Antilles et le golfe du Mexique: durant les expeditions de dragace faites sous la direction de M. Alexandre Agassiz. Archives Museum National d'histoire Naturelle Paris, 6 (2), 127 - 276. https: // doi. org / 10.5962 / bhl. title. 82184"]}

Genus Asterinides Verrill, 1913 Type species. Asterinides folium (Lütken, 1860) (type by original designation). Remarks. Asterinides includes four valid species, all of them originally described in different genera: A. folium, A. hartmeyeri (Döderlein, 1910), A. pilosa (Perrier, 1881) and A. pompom (Clark, 1983). All of these species occur in the Western Atlantic Ocean, especially in the Caribbean Sea, and A. folium is the only one found in the southern hemisphere., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on page 33, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Lutken, C. F. (1860) Bidrag til kundskab om de ved kysterne af Mellenog Sydde ved kysterne af Mellenog Sydundskab. Videnskabelige Meddelelser fra den Naturhistoriske Forening i Kjobenhavn, 1859, 25 - 97.","Doderlein, L. & Hartmeyer, R. (1910) Westindische Seeigel und Seesterne. Zoologische Jahrbucher Supplement, 11 (2), 145 - 156.","Perrier, E. (1881) Description sommaire des especes nouvelles d'Asteries. Museum of Comparative Zoology Bulletin, 9, 1 - 31.","Clark, A. M. (1983) Notes on Atlantic and other Asteroidea. 3. The families Ganeriidae and Asterinidae, with description of a new asterinid genus. Bulletin of the British Museum of Natural History, 45 (7), 359 - 380. https: // doi. org / 10.5962 / bhl. part. 28006"]}

Genus Nymphaster Sladen, 1889 Type species. Nymphaster arenatus (Perrier, 1881) (type by subsequent designation by Fisher, 1917). Remarks. The genus Nymphaster includes 16 species and only N. arenatus occurs in the Atlantic Ocean (Mah, 2020b). Halpern (1970b) identified two species from the Atlantic Ocean, but Clark & Downey (1992) synonymized N. subspinosus (Perrier, 1881) with N. arenatus., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on page 36, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Sladen, W. P. (1889) Asteroidea. Report of the Scientific Results of H. M. S. Challenger, 30, 1 - 893. https: // doi. org / 10.1111 / j. 1096 - 3642.1882. tb 02281. x","Perrier, E. (1881) Description sommaire des especes nouvelles d'Asteries. Museum of Comparative Zoology Bulletin, 9, 1 - 31.","Fisher, W. K. (1917) Notes on Asteroidea. Annals and Magazine of Natural History, Series 8, 20 (116), 166 - 172. https: // doi. org / 10.1080 / 00222931709486985","Mah, C. L. (2020 b) New species, occurrence records and observations of predation by deep-sea Asteroidea (Echinodermata) from the North Atlantic by NOAA ship Okeanos Explorer. Zootaxa, 4766 (2), 201 - 260. https: // doi. org / 10.11646 / zootaxa. 4766.2.1","Halpern, J. A. (1970 b) Goniasteridae (Echinodermata: Asteroidea) of the straits of Florida. Bulletin of Marine Science, 20 (1), 193 - 286.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659"]}

Family Astropectinidae Gray, 1840 Genus Astropecten Gray, 1840 Type species. Astropecten aranciacus (Linnaeus, 1758) (by subsequent designation by Fisher, 1908). Remarks. With more than 105 species described, Astropecten is one of the most diverse sea star genera (Zulliger & Lessios 2010). Of these species, 25 occur in the Atlantic Ocean and eight occur in Brazil: Astropecten acutiradiatus, A. alligator Perrier, 1881, A. antillensis, A. articulatus (Say, 1825), A. brasiliensis brasiliensis, A. cingulatus, A. duplicatus Gray, 1840 and A. marginatus. In the molecular phylogeny of the genus Astropecten proposed by Zulliger & Lessios (2010), the West Atlantic species form a clade with the East Pacific species, most likely separated by the Isthmus of Panama as suggested by Tortonese (1956). Sister to this clade is a clade composed of East Atlantic and Mediterranean species. Diet of this genus is generally composed of bivalves, gastropods and small crustaceans (Christensen 1970; Ribi et al. 1977; Ventura 1991). Species of the genus Astropecten are used for medicinal purposes in the northeastern Brazil, where the tea of the whole specimen is drunk to treat asthma and cough, to prevent abortion and to reduce menstrual bleeding (Alves & Rosa 2006)., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 7-8, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Gray, J. E. (1840) A synopsis of the genera and species of the class Hypostoma (Asterias Linnaeus). Annals of the Magazine of Natural History, Series 1, 6 (37), 175 - 184 + 275 - 290. https: // doi. org / 10.1080 / 03745484009443296","Linnaeus, C. (1758) Systema Naturae. Impensis Direct, Laurentii Salvii, Holmiae, 824 pp.","Fisher, W. K. (1908) Necessary changes in the nomenclature of starfishes. Smithsonian Miscellaneous collections (Quaterly), 52, 87 - 93.","Zulliger, D. E. & Lessios, H. A. (2010) Phylogenetic relationships in the genus Astropecten Gray (Paxillosida: Astropectinidae) on a global scale: molecular evidence for morphological convergence, species-complexes and possible cryptic speciation. Zootaxa, 2504 (1), 1 - 19. https: // doi. org / 10.11646 / zootaxa. 2504.1.1","Perrier, E. (1881) Description sommaire des especes nouvelles d'Asteries. Museum of Comparative Zoology Bulletin, 9, 1 - 31.","Say, T. (1825) On the species of the Linnaean genus Asterias inhabiting the coast of the United States. Journal of the Academy of Natural Sciences, Philadelphia, 1 (5), 151 - 154.","Tortonese, E. (1956) Su alcune specie di Astropectinidae, con descrizone di una nuovo Astropecten (Asteroidea). Annali Museo Storia Naturale Genova, 68, 319 - 334.","Christensen, A. M. (1970) Feeding biology of the sea star Astropecten irregularis Pennant. Ophelia, 8, 1 - 134. https: // doi. org / 10.1080 / 00785326.1970.10429554","Ribi, G., Scharer, R. & Ochsner, P. (1977) Stomach contents and size frequency distribution of two co-existing seastar species Astropecten aranciacus and A. bispinosus with reference to competition. Marine Biology, 43, 181 - 185. https: // doi. org / 10.1007 / BF 00391266","Ventura, C. R. R. (1991) Distribuicao, abundancia e habito alimentar de Asteroidea (Echinodermata) de fundos inconsolidados da plataforma continental do Cabo Frio, RJ. Unpublished Master Thesis, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, 101 pp."]}

Genus Oreaster Müller & Troschel, 1842 Type species. Oreaster reticulatus (Linnaeus, 1758) (type by original designation by Döderlein, 1916). Remarks. The genus Oreaster is composed of only two species: O. clavatus Müller & Troschel, 1842 (East Atlantic) and O. reticulatus (West Atlantic). The molecular data published by Janies et al. (2019) indicated that there was a new species of Oreaster from the Gulf of Mexico. However, Collins et al. (2020) concluded that the specimens analyzed by Janies et al. (2019) are more closely related to Valvaster striatus (Lamarck, 1816), a species considered endemic to the Indo-Pacific., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on page 51, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Muller, J. & Troschel, F. H. (1842) System der Asteriden. Friedrich Viewig und Sohn, Braunschweig, 134 pp.","Linnaeus, C. (1758) Systema Naturae. Impensis Direct, Laurentii Salvii, Holmiae, 824 pp.","Doderlein, L. (1916) Uber die Gattung Oreaster und Venvandte. Zoologische Jahrbucher, Abteilung fur Systematik, 40, 409 - 440.","Janies, D., Hernandez-Diaz, Y. Q., Solis-Marin, F. A., Lopez, K., Alexandrov, B., Galac, M., Herrera, J., Cobb, J., Ebert, T. A. & Bosch. I. (2019) Discovery of adults linked to cloning oceanic starfish larvae (Oreaster: Asteroidea: Echinodermata). The Biological Bulletin, 236, 174 - 185. https: // doi. org / 10.1086 / 703233","Collin, R., Venera-Ponton, D. E., Paulay, G. & Boyle, M. J. (2020) World Travelers: DNA Barcoding Unmasks the Origin of Cloning Asteroid Larvae from the Caribbean. Biology Bulletin, 239 (2), 73 - 79. https: // doi. org / 10.1086 / 710796","Lamarck, J. B. (1816) Histoire naturelle des animaux sans vertebres. Meline, Bruxelles, 568 pp."]}

Genus Ophidiaster L. Agassiz, 1836 Type species. Ophidiaster ophidianus (Lamarck, 1816) (type by monotypy). Remarks. The genus Ophidiaster is composed of 24 species, and six of them are recorded in the Atlantic Ocean: O. alexandri Verrill, 1915, O. bayeri Clark, 1948, O. bullisi (Downey, 1970), O. guildingi Gray, 1840, O. ophidianus (Lamarck, 1816) and O. reyssi Sibuet, 1977. Only O. alexandri and O. guildingi are recorded in Brazil until this date., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on page 48, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Lamarck, J. B. (1816) Histoire naturelle des animaux sans vertebres. Meline, Bruxelles, 568 pp.","Verrill, A. E. (1915) Report on the Starfishes of the West Indies, Florida, and Brazil, Including Those Obtained by the Bahama Expedition from the University of Iowa in 1893. Bulletin of the State of University of Iowa, 7, 1 - 232, 29 pls. https: // doi. org / 10.5962 / bhl. title. 12035","Clark, A. H. (1948) Two new starfishes and a new brittle star from Florida and Alabama. Proceedings of the Biological Society of Washington, 61, 55 - 66.","Downey, M. E. (1970) Drachmaster bullisi new genus and species of Ophidiasteridae (Echinodermata: Asteroidea) with a key to the Caribbean species of the family. Proceedings of the Biological Society of Washington, 83 (6), 77 - 82.","Gray, J. E. (1840) A synopsis of the genera and species of the class Hypostoma (Asterias Linnaeus). Annals of the Magazine of Natural History, Series 1, 6 (37), 175 - 184 + 275 - 290. https: // doi. org / 10.1080 / 03745484009443296","Sibuet, M. (1977) Ophidiaster reyssi nouvelle espece d'Asteride bathyale de l'ocean Atlantique. Bulletin du Museum national d'Histoire naturelle, 343 (494), 1085 - 1090."]}

Ophidiaster guildingi Gray, 1840 Figures 26–27 Ophidiaster guildingii Gray, 1840: 284. Ophidiaster guildingii — Tommasi & Aron 1988: 3; Hendler et al. 1995: 79, fig. 22; Madeira et al. 2019: 93–94. Ophidiaster guildingi — Clark & Downey 1992: 281, fig. 44c, d, pl. 69A, B; Benavides-Serrato et al. 2011: 176; Sandino et al. 2017: S294; Borrero-Peìrez et al. 2019: 5; Cunha et al. 2020: 41, fig. 6; Mah 2020b: 238, fig. 18A–D. Material examined (3 specs, 7–22 mm R). BRAZIL. Bahia, Abrolhos, Minerva Seamount (17°03’S, 37°37’W)— 69 m, 18.viii.2012, 3 specs, R 7–22 mm (MZUSP 2102). Comparative material. BRAZIL. Espírito Santo: Trindade Island (20°29’– 20°32’S; 29°19’– 29°20’W)— Enseada dos Portugueses, 12 m, 15.vii.2013, 1 spec, R 9 mm (MZUSP 1563); Enseada do Príncipe, Pedra da Garoupa, 10.4 m, 16.vii.2013, 1 spec, R 25 mm (MZUSP 1576); Enseada das Orelhas, 8.9 m, 8.i.2015, 1 spec, R 62 mm (MZUSP 1570). Description of largest specimens (R 17–22 mm). Disc small; average R/r 4.9 (Fig. 26A–B). Five long, cylindrical, constricted at base. Madreporite subcircular (Fig. 26D), evident in interradial area. Anus conspicuous in center of disc, surrounded by 10 spines. Terminal plate round, smaller than arm diameter, with about 8 tubercles (Fig. 26G). Abactinal plates (Fig. 26C) cruciform, imbricated, covered by flattened granules, and arranged in seven regular rows. Papular areas forming regular rows parallel to abactinal plates; with 4–8 pores proximally (Fig. 26C) and 3–5 pores distally. Marginal plates similar to abactinal plates. One row of imbricate actinal plates; actinal papular areas with 4–6 papular pores (Fig. 26H). Each inferomarginal plate connected to two adjacent actinal plates via a rod-like plate. One row of adambulacral spines, two subequal spines per plate (Fig. 26F). One subambulacral spine per plate; spine flattened, tapering, twice as large as adambulacral spine. Space between adambulacral and subambulacral spines (Fig. 26E). Oral spines (Fig. 26E) similar to adambulacral spines. Tube feet in two rows, sucking disc with perforated plates. Pedicellariae absent. Ontogenetic variation (R 7 mm). Average R/r 2.3. The specimen has short and wide arms, not constricted at the base (Fig. 27A–B), and a relatively large terminal plate forming the tip of the arm (Fig. 27G). The number of spines, tubercles, and papular areas are reduced in some areas, the anus is surrounded by 7–9 spines, terminal plates have 4–5 tubercles, papular areas have 1–3 pores in the abactinal region, and 1–2 pores in the actinal region. Also, abactinal plates are round and not imbricated (Fig. 27C), and adambulacral spines equal (Fig. 27E). Coloration. No record of coloration of in vivo specimens from this region, but specimens from Trindade Island are dark brown (Cunha et al. 2020). Juveniles from the Caribbean are purple (Hendler et al. 1995). Specimens in ethanol are pale brown. Distribution. Eastern Atlantic: Cape Verde, Gulf of Guinea, Ascension Island. Western Atlantic: U.S.A. (FL, GA, TX), Bermuda, The Bahamas, Mexico, Cuba, Belize, Turks and Caicos, Anguilla, Dominican Republic, Puerto Rico, Virgin Islands, Trinidad and Tobago, Antilles, Nicaragua, Panama, Colombia (Pawson 1978; Clark & Downey 1992; Entrambasaguas 2003; Sandino et al. 2017; Borrero-Peìrez et al. 2019; Mah 2020b). BRAZIL: Bahia, Trindade Island, Rio Grande do Sul (Tommasi 1970; Carrera-Rodriguez & Tommasi 1977; Tommasi & Aron 1988; Cunha et al. 2020). Depth. 0–445 m (Mah 2020b). Biological notes. In Bahia, this species is found in rubble and coral reef ecosystems, calcareous rocks and gravel (Tommasi & Aron 1988). Although O. guildingi is not widely distributed, it is generally abundant locally. Ophidiaster guildingi is a slow-moving species (Hendler et al. 1995) and can reproduce asexually and regenerate a new individual from a single arm (Haeckel 1878). The specimens from Bahia did not show signs of asexual reproduction. Holotype. The type specimen of O. guildingi has not been found (Clark & Downey 1992). Type locality. St Thomas, Virgin Islands (Clark & Downey 1992). Remarks. This species is generally small, growing only up to R 62 mm; according to H.L. Clark (1933), O. guildingi reaches maturity at R 45 mm. As noted above, the number of pores per papular area varies with ontogeny. Specimens from Trindade Island with R 9 mm have 1–2 pores, with R 25 mm have 3–6 pores and with R 62 mm have 5–12 pores. At R 33 mm, specimens have 5–12 pores (Downey 1973), and the maximum number of pores is 15 (Clark & Downey 1992). Clark & Downey (1992) also noted the rare occurrence of pedicellariae in large specimens, but Cunha et al. (2020) did not find pedicellariae in the specimens from Trindade Island. Ophidiaster guildingi differs from O. alexandri , O. bullisi and O. reyssi by having only one row of actinal plates (vs. 3–5 rows in O. alexandri; two rows in O. bullisi; and four rows in O. reyssi); from O. bayeri by having only one row of subambulacral spines (vs. two rows); from O. ophidianus by having up to 15 pores per papular area (vs. more than 20 pores) and flat and tapering spines subambulacral spines (vs. cylindrical and blunt) (Clark 1921; Clark & Downey 1992; Madeira et al. 2019)., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 48-51, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Gray, J. E. (1840) A synopsis of the genera and species of the class Hypostoma (Asterias Linnaeus). Annals of the Magazine of Natural History, Series 1, 6 (37), 175 - 184 + 275 - 290. https: // doi. org / 10.1080 / 03745484009443296","Tommasi, L. R. & Aron, M. A. (1988) Equinodermos da plataforma continental do sudeste do estado da Bahia. Relatos do Instituto Oceanografico da Universidade de Sao Paulo, 19, 1 - 6.","Hendler, G., Muller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea Stars, Sea Urchins and Allies: Echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp.","Madeira, P., Kroh, A., Cordeiro, R., Frias, D. E., Martins, A. M. & Avila, S. P. (2019) The Echinoderm Fauna of the Azores (NE Atlantic Ocean). Zootaxa, 4639 (1), 1 - 231. https: // doi. org / 10.11646 / zootaxa. 4639.1","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Sandino, O. B., Solis-Marin, F. A., Caballero-Ochoa, A. A., Conejeros-Vargas, C. A., Marquez-Borras, F., Ayala-Aguilera, A. P. & Laguarda-Figueras, A. (2017) Equinodermos de Nicaragua: nuevos registros del Pacifico y Caribe Sur. Revista de Biologia Tropical, 65, 288 - 298. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31696","Cunha, R., Tavares, M. & Mendonca, J. B. (2020) Asteroidea (Echinodermata) from shallow-waters of the remote oceanic archipelago Trindade and Martin Vaz, southeastern Atlantic, with taxonomic and zoogeographical notes. Zootaxa, 4742 (1), 31 - 56. https: // doi. org / 10.11646 / zootaxa. 4742.1.2","Mah, C. L. (2020 b) New species, occurrence records and observations of predation by deep-sea Asteroidea (Echinodermata) from the North Atlantic by NOAA ship Okeanos Explorer. Zootaxa, 4766 (2), 201 - 260. https: // doi. org / 10.11646 / zootaxa. 4766.2.1","Pawson, D. L. (1978) The echinoderm fauna of Ascension Island, South Atlantic Ocean, Smithsonian. Contributions Marine Sciences, 2, 1 - 31. https: // doi. org / 10.5479 / si. 01960768.2.1","Entrambasaguas, L. (2003) Estudio de la fauna de equinodermos del archipielago de Cabo Verde: escalas de variabilidad espacial y factores que explican su distribucion. Tesis de licenciatura no publicada. Universidad de Murcia, Murcia. [unknown pagination]","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Carrera-Rodriguez, C. J. & Tommasi, L. R. (1977) Asteroidea de la plataforma continental de Rio Grande do Sul (Brasil), coleccionados durante los viajes del N / Oc. \" Prof. W. Besnard \" para el proyecto Rio Grande do Sul. Boletim do Instituto Oceanografico da Universidade de Sao Paulo, 26 (1), 51 - 130. https: // doi. org / 10.1590 / S 0373 - 55241977000100004","Haeckel, E. (1878) Die Kometenform der Seesterne und der Generationswechsel der Echinodermum. Zeitschrift fur Wissenschaftliche Zoologie, 30 (Supplement), 424 - 445.","Clark, H. L. (1933) A handbook of the littoral echinoderms of Porto Rico and the other West Indian Islands. Scientific Survey of Puerto Rico, 16, 1 - 147.","Downey, M. E. (1973) Starfishes from the Caribbean and the Gulf of Mexico. Smithsonian Contributions Zoology, 126, 1 - 158. https: // doi. org / 10.5479 / si. 00810282.126","Clark, H. L. (1921) The echinoderm fauna of Torres Strait: Its composition and its origin. Carnegie Institution of Washington Publication, 10, 1 - 223. https: // doi. org / 10.5962 / bhl. title. 14613"]}

Plinthaster dentatus (Perrier, 1884) Figure 21 Pentagonaster dentatus Perrier, 1884: 242–243, pl. 8, fig. 3 (instead of pl. 3, fig. 8 as stated in p. 242). Plinthaster dentatus — Clark & Downey 1992: 260, pl. 61, figs. D–E; Sumida et al. 2001: 26, fig. 9; Entrambasaguas 2008: 55; Benavides-Serrato et al. 2011: 168; Soaréz 2016: 78; Sandino et al. 2017: S294; Serrano et al. 2017; Borrero-Peìrez et al. 2019: 5; Madeira et al. 2019: 90; Mah 2020b: 232, fig. 14D. Material examined (1 spec, 20 mm R). BRAZIL. Bahia, Morro de São Paulo (13°42’S; 38°64’W), 801 m, 19.vi.2000, 1 spec, R 20 mm (EQMN 2346). Comparative material. GRENADA. Off Grenada, 762 m, 28.ii.1879, 1 spec, R 34 mm (MCZ AST 408, lectotype); BRAZIL. Rio de Janeiro, 1227 m, 30.xii.1887, 1 spec, R 15 mm (YPM IZ 9526. EC). Description (R 20 mm). Body stellate, flat, arms short (Fig. 21A–B); R/r 2.0; 7 SM plates; R/SM# 2.86. Abactinal plates polygonal, flat, bare in center and surrounded by one row of granules (Fig. 21C). Some plates with pedicellariae and crystal bodies (Fig. 21D). Plates irregularly arranged in the center of disc, regularly arranged towards the arms and absent at arm tip. Madreporite large (Fig. 21A, D). Terminal plate large, naked, tapering distally (Fig. 21E). Papulae large, single. Marginal plates block-like, with scattered granules (note that plates in specimen analyzed are abraded), forming an ambitus and surrounded by one row of granules; 14 per interradius. Superomarginal plates naked; three distalmost pairs in contact medially (Fig. 21E). Actinal areas large. Inferomarginal plates similar to superomarginal plates, but with some granules along edge. Actinal plates flat, large, covered with regularly spaced, coarse granules (Fig. 21F). Five blunt adambulacral spines. Two rows of subambulacral spines, smaller than adambulacral spines: first row with 3–4 spines, second row with 3–5 spines per plate (Fig. 21G). Oral plates with nine subprismatic spines (Fig. 21H); spines increasing in size toward mouth. Bivalved pedicellariae on actinal plates, near ambulacral furrow. Coloration. No record of coloration of in vivo specimens from this region. Halpern (1970b) mentioned that live specimens have an orange abactinal region and creamy actinal. Specimen in ethanol is beige with brown spots. Distribution. Pacific Ocean: New South Wales, Australia and Kermadec Islands, New Zealand. Eastern Atlantic: Ireland, Bay of Biscay, Spain, Azores, Canary Islands, Morocco, Cape Verde, Liberia, Gulf of Guinea. Western Atlantic: U.S.A. (NC, SC, FL, LA, TX), The Bahamas, Mexico, Cuba, Dominican Republic, Jamaica, Anguilla, Trinidad and Tobago, Honduras, Nicaragua, Panama, Colombia, Venezuela, Uruguay (Clark 1941; Gray et al. 1968; Halpern 1970b; Sibuet 1977; Clark & Downey 1992; Clark & McKnight 2001; Alvarado & Solís-Marín 2013; Soaréz 2016; Sandino et al. 2017; Serrano et al. 2017; Borrero-Peìrez et al. 2019; Madeira et al. 2019; Mah 2020b). BRAZIL: Maranhão, Bahia, Espírito Santo (Halpern 1970b; Ventura et al. 2007; Gondim et al. 2014) and Rio de Janeiro (YPM IZ 9526.EC, not analyzed). Depth. 229–2910 m (Clark & Downey 1992). Biological notes. Plinthaster dentatus is a deep-water species and its biology is not well studied. This species feeds on foraminifera, crinoid arms, corals, encrusting organisms and geodiid sponges (Halpern 1970a; Mah 2020b). Lectotype. MCZ 408, by subsequent designation of Halpern (1970b). Perrier (1884) mentioned three specimens in his paper, two from off Cuba (one of them R 14 mm) and one from off Grenada (MCZ 408; about R 34 mm), but we could not locate the specimens from off Cuba. Type locality. Off Grenada. Remarks. Plinthaster dentatus is a widely distributed species and the intraspecific variation described is remarkable. Unfortunately, most descriptions do not detail which variation was due to ontogeny. Sumida et al. (2001) provided morphological data on specimens from Ireland ranging from R 1.1 to 7.6 mm and concluded that juveniles of P. dentatus have isometric growth. Next, we compare the specimens described by them with our specimen and the specimens described by Perrier (1884; R 14 mm and 34 mm), Halpern (1970b; R 45 mm), Clark & Downey (1992; R 45 mm) and Mah (2018; R 55–70 mm). The number of marginal plates per interradius increases from 2–6 in R Plinthaster dentatus differs from the other species by having 12–20 marginal plates per interradius (vs. 6–8 marginal plates in P. lenaigae and P. untiedtae, and 8–10 in P. ceramoidea) (Mah 2018)., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 40-42, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Perrier, E. (1884) Memoire sur les etoiles de mer recueillies dans la mer des Antilles et le golfe du Mexique: durant les expeditions de dragace faites sous la direction de M. Alexandre Agassiz. Archives Museum National d'histoire Naturelle Paris, 6 (2), 127 - 276. https: // doi. org / 10.5962 / bhl. title. 82184","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Sumida, P. Y. G., Tylerp, A. & Billettd, S. M. (2001) Early juvenile development of deep-sea asteroids of the NE Atlantic Ocean, with notes on juvenile bathymetric distributions. Acta Zoologica, 82, 11 - 40. https: // doi. org / 10.1046 / j. 1463 - 6395.2001.00058. x","Entrambasaguas, L. (2008) Estudio fauniistico y ecoloigico de los equinodermos del archipieilago de Cabo Verde. Tesis doctoral no publicada, Universidad de Murcia, Murcia, 315 pp.","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Sandino, O. B., Solis-Marin, F. A., Caballero-Ochoa, A. A., Conejeros-Vargas, C. A., Marquez-Borras, F., Ayala-Aguilera, A. P. & Laguarda-Figueras, A. (2017) Equinodermos de Nicaragua: nuevos registros del Pacifico y Caribe Sur. Revista de Biologia Tropical, 65, 288 - 298. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31696","Serrano, A., Cartes, J. E., Papiol, V., Punzon, A., Garcia-Alegre, A., Arronte, J. C., Rios, P., Lourido, A., Frutos, I. & Blanco, M. (2017) Epibenthic communities of sedimentary habitats in a NE Atlantic deep seamount (Galicia Bank). Journal of Sea Research, 130, 154 - 165. https: // doi. org / 10.1016 / j. seares. 2017.03.004","Madeira, P., Kroh, A., Cordeiro, R., Frias, D. E., Martins, A. M. & Avila, S. P. (2019) The Echinoderm Fauna of the Azores (NE Atlantic Ocean). Zootaxa, 4639 (1), 1 - 231. https: // doi. org / 10.11646 / zootaxa. 4639.1","Mah, C. L. (2020 b) New species, occurrence records and observations of predation by deep-sea Asteroidea (Echinodermata) from the North Atlantic by NOAA ship Okeanos Explorer. Zootaxa, 4766 (2), 201 - 260. https: // doi. org / 10.11646 / zootaxa. 4766.2.1","Halpern, J. A. (1970 b) Goniasteridae (Echinodermata: Asteroidea) of the straits of Florida. Bulletin of Marine Science, 20 (1), 193 - 286.","Clark, H. L. (1941) Reports on the scientific results of the Atlantis Expedition to the West Indies, under the joint auspices of the University of Havana and Harvard University. The echinoderms (other than holothurians). Memorias de la Sociedad Cubana de Historia Natural Felipe Poey, 15 (1), 1 - 154.","Gray, I. E., Downey, M. E. & Cerame-Vivas, M. J. (1968) Sea-Stars of North Carolina. Fisheries Bulletin, 67 (1), 127 - 163, 40 figs.","Sibuet, M. (1977) Ophidiaster reyssi nouvelle espece d'Asteride bathyale de l'ocean Atlantique. Bulletin du Museum national d'Histoire naturelle, 343 (494), 1085 - 1090.","Clark, H. E. S. & McKnight, D. G. (2001) The Marine Fauna of New Zealand: Echinodermata: Asteroidea (sea-stars), Order Valvatida. NIWA Biodiversity Memoir, Wellington, 270 pp.","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Vol. 1. Springer, Berlin, 658 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Ventura, C. R. R., Verissimo, I., Nobre, C. C. & Zama, P. C. (2007) Filo Echinodermata. In: Lavrado, H. P. & Viana, M. S. (Eds.), Atlas de invertebrados marinhos da regiao central da Zona Economica Exclusiva brasileira. Parte 1. Museu Nacional do Rio de Janeiro, Serie Livros 25. Museu Nacional do Rio de Janeiro, Rio de Janeiro, pp. 258.","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Halpern, J. A. (1970 a) A monographic revision of the goniasterid sea stars of the North Atlantic. Unpublished PhD Dissertation, University of Miami, Florida, 253 pp.","Mah, C. L. (2018) New genera, species and occurrence records of Goniasteridae (Asteroidea; Echinodermata) from the Indian Ocean. Zootaxa, 4539 (1), 1 - 116. https: // doi. org / 10.11646 / zootaxa. 4539.1.1","Downey, M. E. (1973) Starfishes from the Caribbean and the Gulf of Mexico. Smithsonian Contributions Zoology, 126, 1 - 158. https: // doi. org / 10.5479 / si. 00810282.126","Perrier, E. (1881) Description sommaire des especes nouvelles d'Asteries. Museum of Comparative Zoology Bulletin, 9, 1 - 31."]}

Astropecten acutiradiatus Tortonese, 1956 Astropecten acutiradiatus Tortonese, 1956: 323–324, pl.8. Astropecten acutiradiatus — Brito 1962: 3; 1968: 8; Tommasi 1970: 6, fig. 18; Clark & Downey 1992: 26–27, fig. 9b, pls. 5C, D. Distribution. Curaçao, Venezuela (Clark & Downey 1992). BRAZIL: Bahia, Rio de Janeiro (Tortonese 1956; Brito 1968; Tommasi 1970; Clark & Downey 1992; Ventura et al. 2007). Depth. 35–66 (475?) m (Clark & Downey 1992; Ventura et al. 2007). Biological notes. Specimens of A. acutiradiatus live on sandy bottoms and feed mainly on crustaceans and gastropods (Ventura et al. 2007). Holotype. Museo Civico di Storia Naturale di Genova, C.E. 36017 (Clark & Downey 1992). Type locality. Baía de Guanabara, Rio de Janeiro, Brazil (Brito 1968). Remarks. Despite four years of collection in Baía de Guanabara, Brito and his team never found additional specimens of A. acutiradiatus (Brito 1968). Ventura et al. (2007) recorded the species for Bahia, but unfortunately, we did not have access to this material to make any further observations. Astropecten acutiradiatus is poorly known and ontogenetic variation has not been documented. The arms of the holotype are long and narrow, unlike any other Brazilian Astropecten. Clark & Downey (1992) suggested that this species should be considered a subspecies of Astropecten articulatus. Although Tortonese (1956) mentioned the similarity between A. acutiradiatus and A. articulatus, he did not mention characters used to separate these species. A revision with the examination of more specimens is necessary to elucidate this question., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on page 8, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Tortonese, E. (1956) Su alcune specie di Astropectinidae, con descrizone di una nuovo Astropecten (Asteroidea). Annali Museo Storia Naturale Genova, 68, 319 - 334.","Brito, I. M. (1962) Ensaio de catailogo dos equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zooloigicos, 13, 1 - 10.","Brito, I. M. (1968) Asteroiides e equinoiides do Estado da Guanabara e adjace ˆ ncias. Boletim do Museu Nacional, 260, 1 - 51.","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Ventura, C. R. R., Verissimo, I., Nobre, C. C. & Zama, P. C. (2007) Filo Echinodermata. In: Lavrado, H. P. & Viana, M. S. (Eds.), Atlas de invertebrados marinhos da regiao central da Zona Economica Exclusiva brasileira. Parte 1. Museu Nacional do Rio de Janeiro, Serie Livros 25. Museu Nacional do Rio de Janeiro, Rio de Janeiro, pp. 258."]}

Luidia clathrata (Say, 1825) Figures 9–10 Asterias clathrata Say, 1825: 142. Luidia clathrata — Tommasi 1958: 9, fig. 1, pl. 2; Brito 1962: 4; 1968: 11–12, fig. 2, pl. 2; Tommasi & Aron 1987: 3; Tommasi et al. 1988: 6; Clark & Downey 1992: 13, figs. 4d, 5e–g, 6g, i, 8g, pl. 4B; Magalhães et al. 2005: 63; Ventura et al. 2007: 237; Manso et al. 2008: 185, fig. 7a–e; Lima & Fernandes 2009: 58; Magris & Deìstro 2010: 59; Xavier 2010: 75; Benavides-Serrato et al. 2011: 99–100; Gondim et al. 2014: 10–11, figs. 3e–h; Sandino et al. 2017: S294; Bueno et al. 2018: 178–179, fig. 8; Gurjão & Lotufo 2018: 11; Miranda 2018: 14, fig. 10C; Patrizzi & Dobrovolski 2018: 182; Borrero-Peìrez et al. 2019: 4; Torres & Torres 2019: 413. Material examined (1 spec, 57 mm R). BRAZIL. Bahia, Todos os Santos Bay (13°10’S; 38°45’W)— 15 m, 15.iii.1997, 1 spec, R 57 mm (UFBA 330). Comparative material. U.S.A., North Carolina, Cape Hatteras, 26 m, 19.x.1884, 1 spec, R 70 mm (NMNH 8507, neotype). Description (R 57 mm). Flat disc (Fig. 9A); R/r 5.2. Five tapering arms. Abactinal plates covered by paxillae (Fig. 10A) with robust granules (3–7) in center and a fringe of small peripheral spinelets. Paxillae of disc and arms smaller, irregularly arranged (Fig. 10F). Paxillae from periphery of arms in five regular rows, square to rectangular in abactinal view (Fig. 10D); paxillae from three outer rows equal to or larger than those from inner rows. Madreporite often with irregular outline and hidden by paxillae (Fig. 10H). Superomarginal plates paxilliform. Inferomarginal plates elongated and separated by a gap. Marginal region of plates with two unequal, denticulate spines (Fig. 10B); around these, several small spines. Actinal surface of plate densely covered by flat spines, large in central region and minute in marginal region (Fig. 10G). Four adambulacral spines. Two curved spines, innermost spine smallest; two flattened, truncated spines of same size, side by side above (Fig. 10C, E). Oral plate with long spines, apical ones larger (Fig. 10I). Tube feet in two rows, sucking disc lacking. Pedicellariae absent. Coloration. No record of coloration of in vivo specimens from this region. In ethanol, specimens are pale beige to white. Specimens from the Gulf of Mexico have a noticeable grey band on the dorsal surface of the inferomarginal plates, above the fringe spines that is retained for years even in alcohol (Janessa Fletcher, pers. comm. on 4 Dec 20, “Picture Guide to the SEAMAP specimens of the Eastern Gulf of Mexico ”, unpublished). The dry specimen in Hopkins & Knott (2010) has greyish upper arm surface, white ocular tips, white inferomarginal spines and off-white actinal surface. Frequently shows darker grey dorsal central arm stripe, but it is not encountered in the neotype. Distribution. U.S.A. (NC, FL), Bermuda, Gulf of Mexico, Mexico, The The Bahamas, Caribbean Sea, Cuba, Dominican Republic, Puerto Rico, Belize, Honduras, Nicaragua, Panama, Colombia, Venezuela, Guyana (Sandino et al. 2017; Borrero-Peìrez et al. 2019; Mah 2020a). BRAZIL: Amapá, Pará, Paraíba, Bahia, Rio de Janeiro, São Paulo, Paraná, Santa Catarina (Rathbun 1879; Bernasconi 1943; Tommasi 1958, 1970; Brito 1960, 1968; Downey 1973; Walenkamp 1976; 1979; Clark & Downey 1992; Hendler et al. 1995; Benavides-Serrato et al. 2005; Mag- alhães et al. 2005; Alvarado et al. 2008; Manso et al. 2008; Lima & Fernandes 2009; Xavier 2010; Gondim et al. 2014; Bueno et al. 2018; Miranda 2018; Torres & Torres 2019). Depth. 0–175 m (Clark & Downey 1992). Biological notes. The specimen described here was found on a silty soft bottom environment, just outside Todos os Santos Bay; additional specimens have been found in sandy and muddy sediments inside of the Bay. Hendler et al. (1995) also reported L. clathrata ’s preference for protected environments with fine sediments, sometimes in estuarine conditions. This species has a generalized diet, feeding on foraminifera and small to medium-sized invertebrates, including polychaetes, crustaceans and ophiuroids (Lawrence et al. 1974; Pechaszadeh & Lera 1983; Tararam et al. 1993), but Schwartz & Porter (1977) reported a strong preference for scallops in North Carolina. Caribbean populations of L. clathrata have seasonal reproduction (Lawrence 1973; Dehn 1980a, b; Watt & Lawrence 1990; Pomory & Lares 2000). Luidia clathrata is classified as “Least Concern” by the Ministry of the Environment (MMA 2018). According to Gurjão & Lotufo (2018), its harvesting in Brazil is currently prohibited. Neotype. NMNH 8507, designated by Hopkins & Knott (2010) because the holotype has never been found. Type locality. Cape Hatteras, NC, U.S.A. Remarks. The specimen described differs from the neotype by having two inferomarginal spines (vs. 3) and 3–7 granules in central region of paxillae (vs. 7–11). Walenkamp (1976) reported that ontogenetic changes in this species include an increase in the number of large lateral spines (1–2 in smaller specimens; 3 in adults), and the drawing of L. clathara ’s paxilla in Clark & Downey (1992, fig. 6g) indicates that larger specimens have more than 10 central granules (not quantified, but described as being numerous). Also, the specimens analyzed by Gondim et al. (2014), measuring 26–42 mm R (R/r 6.32–6.48), have two inferomarginal spines and 1–6 granules. Adults of L. clathara may reach 160 mm of R and the average R/r for adults is between 7–8. According to H.L. Clark (1933), adult specimens have R 100 mm or more., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 21-22, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Say, T. (1825) On the species of the Linnaean genus Asterias inhabiting the coast of the United States. Journal of the Academy of Natural Sciences, Philadelphia, 1 (5), 151 - 154.","Tommasi, L. R. (1958) Os equinodermos do litoral de Sao Paulo. II. Diadematidae, Schizasteridae, Brissidae, Cidaroidea (Echinoidea) e Asteroidea do Bentos Costeiro. Contribuicoes avulsas do Instituto Oceanografico, 2, 1 - 27, 6 pls.","Brito, I. M. (1962) Ensaio de catailogo dos equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zooloigicos, 13, 1 - 10.","Brito, I. M. (1968) Asteroiides e equinoiides do Estado da Guanabara e adjace ˆ ncias. Boletim do Museu Nacional, 260, 1 - 51.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 9 (1), 61 - 65. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Ventura, C. R. R., Verissimo, I., Nobre, C. C. & Zama, P. C. (2007) Filo Echinodermata. In: Lavrado, H. P. & Viana, M. S. (Eds.), Atlas de invertebrados marinhos da regiao central da Zona Economica Exclusiva brasileira. Parte 1. Museu Nacional do Rio de Janeiro, Serie Livros 25. Museu Nacional do Rio de Janeiro, Rio de Janeiro, pp. 258.","Manso, C. L. C., Alves, O. F. S. & Martins, L. R. (2008) Echinodermata da Baia de Todos os Santos e da Baia de Aratu (Bahia, Brasil). Biota Neotropica, 8 (3), 179 - 196. https: // doi. org / 10.1590 / S 1676 - 06032008000300017","Lima, E. J. B. & Fernandes, M. L. B. (2009) Diversidade de equinodermos (Echinodermata) no estado de Pernambuco (Brasil). Revista Brasileira de Zoociencias, 11, 55 - 63.","Xavier, L. A. R. (2010) Inventario dos equinodermos do Estado de Santa Catarina, Brasil. Brazilian Journal of Aquatic Science and Technology, 14, 73 - 78. https: // doi. org / 10.14210 / bjast. v 14 n 2. p 73 - 78","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Sandino, O. B., Solis-Marin, F. A., Caballero-Ochoa, A. A., Conejeros-Vargas, C. A., Marquez-Borras, F., Ayala-Aguilera, A. P. & Laguarda-Figueras, A. (2017) Equinodermos de Nicaragua: nuevos registros del Pacifico y Caribe Sur. Revista de Biologia Tropical, 65, 288 - 298. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31696","Bueno, M. L., Alitto, R. A. S., Guilherme P. D. B., Domenico, M. D. & Borges, M. (2018) Guia ilustrado dos Echinodermata da porcao sul do Embaiamento Sul Brasileiro. Pesquisa e Ensino em Ciencias Exatas e da Natureza, 2 (2), 169 - 237. https: // doi. org / 10.29215 / pecen. v 2 i 2.1071","Gurjao, L. M. & Lotufo, T. M. C. (2018) Native species exploited by marine aquarium trade in Brazil. Biota Neotropica, 18 (3), e 20170387. https: // doi. org / 10.1590 / 1676 - 0611 - bn- 2017 - 0387","Miranda, A. P. S. (2018) Diversidade e distribuicao de equinodermos da Plataforma Continental Amazonica. Unpublished Honors Thesis, Universidade Federal do Para, Belem, 62 pp.","Patrizzi, N. & Dobrovolski, R. (2018) Integrating climate change and human impacts into marine spatial planning: a case study of threatened starfish species in Brazil. Ocean & Coastal Management, 161, 177 - 188. https: // doi. org / 10.1016 / j. ocecoaman. 2018.05.003","Torres, V. S. & Torres, F. S. S. (2019) Inventario preliminar dos invertebrados marinhos observados na praia da Pinheira, Palhoca, SC, Brasil. UNISANTA Bioscience, 8 (4), 408 - 422.","Hopkins, T. S. & Knott, K. E. (2010) The establishment of a neotype for Luidia clathrata (Say, 1825) and a new species within the genus Luidia (Asteroidea: Paxillosida: Luidiidae). In: Harris, L. G., Bottger, S. A., Walker, C. W. & Lesser, M. P. (Eds.), Proceedings of the 12 th International Echinoderm Conference, 7 - 11 August 2006, Durham, New Hampshire. Taylor & Francis Group, London, pp. 207 - 212. https: // doi. org / 10.1201 / 9780203869543 - c 32","Mah, C. L. (2020 a) World Asteroidea Database. Available from: http: // www. marinespecies. org / asteroidea / (accessed 30 April 2020)","Rathbun, R. (1879) A list of the Brazilian echinoderms, with notes on their distribution. Transactions of the Connecticut Academy of Arts and Sciences, 5 (3), 139 - 158. https: // doi. org / 10.5962 / bhl. title. 16126","Bernasconi, I. (1943) Los Asteroideos sulamericanos de la familia Luidiidae. Anales del Museo Argentino de Ciencias Naturales, 7, 1 - 20.","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Brito, I. M. (1960) Asteroides dos estados do Rio de Janeiro e de Sao Paulo. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 5, 1 - 13.","Downey, M. E. (1973) Starfishes from the Caribbean and the Gulf of Mexico. Smithsonian Contributions Zoology, 126, 1 - 158. https: // doi. org / 10.5479 / si. 00810282.126","Walenkamp. J. H. C. (1976) The asteroids of the coastal waters of Surinam. Zoologische Verhandelingen, 147, 1 - 91, 18 pls.","Walenkamp, J. H. C. (1979) Asteroidea (Echinodermata) from the Guyana shelf. Zoologische Verhandelingen, 170, 1 - 97.","Hendler, G., Muller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea Stars, Sea Urchins and Allies: Echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp.","Benavides-Serrato, M., Borrero-Perez, G. H., Navas, G. R. & Solano, O. D. (2005) Listado taxonomico de los asteroideos (Echinodermata: Asteroidea) de la plataforma y el talud superior del caribe colombiano. Revista de Biologia Tropical, 53 (13), 171 - 194.","Alvarado, J. J., Solis-Marin, F. A. & Ahearn, C. (2008) Equinodermos (Echinodermata) del Caribe Centroamericano. Revista de Biologia Tropical, 56, 37 - 55.","Lawrence, J. M., Erwin, K. & Turner, R. L. (1974) Stomach contents of Luidia clathrata (Asteroidea). Florida Scientist, 37, 8.","Pechaszadeh, P. E. & Lera, M. E. (1983) Alimentacion de tres especies tropicles de Luidia (Echinodermata: Asteroidea) en Golfo Triste, Venezuela. Caribbean Journal of Science, 19, 1 - 6.","Tararam, A. S., Wakabara, Y. & Equi, M. B. (1993) Habitos alimentares de onze especies de megafauna bentica da plataforma continental de Ubatuba, SP. Publicacao Especial Instituto Oceanografico, 10, 159 - 167.","Schwartz, F. J. & Porter, H. J. 1977. Fishes, macroinvertebrates, and their ecological interrelationships with a calico scallop bed off North Carolina. Fishery Bulletin, 75 (2), 427 - 446.","Lawrence, J. M. (1973) Level, content, and caloric equivalents of the lipid, carbohydrate, and protein in the body components of Luidia clathrata (Echinodermata: Asteroidea: Platyasterida) in Tampa Bay. Journal of Experimental Marine Biology and Ecology, 11, 263 - 274. https: // doi. org / 10.1016 / 0022 - 0981 (73) 90026 - 9","Dehn, P. F. (1980 a) Growth and reproduction in Luidia clathrata (Say) (Echinodermata: Asteroidea). Unpublished PhD Dissertation, University of South Florida, Tampa, Florida, 183 pp.","Pomory, C. M. & Lares, M. T. (2000) Rate of regeneration of two arms in the field and its effect on body components in Luidia clathrata. Journal of Experimental Marine Biology and Ecology, 254, 211 - 230. https: // doi. org / 10.1016 / S 0022 - 0981 (00) 00281 - 1","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp.","Clark, H. L. (1933) A handbook of the littoral echinoderms of Porto Rico and the other West Indian Islands. Scientific Survey of Puerto Rico, 16, 1 - 147."]}

Astropecten antillensis Lu ̈tken, 1860 Figures 2–3 Astropecten antillensis Lu ̈tken, 1860: 47. Astropecten antillensis — Clark & Downey 1992: 29–30, figs. 9c, 11c, d, pls 9C, D; Benavides-Serrato et al. 2011: 114; Williams et al. 2013: 456; Nisperuza et al. 2016: 115; Soaréz 2016: 78; Borrero-Peìrez et al. 2019: 4; Cunha et al. 2020: 47, fig. 9. Material examined (3 specs, 20–43 mm R). BRAZIL. Bahia (12°52’– 12°56’S; 38°12’– 38°30’W)— Busca Vida beach, Guarajuba, 45 m, ii.2008, 1 spec, R 20 mm (UFBA 949). Salvador: Plataforma beach, 16 m, 27.ix.2010, 1 spec, R 21 mm (UFBA 1183); Porto beach, 13 m, vi.2012, 1 spec, R 43 mm (UFBA 1620). Comparative material. VIRGIN ISLAND. St. Thomas, 4 specs, R 10–56 mm (NHMD 76172, syntypes). Description (R 43 mm). Disc half size of arms; R/r 2.5; 22 SM plates; R/SM# 1.95. Arms long and narrow (Fig. 2A–C). Abactinal region covered by paxillae with 1–5 central and 10–12 peripheral spinelets (Fig. 2D). Number of paxillae spinelets increases toward center of disc. Madreporite small, partially hidden among paxillae (Fig. 2D). Superomarginal and inferomarginal plates organized in one row each; inferomarginal plates not projected beyond superomarginal plates. Superomarginal plates covered by spines, fine granules and spinelets. First superomarginal plate with one spine, others with two spines (Fig. 2E). Spine on first plate twice as large as spines from other plates. In plates with two spines, inner spine smaller than outer spine. Inferomarginal plates separated by a gap. Fringe with two layers of flattened and pointed spines, oblique, top layer with two large spines of equal size and another thinner and smaller spine (Fig. 2F), bottom layer with three small spines. Actinal surface of inferomarginal plates lacks squamules (short, flat, scalelike spines; from Cobb et al. 2019); central region naked, sometimes with long spines similar to lateral fringe spines (Fig. 2H). Three equal-sized adambulacral spines forming a single row. Subambulacral row with three spines, central spine larger than adjacent ones (Fig. 2I); sometimes a second row with 1–3 spines present. Oral spines with broad, blunt tip (Fig. 2G). Pedicellariae absent. Ontogenetic variation (R 20–21 mm). Average R/r 4.1; 19 SM plates (R 20 mm); R/SM# 1.0. Paxillae with 1–3 central and 5–9 peripheral spines (Fig. 3D). Inferomarginal plates project beyond superomarginal plates (Fig. 3E). Superomarginal plates with only one spine; spine on first superomarginal plate relatively larger than that of larger specimen (Fig. 3A, F). Actinal region of inferomarginal plates with fewer squamules than those of larger specimens. Coloration. Yellowish body with white inferomarginal fringe; Caribbean specimens are yellowish, reddish and orange in situ (Clark & Downey 1992; Benavides-Serrato et al. 2005); white or pale brown in ethanol. Distribution. Gulf of Mexico, Cuba, Antilles, Puerto Rico, Colombia (Sladen 1889; Bayer et al. 1970; Clark & Downey 1992; Abreu-Pérez et al. 2005; Williams et al. 2013; Nisperuza et al. 2016; Soaréz 2016; Borrero-Peìrez et al. 2019). BRAZIL: Bahia and possibly Trindade Island (John 1948; Cunha et al. 2020). Depth. 3–278 m (Clark & Downey 1992). Biological notes. In Bahia, Astropecten antillensis lives in nutrient-poor regions with sandy and muddy bottoms. This species can also be found in rubble bottoms (Zoppi de Roa 1967; Alvarado & Solís-Marín 2013). Syntype. NHMD 76172 [previously as ZMUC AST 4], 76371 (Tom Schiøtte, per. comm). Type locality. St. Thomas, Virgin Islands. Remarks. John (1948) noticed that some of the specimens identified as A. brasiliensis by Sladen (1889) were actually A. antillensis; John’s publication was the first record of this species in Brazil. H.L. Clark (1901) most likely misidentified the Astropecten specimens collected in Puerto Rico because in his identification key, he reported that specimens of A. antillensis do not have spines on marginal plates (given that Clark mentioned in the description following the key that the specimens had inferomarginal spines, “marginal” probably referred to the superomarginal plates)., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 8-10, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Williams, S. M., Benavides-Serrato, M., Garcia-Arraras, J. E., Hernandez-Delgado, E. A. & Rodriguez-Barreras, R. (2013) Review of Echinoderm Research in Puerto Rico, with the Focus on Biological and Ecological Aspects. In: Alvarado, J. J. & Solis-Marin, F. A. (Eds.), Echinoderm Research and Diversity in Latin America. Springer, Berlin, pp. 437 - 469. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9 _ 14","Nisperuza, P. C., Padilla, C. J. & Quiros, R. J. (2016) Current status of the study of the caribbean cordobes echinoderms, Colombia. Revista Colombiana de Ciencia Animal, 8 (1), 112 - 119. https: // doi. org / 10.24188 / recia. v 8. n 1.2016.234","Cunha, R., Tavares, M. & Mendonca, J. B. (2020) Asteroidea (Echinodermata) from shallow-waters of the remote oceanic archipelago Trindade and Martin Vaz, southeastern Atlantic, with taxonomic and zoogeographical notes. Zootaxa, 4742 (1), 31 - 56. https: // doi. org / 10.11646 / zootaxa. 4742.1.2","Cobb, J. C., Lawrence, J. M., Herrera, J. C. & Janies, D. (2019) A new species of Astropecten (Echinodermata: Asteroidea: Paxillosida: Astropectinidae) and a comparison of the Astropecten species from the Gulf of Mexico and the East Florida Shelf. Zootaxa, 4612 (3), 301 - 325. https: // doi. org / 10.11646 / zootaxa. 4612.3.1","Benavides-Serrato, M., Borrero-Perez, G. H., Navas, G. R. & Solano, O. D. (2005) Listado taxonomico de los asteroideos (Echinodermata: Asteroidea) de la plataforma y el talud superior del caribe colombiano. Revista de Biologia Tropical, 53 (13), 171 - 194.","Sladen, W. P. (1889) Asteroidea. Report of the Scientific Results of H. M. S. Challenger, 30, 1 - 893. https: // doi. org / 10.1111 / j. 1096 - 3642.1882. tb 02281. x","Bayer, F., Voss, G. & Robin, R. (1970) Bioenvironmental and radiological safety feasibility studies Atlantic-Pacific interoceanic canal. In: Report on the marine fauna and benthic shelf slope communities of the isthmian region. Vol. 1. University of Miami, Rosenstiel School of Marine and Atmospheric Science, Florida, pp. 99.","Abreu-Perez, M., Solis-Marin, F. A. & Laguarda-Figueras, A. (2005) Catalogo de los equinodermos (Echinodermata: Asteroidea y Ophiuroidea) neritico-bentonicos del Arquipielago Cubano. Revista de Biologia Tropical, 53, 29 - 52.","John, D. D. (1948) Notes on the asteroids in the British Museum (Natural History). I. The species of Astropecten. Novitates Zoologicae, 42, 485 - 508.","Zoppi De Roa, E. (1967) Contribucion al estudio de los equinodermos de Venezuela. Acta Biologica de Venezuela, 5, 267 - 333.","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Vol. 1. Springer, Berlin, 658 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Clark, H. L. (1901) The echinoderms of Porto Rico. Bulletin of the United States Fish Commission, 20, 233 - 263, 4 pls."]}

Astropecten marginatus Gray, 1840 Figures 6–7 Astropecten marginatus Gray, 1840: 181. Astropecten marginatus — Brito 1962: 3; 1968: 7, pl. 4, fig. 1; Nomura & Fausto Filho 1966: 19; Lima-Verde 1969: 11; Carrera- Rodriguez & Tommasi 1977: 88–89; Tommasi et al. 1988: 5; Clark & Downey 1992: 41–42, fig. 10e, pl. 11A–B; Gondim et al. 2008: 155; Ventura et al. 2008: 177–178; Lima & Fernandes 2009: 58; Magris & Deìstro 2010: 59; Xavier 2010: 75; Gondim et al. 2014: 22–24, figs. 6a–e, 12b; Guilherme & Rosa 2014; Fernandez et al. 2017; Bueno et al. 2018: 177, fig. 6; Gurjão & Lotufo 2018: 10; Miranda 2018: 14; Patrizzi & Dobrovolski 2018: 182; Torres & Torres 2019: 412; Turra et al. 2019. Astropecten richardi — Perrier 1875: 372. Material examined (21 specs, 10–50 mm R). BRAZIL. Bahia, Santa Cruz de Cabrália (16°14’– 16°15’S; 38°00’– 38°58’W)— 7 m, 16.xi.2010, 5 specs, R 11–25 mm (MZUSP 2100); 6 m, 16.xi.2010, 6 specs, R 10–28 mm (MZUSP 2101); 7 m, 16.xi.2010, 5 specs, R 30–50 mm (MZUSP 2100); 6 m, 16.xi.2010, 5 specs, R 30–40 mm (MZUSP 2101). Description (R 30–50 mm). Disc broad and flattened, arms short and flattened (Fig. 6A–C); R/r 3.3; 39 SM plates (R 50 mm); R/SM# 1.3. Abactinal region covered by paxillae as follows: proximal region of arms and disc with 10–12 central and 15 peripheral spinelets (Fig. 6D), midline region of arms with 5–8 central and nine peripheral spinelets, and distal region of arms with 1–3 central and seven peripheral spinelets. Madreporite large, not hidden by paxillae (Fig. 6E). Marginal plates horizontal and elongated. Superomarginal plates without spines and covered by granules: central granules robust, peripheral granules elongated, hyaline and thin. Inferomarginal plates projected beyond superomarginal plates. Fringe with two layers of spines, parallel, top layer with two spines of equal size, and a third proximal 3x smaller and thinner (Fig. 6G); bottom layer with 4–5 small spines, proximal spine smaller and thinner (Fig. 6I). Actinal region of inferomarginal plates lacking squamules; central region naked, sometimes with long spines similar to lateral fringe spines (Fig. 6J). Three adambulacral spines forming single line, central spine longer than adjacent ones (Fig. 6K). One large subambulacral spine in center of plate, largest spine in adambulacral plate, surrounded by acicular spinelets; spinelets between large spine and furrow spines. Oral spines flattened with blunt tip (Fig. 6F). Pedicellariae absent. Ontogenetic variation (R 10–28 mm). Average R/r 3.0; 22 SM plates (R 20 mm); R/SM# 1.1. Proximal paxillae with 3–5 central and 8–10 peripheral spinelets (Fig. 7C), distal paxillae with 0–1 central and 4–5 peripheral spinelets. Inferomarginal fringe with two layers as in larger specimen, but proximal spine in top layer 5x smaller than adjacent spines, and bottom layer with 3–4 small spines, proximal spine smallest (Fig. 7F). Three adambulacral spines forming single line, central spine largest in adambulacral plate (Fig. 7E). Second subambulacral row as in larger specimen, with central spine largest than adjacent spines, but not largest in adambulacral plate; third row with 2–4 sub-equal spines. Coloration. Abactinal surface greyish in center of disc and arms, margins orangish to creamy; inferomarginal spines fade from a bluish-grey base to creamy tips. Specimens in ethanol are pale brown. Distribution. Gulf of Mexico, Mexico, Dominican Republic, Puerto Rico, Costa Rica, Colombia, Venezuela, Guyana, Suriname, French Guyana (Walenkamp 1976; Clark & Downey 1992; Alvarado et al. 2008; Benavides- Serrato et al. 2005, 2011; Pawson et al. 2009, Alvarado & Solís-Marín 2013). BRAZIL: Amapá, Ceará, Rio Grande do Norte, Paraíba, Pernambuco, Bahia, Rio de Janeiro, São Paulo, Paraná, Santa Catarina, Rio Grande do Sul (Bernasconi 1955; Tommasi 1958; Brito 1962; Lima-Verde 1969; Netto 2006; Gondim et al. 2008; Pelaes 2008; Lima & Fernandes 2009; Xavier 2010; Gondim et al. 2014; Bueno et al. 2018; Miranda 2018; Torres & Torres 2019; Turra et al. 2019). Depth. 1–130 m (Clark & Downey 1992). Biological notes. This species lives in muddy, sandy and rubble bottoms (Alvarado & Solís-Marín 2013) and most likely feeds during the day, preferably on gastropods, bivalves and crustaceans (Bitter 1984; Zoldan 2005; Guilherme & Rosa 2014; Fernandez et al. 2017). In Caraguatatuba Bay, São Paulo, A. marginatus lives in sympatry with L. senegalensis, where they partition food resources (Fernandez et al. 2017). The reproductive cycle of A. marginatus in this region is annual and spawning happens mainly during the rainy season (Turra et al. 2019). Netto (2006) found many specimens gathered in shallow pools at night with 2–5 arms facing upwards, possibly engaged in gas exchange as the exposure of the tube feet into the water column should increase its breathing surface (Farmanfarmaian 1966). Astropecten marginatus is classified as “Vulnerable” (baseline data indicate a reduction in population by at least 30% over the next 100 years) by the Ministry of the Environment (MMA 2018). According to Gurjão & Lotufo (2018), its harvesting in Brazil is currently prohibited. Holotype. Gray (1840, p. 178) mentioned that the specimens he described were deposited in the collections of the Zoological Society of London (especially South American specimens collected by Mr. Hugh Cuming) and of the British Museum, where he worked. However, Andrew Cabrinovic told us that the type has not been found at the NHM-UK (pers. comm. on 20 Feb 20). Type locality. Not reported by Gray (1840). Taxonomic remarks of the genus Astropecten As noted by Cobb et al. (2019), telling some of the Astropecten species apart is not trivial. We found several instances of misidentification in prior work from Northeastern Brazil and we hope that the present work will reduce errors in future studies. Phylogenetic relationships between some species of Brazilian Astropecten were addressed by Zulliger & Lessios (2010) and Cobb et al. (2019). Their molecular cladograms indicate that A. antillensis is more closely related to A. articulatus than to A. cingulatus. The relationship between these species and A. marginatus is uncertain as this species could be equally related to A. antillensis, A. articulatus and A. cingulatus or most closely related to A. cingulatus (Zulliger & Lessios, 2010). Astropencten brasiliensis was not included in these studies. Astropecten acutiradiatus differs from A. alligator by having an enlarged spine in the first subambulacral row (vs. three (sub)equal spines); from A. antillensis by having an enlarged distal spine in the first subambulacral row (vs. enlarged central spine); from A. articulatus by having clavate paxillar spinelets (vs. paxillar spinelets granulose); from A. brasiliensis brasiliensis and A. duplicatus by the absence of enlarged spines on the distal superomarginal plates (vs. presence of enlarged spines on the distal superomarginal plates); from A. cingulatus and A. marginatus by having spines on the proximal superomarginal plates (vs. superomarginal spines absent). Astropecten antillensis differs from A. acutiradiatus, A. articulatus, A. brasiliensis brasiliensis and A. duplicatus by having an enlarged central spine in the first subambulacral row (vs. enlarged distal spine); from A. alligator by having an enlarged spine in the first subambulacral row (vs. three (sub)equal spines); from A. cingulatus and A. marginatus by having spines on the superomarginal plates (vs. superomarginal spines absent). Astropecten antillensis also differs from A. brasiliensis brasiliensis because its inner superomarginal (not the first pair) spine is smaller than the outer spine (vs. inner spine larger than outer spine). Astropecten brasiliensis brasiliensis differs from A. marginatus and A. cingulatus by having superomarginal spines (vs. superomarginal spines absent); from A. articulatus and A. duplicatus by having subambulacral spines with blunt tips (vs. spines truncated in two), and inferomarginal fringe spines oblique (vs. inferomarginal fringe spines horizontal); and from A. alligator by having an enlarged distal spine in the first subambulacral row (vs. (sub)equal spines). Astropecten cingulatus differs from A. marginatus by having inferomarginal plates densely covered with overlapping squamules and spines (vs. inferomarginal plates bare or with sparse, widely spaced squamules), and these two species differ from the other Brazilian Astropecten by having no spines on the superomarginal plates (vs. spines present)., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 15-18, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Gray, J. E. (1840) A synopsis of the genera and species of the class Hypostoma (Asterias Linnaeus). Annals of the Magazine of Natural History, Series 1, 6 (37), 175 - 184 + 275 - 290. https: // doi. org / 10.1080 / 03745484009443296","Brito, I. M. (1962) Ensaio de catailogo dos equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zooloigicos, 13, 1 - 10.","Brito, I. M. (1968) Asteroiides e equinoiides do Estado da Guanabara e adjace ˆ ncias. Boletim do Museu Nacional, 260, 1 - 51.","Nomura, H. & Fausto Filho, J. (1966) Shrimp survey in coastal and offshore waters of northeastern Brazil. Arquivos da Estacao de Biologia Marinha da Universidade Federal do Ceara, 6, 15 - 29.","Lima-Verde, J. S. (1969) Primeira contribuicao ao inventario dos equinodermas do Nordeste brasileiro. Arquivos de Ciencias do Mar, 9 (1), 9 - 13.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Gondim, A., Lacouth, P., Alonso, C. & Manso, C. (2008) Echinodermata da praia do Cabo Branco, JoaTo Pessoa, Paraiiba, Brasil. Biota Neotropica, 8, 151 - 159. https: // doi. org / 10.1590 / S 1676 - 06032008000200016","Lima, E. J. B. & Fernandes, M. L. B. (2009) Diversidade de equinodermos (Echinodermata) no estado de Pernambuco (Brasil). Revista Brasileira de Zoociencias, 11, 55 - 63.","Xavier, L. A. R. (2010) Inventario dos equinodermos do Estado de Santa Catarina, Brasil. Brazilian Journal of Aquatic Science and Technology, 14, 73 - 78. https: // doi. org / 10.14210 / bjast. v 14 n 2. p 73 - 78","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Guilherme, P. D. & Rosa, L. C. (2014) Seasonal variation in body size and diet of the sea star Astropecten marginatus (Paxillosida, Astropectinidae) off coast of Parana, Southern Brazil. Revista de Biologia Tropical, 62, 59 - 68. https: // doi. org / 10.15517 / rbt. v 62 i 1.7888","Fernandez, W. S., Dias, G. M., Majer, A., Delboni, C., Denadai, M. R. & Turra, A. (2017) Resource partitioning between sympatric starfish from tropical unconsolidated substrate: implications for coexistence and top-down control on benthic prey. Estuarine, Coastal and Shelf Science, 196, 141 - 149. https: // doi. org / 10.1016 / j. ecss. 2017.06.039","Bueno, M. L., Alitto, R. A. S., Guilherme P. D. B., Domenico, M. D. & Borges, M. (2018) Guia ilustrado dos Echinodermata da porcao sul do Embaiamento Sul Brasileiro. Pesquisa e Ensino em Ciencias Exatas e da Natureza, 2 (2), 169 - 237. https: // doi. org / 10.29215 / pecen. v 2 i 2.1071","Gurjao, L. M. & Lotufo, T. M. C. (2018) Native species exploited by marine aquarium trade in Brazil. Biota Neotropica, 18 (3), e 20170387. https: // doi. org / 10.1590 / 1676 - 0611 - bn- 2017 - 0387","Miranda, A. P. S. (2018) Diversidade e distribuicao de equinodermos da Plataforma Continental Amazonica. Unpublished Honors Thesis, Universidade Federal do Para, Belem, 62 pp.","Patrizzi, N. & Dobrovolski, R. (2018) Integrating climate change and human impacts into marine spatial planning: a case study of threatened starfish species in Brazil. Ocean & Coastal Management, 161, 177 - 188. https: // doi. org / 10.1016 / j. ocecoaman. 2018.05.003","Torres, V. S. & Torres, F. S. S. (2019) Inventario preliminar dos invertebrados marinhos observados na praia da Pinheira, Palhoca, SC, Brasil. UNISANTA Bioscience, 8 (4), 408 - 422.","Turra, A., Checon, H. H., Fernandez, W. S., Majer, A. P., Delboni, C. G., Denadai, M. R. & Dias, G. M. (2019) The importance of embayed coastal areas for the life cycle of two sympatric starfish species (Echinodermata, Asteroidea). Estuarine, Coastal and Shelf Science, 219, 128 - 138. https: // doi. org / 10.1016 / j. ecss. 2019.01.028","Perrier, E. (1875) Revision de la collection de stellerides du Museum d'Histoire Naturelle de Paris. Archives de Zoologie Experimentale et Generate, 4, 265 - 450.","Walenkamp. J. H. C. (1976) The asteroids of the coastal waters of Surinam. Zoologische Verhandelingen, 147, 1 - 91, 18 pls.","Alvarado, J. J., Solis-Marin, F. A. & Ahearn, C. (2008) Equinodermos (Echinodermata) del Caribe Centroamericano. Revista de Biologia Tropical, 56, 37 - 55.","Pawson, D. L., Vance, D. J., Messing, C. G., Solis-Marin, F. A. & Mah, C. L. (2009) Echinodermata of the Gulf of Mexico. In: Felder, D. L. & Camp, D. K. (Eds.), Gulf of Mexico - Origins, Waters, and Biota. Texas A & M University Press, College Station, Texas, pp. 1177 - 1204.","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Vol. 1. Springer, Berlin, 658 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Bernasconi, I. (1955) Equinoideos y asteroideos de la coleccion del instituto oceanografico de la Universidad de San Pablo. Primera contribucion. Boletim do Instituto Oceanografico, 6, 51 - 57. https: // doi. org / 10.1590 / S 0373 - 55241955000100002","Tommasi, L. R. (1958) Os equinodermos do litoral de Sao Paulo. II. Diadematidae, Schizasteridae, Brissidae, Cidaroidea (Echinoidea) e Asteroidea do Bentos Costeiro. Contribuicoes avulsas do Instituto Oceanografico, 2, 1 - 27, 6 pls.","Netto, L. F. (2006) Echinodermata do Canal de Sao Sebastiao, Sao Sebastiao. Unpublished Master Thesis, Universidade de Sao Paulo, Sao Paulo, 201 pp.","Pelaes, M. A. L. (2008) Composicao, distribuicao espacial e temporal das especies de Echinodermata na plataforma rasa do sul da Bahia num trecho entre Itacare e Canavieiras. Unpublished Master Thesis, Universidade Estadual de Santa Cruz, Ilheus, Bahia, 97 pp.","Bitter, R. (1984) Composicion de la dieta de Astropecten marginatus (Echinodermata: Asteroidea). Golfo Triste, region Centro- Occidental. Boletin del Instituto Oceanografico, Universidad de Oriente, 23, 169 - 176.","Zoldan, L. V. (2005) Analise do conteudo estomacal de duas especies de estrela-do-mar: Astropecten marginatus (Gray, 1840) e Luidia senegalensis (Lamarck, 1816) na enseada de Caraguatatuba, SP, sudeste do Brasil. Unpublished Honors Thesis, Centro Universitario da Fundacao de Ensino Octavio Bastos, Sao Joao da Boa Vista, Sao Paulo, 41 pp.","Farmanfarmaian, A. (1966) The respiratory physiology of echinoderms. In: Boolootian, R. A. (Ed.), Physiology of Echinodermata. John Wiley & Sons, New York, pp. 245 - 265.","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp.","Cobb, J. C., Lawrence, J. M., Herrera, J. C. & Janies, D. (2019) A new species of Astropecten (Echinodermata: Asteroidea: Paxillosida: Astropectinidae) and a comparison of the Astropecten species from the Gulf of Mexico and the East Florida Shelf. Zootaxa, 4612 (3), 301 - 325. https: // doi. org / 10.11646 / zootaxa. 4612.3.1","Zulliger, D. E. & Lessios, H. A. (2010) Phylogenetic relationships in the genus Astropecten Gray (Paxillosida: Astropectinidae) on a global scale: molecular evidence for morphological convergence, species-complexes and possible cryptic speciation. Zootaxa, 2504 (1), 1 - 19. https: // doi. org / 10.11646 / zootaxa. 2504.1.1"]}

Genus Linckia Nardo, 1834 Type species. Linckia laevigata (Linnaeus, 1758) (type by original designation). Remarks. The genus Linckia is composed of nine species, only three of these are recorded for the Atlantic Ocean: L. bouvieri Perrier, 1875, L. guildingi and L. nodosa Perrier, 1875, the last two have been recorded in Brazil., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on page 42, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Nardo, J. D. (1834) De Asteriis. Isis von Oken, Encyclopaedische Zeitun, 27, 716 - 717.","Linnaeus, C. (1758) Systema Naturae. Impensis Direct, Laurentii Salvii, Holmiae, 824 pp.","Perrier, E. (1875) Revision de la collection de stellerides du Museum d'Histoire Naturelle de Paris. Archives de Zoologie Experimentale et Generate, 4, 265 - 450."]}

Asterinides folium (Lütken, 1860) Figures 17–18 Asteriscus folium Lütken, 1860: 60. Asterina folium — Clark & Downey 1992: 182, pl. 421, figs. 31–32; Hendler et al. 1995: 74, fig. 18. Asterinides folium — O’Loughlin 2002: 293, fig. 7; O’Loughlin & Waters 2004: 17, fig. 8; Oliveira et al. 2010: 3, fig. 2a; Bena- vides-Serrato et al. 2011: 150; Gondim et al. 2014: 24, fig. 7a–e; Sandino et al. 2017: S294; Souto & Martins 2017: 304; Cunha et al. 2020: 46. Material examined (8 specs, 2–7 mm R). BRAZIL. Bahia (12°47’– 13°05’S; 38°08’– 38°39’W)— Busca Vida beach, Guarajuba, 23 m, 1.ii.2006, 1 spec, R 2 mm (UFBA 685); 23 m, i.2010, 4 specs, R 2–2.6 mm (UFBA 983, 1076); 25 m, vii.2010, 1 spec, R 1.5 mm (UFBA 1163). Todos os Santos Bay, 12 m, 5.iv.1997, 1 spec, R 2 mm (UFBA 1107). Salvador, Itapuã beach, 1 m, 19.ii.2007, 1 spec, R 7 mm (UFBA 528). Comparative material. Asterinides pilosa: BRITISH VIRGIN ISLAND. Caribbean Sea, 9–12 m, 11.v.1976, 2 specs, R 3–3.8 mm (CASIZ 112487). Asterinides pompom: The Bahamas. Whitehorse Cay, 9 m, 28.vi.1998, 1 spec, R 13 mm (CASIZ 117432). Description of largest specimen (R 7 mm). Body stellate (Fig. 17A–D), disc inflated; R/r 1.4. Five arms. Abactinal plates regularly arranged, imbricated (Fig. 17C), with tufts of spinelets (up to 12 in proximal plates) in proximal edge, and thick crystalline bodies. Terminal plates naked. Madreporite large, suboval, sunk between proximal plates (Fig. 18B). Primary plates heart-shaped (Fig. 18A). Papular areas small, conspicuous, with one papula (Fig. 18B), forming a longitudinal series of six areas along each arm. Superomarginal plates with 15–20 slender spines. Inferomarginal plates with tufts of 17–20 spinelets projected outwards (Fig. 18E). Actinal plates with 2–3 pointed spines, central spine often longest (Fig. 18F). Three adambulacral spines, webbed, central spine often longest (Fig. 18D). Two to three subambulacral spines, webbed (Fig. 18D). Oral plates with six oral spines and four suboral spines, all webbed (Fig. 18C). Tube feet in two rows, sucking disc present. Pedicellariae absent. Ontogenetic variation (R 1.5–2.6 mm). Average R/r 1.4. Differs from the largest specimen by having a pentagonal shape; juxtaposed abactinal plates; primary plates rosette-shaped (Fig. 17E); a pair of papular area per interradius, in proximal region; few or no actinal spines; tufts of inferomarginal spines conspicuous; two oral spines. Also, the smallest specimen (R 1.5 mm) has only one spine per adambulacral plate. Coloration. Specimen in vivo has scarlet red abactinal region and white to yellowish actinal region, with orange edge. Specimens in ethanol are white to beige. Distribution. Bermuda, U.S.A. (FL), Gulf of Mexico, Mexico, The The Bahamas, Puerto Rico, Belize, Jamaica, Nicaragua, Panama, Colombia, Venezuela (Verrill 1915; Clark & Downey 1992; Hendler et al. 1995; Alvarado et al. 2008; Benavides-Serrato et al. 2011; Sandino et al. 2017). BRAZIL: Paraíba, Bahia, Trindade Island, Rio de Janeiro (Brito 1968, 1971; Tommasi 1970; Clark & Downey 1992; Alvarado & Solís-Marín 2013; Gondim et al. 2014; Souto & Martins 2017; Cunha et al. 2020). Depth. 0–25 m (Clark & Downey 1992; present paper). Biological notes. Specimens from Bahia live under rocks, in sandy and calcareous bottoms, often in isolation. Specimens from Trindade Island also live under rocks but in densities of 25 specimens per site (Brito 1968; 1971). Specimens from the Caribbean are also solitary and their coloration varies with size: juvenile specimens are white, intermediate specimens are red, yellow or yellowish, and adult specimens are blue and blue-green (Hendler et al. 1995). Souto & Martins (2017) reported that the sites from Camaçari, Bahia, are recruitment areas for A. folium populations since only small specimens were found at those sites; their coloration was not recorded by collectors. The larger specimen was collected in the shallow subtidal region. Syntype. NHMD 76237 [previously as ZMUC AST 69] (Tom Schiøtte, per. comm). Type locality. St. Thomas, U.S. Virgin Islands (Lütken, 1860). Remarks. Clark & Downey (1992), O’Loughlin (2002) and O’Loughlin & Waters (2004) reported additional intraspecific variation not observed here, including the presence of 4–6 arms (vs. 5 arms); up to 20 spinelets in the proximal plates (vs. up to 12); 2–5 spines on the actinal plates (vs. 2–3); 3–5 adambulacral spines (vs. three). The specimens described in these papers were up to R 19 mm (vs. up to R 7 mm in the present study) and although they do not mention ontogenetic variation, the higher number of spines and spinelets reported may be related to size. H.L. Clark (1933) noted that adult specimens have at least R 10 mm. Asterinides folium differs from A. pilosa by having five arms, rays not clearly defined, and up to five spines in actinal plates (vs. six arms, distinct rays and clusters of small spines), from A. hartmeyeri by having a fringe of 15–20 inferomarginal spinelets (vs. 6–10 spinelets), and from A. pompom by having a stellate shape and single papulae (vs. pentagonal shape with indistinct rays and paired papulae in the middle of the arm). Also, A. pompom has an inflated petaloid area in the abactinal region and a granulose structure with a cluster of thin spinelets in most abactinal plates. For a thorough revision of the genus Asterinides, see O’Loughlin & Waters (2004)., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 33-36, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Lutken, C. F. (1860) Bidrag til kundskab om de ved kysterne af Mellenog Sydde ved kysterne af Mellenog Sydundskab. Videnskabelige Meddelelser fra den Naturhistoriske Forening i Kjobenhavn, 1859, 25 - 97.","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Hendler, G., Muller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea Stars, Sea Urchins and Allies: Echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp.","O'Loughlin, P. M. (2002) New genus and species of southern Australian and Pacific Asterinidae (Echinodermata, Asteroidea). Memoirs of Museum Victoria, 59 (2), 277 - 296. https: // doi. org / 10.24199 / j. mmv. 2002.59.2","O'Loughlin, P. M. & Waters, J. M. (2004) A molecular and morphological revision of genera of Asterinidae (Echinodermata: Asteroidea). Memoirs of Museum Victoria, 61 (1), 1 - 40. https: // doi. org / 10.24199 / j. mmv. 2004.61.1","Oliveira, J. P., Oliveira, J. & Manso, C. L. C. (2010) Inventario da colecao de equinodermos do LABIMAR, Campus Prof °. Alberto Carvalho, Universidade Federal de Sergipe. Scientia Plena, 6, 1 - 14.","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Sandino, O. B., Solis-Marin, F. A., Caballero-Ochoa, A. A., Conejeros-Vargas, C. A., Marquez-Borras, F., Ayala-Aguilera, A. P. & Laguarda-Figueras, A. (2017) Equinodermos de Nicaragua: nuevos registros del Pacifico y Caribe Sur. Revista de Biologia Tropical, 65, 288 - 298. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31696","Souto, C. & Martins, L. (2017) Os equinodermos. In: Nunes, J. M. C. & Matos, M. R. B. (Eds.), Litoral Norte da Bahia: caracterizacao ambiental, biodiversidade e conservacao. Vol. 1. EDUFBA, Salvador, pp. 303 - 315.","Cunha, R., Tavares, M. & Mendonca, J. B. (2020) Asteroidea (Echinodermata) from shallow-waters of the remote oceanic archipelago Trindade and Martin Vaz, southeastern Atlantic, with taxonomic and zoogeographical notes. Zootaxa, 4742 (1), 31 - 56. https: // doi. org / 10.11646 / zootaxa. 4742.1.2","Verrill, A. E. (1915) Report on the Starfishes of the West Indies, Florida, and Brazil, Including Those Obtained by the Bahama Expedition from the University of Iowa in 1893. Bulletin of the State of University of Iowa, 7, 1 - 232, 29 pls. https: // doi. org / 10.5962 / bhl. title. 12035","Alvarado, J. J., Solis-Marin, F. A. & Ahearn, C. (2008) Equinodermos (Echinodermata) del Caribe Centroamericano. Revista de Biologia Tropical, 56, 37 - 55.","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Brito, I. M. (1968) Asteroiides e equinoiides do Estado da Guanabara e adjace ˆ ncias. Boletim do Museu Nacional, 260, 1 - 51.","Brito, I. M. (1971) Contribuicao ao conhecimento dos Equinodermas da Ilha da Trindade, Brasil. Arquivos do Museu Nacional, 54, 261 - 265.","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Vol. 1. Springer, Berlin, 658 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Clark, H. L. (1933) A handbook of the littoral echinoderms of Porto Rico and the other West Indian Islands. Scientific Survey of Puerto Rico, 16, 1 - 147."]}

Linckia guildingi Gray, 1840 Figures 22–23 Linckia guildingii Gray, 1840: 285. Linckia guildingii — Müller & Troschel 1842: 33; Rathbun 1879: 148; Clark 1938: 133; Tommasi 1958: 17; Brito 1962: 3; 1968: 4–5, pl. 1, fig. 3; 1971: 262; Lima-Verde 1969: 11; Tommasi 1970: 9, pl. 9, fig. 27; Tommasi & Aron 1988: 3; Tommasi et al. 1988: 6; Fernandes et al. 2002: 422; Gondim et al. 2008: 154; Carmo et al. 2015; Sandino et al. 2017: S294; Gurjão & Lotufo 2018: 11; Miranda 2018: 14; Patrizzi & Dobrovolski 2018: 182. Linckia guildingi — Clark & Downey 1992: 275, pl. 67, fig. 42; Hendler et al. 1995: 76, figs. 20–21; Pérez-Ruzafa et al. 1999: 47; Williams 2000; Entrambasaguas 2003: 101–106; Entrambasaguas, 2008: 63–64; Benavides-Serrato et al. 2011: 174; Miranda et al. 2012: 144; Gondim et al. 2014: 32, figs. 10a–e, 12d; Alvarado et al. 2017: S276; Souto & Martins 2017: 304–305, fig. 1D; Rubio-Polania et al. 2018: 190; Borrero-Peìrez et al. 2019: 5; Cunha et al. 2020: 38, fig. 5; Prata et al. 2020. Material examined (34 specs, 6–120 mm R). BRAZIL. Bahia (12°45’– 13°54’S; 38°37’– 38°58’W)— Salvador: Amaralina beach, intertidal, 8.iv.2008, 1 spec, R 6 mm (UFBA 627); Pituba beach, intertidal, 26.i.2006, 1 spec, R 45 mm (UFBA 169); intertidal, 4.xi.2010, 1 spec, R 57 mm (UFBA 1213); intertidal, 2.v.2011, 4 specs, R 50–74 mm (UFBA 1360); Itaparica Island, intertidal, 23.xi.1991, 2 specs, R 30–60 mm (UFBA 194); Ponta de Humaitá beach, intertidal, ii.2011, 2 specs, R 45–54 mm (UFBA 1269–1270). Barra Grande beach, Vera Cruz, 16 m, 2.x.2007, 1 spec, R 10 mm (UFBA 529). Frades Island, Ponta de Nossa Senhora, 3 m, 17.x.2008, 8 specs, R 76–104 mm (UFBA 674). Salvador: Farol da Barra beach, 17.i.2007, 1 spec, R 76 mm (UFBA 470); Itapuã beach, intertidal, 1.viii.2007, 1 spec, R 114 mm (UFBA 590); Pituba beach, intertidal, 16.iv.1991, 1 spec, R 95 mm (UFBA 591); intertidal, 5.vii.1997, 2 specs, R 105–120 mm (UFBA 41); intertidal, 2005, 1 spec, R 118 mm (UFBA 639); intertidal, 8.iv.2008, 1 spec, R 87 mm (UFBA 626); intertidal, 2.v.2011, 5 specs, R 82–113 mm (UFBA 1360); Ponta de Humaitá beach, intertidal, xii.2010, 1 spec, R 116 mm (UFBA 1268). Comparative material. BRAZIL. Alagoas, Ipioca coral reef, 1.vii.2007, 1 spec, R 35 mm (UFBA 533). WEST INDIES, 1 spec, R 24 mm (NHM-UK 1953.4.27.68, lectotype); 1 spec, R 15 mm (NHM-UK 1953.4.27.68, paralectotype). Description (R 76–120 mm). Small disc; average R/r 12.5. Five to six (rarely 1, 4 or 7) long, narrow, cylindri- cal arms (Fig. 22A–B). Abactinal surface with small, irregularly arranged, tumid plates covered by granules (Fig. 22C). Marginal plates larger than abactinal plates, tumid, covered by granules and arranged into two longitudinal rows. Papular areas large, only on abactinal surface, with 11–25 pores; number of pores lower near disc; actinal surface without papular areas. One to two madreporites (Fig. 22E) with deep furrows; only one per interradius.Anus inconspicuous. Terminal plates small, oval, covered by granules. Actinal plates (Fig. 22D) arranged into three rows extending almost or completely to tip of arm, covered by granules slightly larger than those on abactinal surface; plates near furrow larger than others. Two blunt adambulacral spines, proximal spine larger. Two rows of large, blunt subambulacral spines; outer spines granular-shaped. Innermost two spines on oral plates smaller than adjacent spines (Fig. 22F). Tube feet in two rows; sucking disc with perforated plates (Fig. 23G–H). Pedicellariae absent. Ontogenetic variation (R 6–74 mm). Average R/r 9.2. Terminal plates proportionally larger than that of large specimen; plate naked or only with a few granules (Fig. 23D). Papular areas small, with 1–5 pores. Madreporite proportionally smaller than that of large specimen, with few gyres (Fig. 23C). Marginal plates prominent in specimens up to R 15 mm. Second row of subambulacral spines in specimens up to R 55 mm almost same size as adjacent granules (Fig. 23E), making it hard to differentiate them. Innermost spines on oral plate prominent (Fig. 23F). Coloration. Specimens in vivo are whitish, pink or light brown; papular areas darker. Specimens in ethanol are beige to brown or light pink. Distribution. Circumtropical (Clark & Downey 1992; Alvarado & Solis-Marin 2013; Gondim et al. 2014; Cunha et al. 2020). BRAZIL: Pará, Paraíba, Pernambuco, Alagoas, Bahia, Espírito Santo, Trindade Island, Rio de Janeiro and São Paulo (Verrill 1868; Rathbun 1879; Verrill 1915; Brito 1960, 1968; Tommasi 1970; Tommasi & Aron 1988; Gondim et al. 2014; Carmo et al. 2015; Souto & Martins 2017; Miranda 2018; Cunha et al. 2020). Depth. 0–298 m (Clark & Downey 1992). Biological notes. Linckia guildingi is a nocturnal species, commonly found under rocks, in rock crevices, on coral reefs, rhodolith beds and in sandy bottoms from Bahia (Alves & Cerqueira 2000; Sampaio 2010; Prata et al. 2020; present paper). Specimens in the Northern Brazil have also been found in muddy bottoms (Miranda 2018). Juveniles of this species reproduce asexually by fission (Clark 1933) and are often found with arms of different sizes (note that our measurements were based on the largest arm). Martins et al. (2012) reported the commercial exploitation of this species for the aquarium trade, but the harvesting of L. guildingi in Brazil is currently prohibited (Gurjão & Lotufo 2018). This species is classified as “Vulnerable” (baseline data indicates that the population size is small [i.e., number of mature individuals per subpopulation is 1000 or less] and currently in decline) by the Ministry of the Environment (MMA 2018). Patrizzi & Dobrovolski (2018) predicted that the habitable range of L. guildingi may have a 10–28-fold expansion under higher atmospheric CO 2 concentrations. The effect of this expansion on the local communities is unknown, but it is likely to cause negative trophic impact (Kordas et al. 2011). Lectotype. NHM-UK 1953.4.27.68. Type locality. West Indies. Remarks. The data presented by Clark & Downey (1992) and the morphology of the type specimens (R 15–24 mm) support the ontogenetic variation described here. According to H.L. Clark (1933), adult individuals vary from R 75–215 mm. Linckia bouvieri and L. nodosa differ from L. guildingi by having smaller papular areas, secondary plates between the primary plates, and large abactinal plates (vs. large papular areas, secondary plates absent, and small abactinal plates). Also, the abactinal plates in L. bouvieri are flat (vs. tumid). Genus Narcissia Gray, 1840 Type species. Narcissia canariensis (d’Orbigny, 1839) (type by monotypy). Remarks. The genus Narcissia is composed of four species: Narcissia ahearnae Pawson, 2007 (from NW Atlantic), Narcissia canariensis (d’Orbigny, 1839) (from East Atlantic), Narcissia gracilis Clark, 1916 (from East Pacific) and Narcissia trigonaria Sladen, 1889 (from West Atlantic). The classification of Narcissia has been controversial since molecular data placed N. trigonaria in an unusual phylogenetic position, sister to a clade with goniasterid and ophidiasterid species (Mah & Foltz 2011). Here, we follow Mah (2020a) and keep Narcissia in the family Ophidiasteridae., Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 42-45, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Gray, J. E. (1840) A synopsis of the genera and species of the class Hypostoma (Asterias Linnaeus). Annals of the Magazine of Natural History, Series 1, 6 (37), 175 - 184 + 275 - 290. https: // doi. org / 10.1080 / 03745484009443296","Muller, J. & Troschel, F. H. (1842) System der Asteriden. Friedrich Viewig und Sohn, Braunschweig, 134 pp.","Rathbun, R. (1879) A list of the Brazilian echinoderms, with notes on their distribution. Transactions of the Connecticut Academy of Arts and Sciences, 5 (3), 139 - 158. https: // doi. org / 10.5962 / bhl. title. 16126","Clark, A. H. (1938) A new genus of starfishes from Puerto Rico. Smithsonian Miscellaneous Collections, 91 (29), 1 - 7, 1 pl.","Tommasi, L. R. (1958) Os equinodermos do litoral de Sao Paulo. II. Diadematidae, Schizasteridae, Brissidae, Cidaroidea (Echinoidea) e Asteroidea do Bentos Costeiro. Contribuicoes avulsas do Instituto Oceanografico, 2, 1 - 27, 6 pls.","Brito, I. M. (1962) Ensaio de catailogo dos equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zooloigicos, 13, 1 - 10.","Brito, I. M. (1968) Asteroiides e equinoiides do Estado da Guanabara e adjace ˆ ncias. Boletim do Museu Nacional, 260, 1 - 51.","Lima-Verde, J. S. (1969) Primeira contribuicao ao inventario dos equinodermas do Nordeste brasileiro. Arquivos de Ciencias do Mar, 9 (1), 9 - 13.","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Tommasi, L. R. & Aron, M. A. (1988) Equinodermos da plataforma continental do sudeste do estado da Bahia. Relatos do Instituto Oceanografico da Universidade de Sao Paulo, 19, 1 - 6.","Fernandes, M. L. B., Tommasi, L. R. & Lima, E. J. B. (2002) Filo Echinodermata de Pernambuco. In: Tabarelli, M. & Silva, J. M. C. (Eds.), Diagnostico da Biodiversidade de Pernambuco. Vol. 2. Massangana, Recife, pp. 405 - 427.","Gondim, A., Lacouth, P., Alonso, C. & Manso, C. (2008) Echinodermata da praia do Cabo Branco, JoaTo Pessoa, Paraiiba, Brasil. Biota Neotropica, 8, 151 - 159. https: // doi. org / 10.1590 / S 1676 - 06032008000200016","Carmo, A. V., Okada, N. B. & Freret-Meurer, N. V. (2015) Caracterizacao do microhabitat da estrela-do-mar Linckia guildingii Gray, 1840 que ocorre na Praia de Aracatiba, Ilha Grande, Angra dos Reis, Brasil. Revista Pensar Biologia, 1 (1), 22.","Sandino, O. B., Solis-Marin, F. A., Caballero-Ochoa, A. A., Conejeros-Vargas, C. A., Marquez-Borras, F., Ayala-Aguilera, A. P. & Laguarda-Figueras, A. (2017) Equinodermos de Nicaragua: nuevos registros del Pacifico y Caribe Sur. Revista de Biologia Tropical, 65, 288 - 298. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31696","Gurjao, L. M. & Lotufo, T. M. C. (2018) Native species exploited by marine aquarium trade in Brazil. Biota Neotropica, 18 (3), e 20170387. https: // doi. org / 10.1590 / 1676 - 0611 - bn- 2017 - 0387","Miranda, A. P. S. (2018) Diversidade e distribuicao de equinodermos da Plataforma Continental Amazonica. Unpublished Honors Thesis, Universidade Federal do Para, Belem, 62 pp.","Patrizzi, N. & Dobrovolski, R. (2018) Integrating climate change and human impacts into marine spatial planning: a case study of threatened starfish species in Brazil. Ocean & Coastal Management, 161, 177 - 188. https: // doi. org / 10.1016 / j. ocecoaman. 2018.05.003","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Hendler, G., Muller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea Stars, Sea Urchins and Allies: Echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp.","Perez-Ruzafa, A., Entrambasaguas, L. & Bacallado, J. J. (1999) Fauna de equinodermos (Echinodermata) de los fondos rocosos infralitorales del archipielago de Cabo Verde. Revista de la Academia Canaria de Ciencias, 11 (3 - 4), 43 - 62.","Williams, S. T. (2000) Species boundaries in the starfish genus Linckia. Marine Biology, 136 (1), 137 - 148. https: // doi. org / 10.1007 / s 002270050016","Entrambasaguas, L. (2003) Estudio de la fauna de equinodermos del archipielago de Cabo Verde: escalas de variabilidad espacial y factores que explican su distribucion. Tesis de licenciatura no publicada. Universidad de Murcia, Murcia. [unknown pagination]","Entrambasaguas, L. (2008) Estudio fauniistico y ecoloigico de los equinodermos del archipieilago de Cabo Verde. Tesis doctoral no publicada, Universidad de Murcia, Murcia, 315 pp.","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Miranda, A. L. S., Lima, M. L. F., Sovierzoski, H. H. & Correia, M. D. (2012) Inventory of the Echinodermata collection from the Universidade Federal de Alagoas. Biota Neotropica, 12 (2), 135 - 146. https: // doi. org / 10.1590 / S 1676 - 06032012000200014","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Alvarado, J. J., Chacon-Monge, J. L., Solis-Marin, F. A., Pineda-Enriquez, T., Caballero-Ochoa, A. A., Rivera, S. S. & Chaves, R. R. (2017) Equinodermos del Museo de Zoologia de la Universidad de Costa Rica. Revista de Biologia Tropical, Universidad de Costa Rica, 65 (1), S 272 - S 287. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31695","Souto, C. & Martins, L. (2017) Os equinodermos. In: Nunes, J. M. C. & Matos, M. R. B. (Eds.), Litoral Norte da Bahia: caracterizacao ambiental, biodiversidade e conservacao. Vol. 1. EDUFBA, Salvador, pp. 303 - 315.","Rubio-Polania, J. C., Torruco-Gomez, D., Gonzalez-Solis, A., Ordaz, J. & Caamal-Jimenez, Y. (2018) Benthic megafauna of outer margins of the continental shelf of Yucatan Peninsula. Regional Studies in Marine Science, 24, 184 - 195. https: // doi. org / 10.1016 / j. rsma. 2018.08.014","Cunha, R., Tavares, M. & Mendonca, J. B. (2020) Asteroidea (Echinodermata) from shallow-waters of the remote oceanic archipelago Trindade and Martin Vaz, southeastern Atlantic, with taxonomic and zoogeographical notes. Zootaxa, 4742 (1), 31 - 56. https: // doi. org / 10.11646 / zootaxa. 4742.1.2","Prata, J., Lucena, R. A., Lima, S. F. B., Weverton, J., Souza, S. & Christoffersen, M. L. (2020) Species richness of Pycnogonida and Echinodermata associated with the reef ecosystems of Morro de Sao Paulo on Tinhare Island in Northeastern Brazil. International Journal of Development Research, 10 (04), 34943 - 34951.","Alvarado, J. J. & Solis-Marin, F. A. (2013) Echinoderm research and diversity in Latin America. Vol. 1. Springer, Berlin, 658 pp. https: // doi. org / 10.1007 / 978 - 3 - 642 - 20051 - 9","Verrill, A. E. (1868) Notes of the radiate on the Museum of Yale College, with descriptions of genera and species. 4. Notice on the corals and echinoderms collected by Prof. C. F. Hartt at the Abrolhos Reefs, Province of Bahia, Brazil, 1867. Transactions of the Connecticut Academy of Arts and Sciences, 1 (2), 351 - 371.","Verrill, A. E. (1915) Report on the Starfishes of the West Indies, Florida, and Brazil, Including Those Obtained by the Bahama Expedition from the University of Iowa in 1893. Bulletin of the State of University of Iowa, 7, 1 - 232, 29 pls. https: // doi. org / 10.5962 / bhl. title. 12035","Brito, I. M. (1960) Asteroides dos estados do Rio de Janeiro e de Sao Paulo. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 5, 1 - 13.","Alves, O. F. S. & Cerqueira, W. R. P. (2000) Echinodermata das praias de Salvador (Bahia, Brasil). Revista Brasileira de Zoologia, 17 (2), 543 - 553. https: // doi. org / 10.1590 / S 0101 - 81752000000200024","Sampaio, F. L. M. (2010) Distribuicao espaco-temporal de mega equinodermos e reproducao assexuada de Linckia guildingii (Echinodermata: Asteroidea) no plato do recife de coroa vermelha, Bahia, Brasil. Unpublished Master Thesis, Universidade Estadual de Santa Cruz, Ilheus, Bahia, 33 pp.","Clark, H. L. (1933) A handbook of the littoral echinoderms of Porto Rico and the other West Indian Islands. Scientific Survey of Puerto Rico, 16, 1 - 147.","Martins, L., Souto, C., Magalhaes, W. F., Alves, O. F. S., Rosa, I. L. & Sampaio, C. L. S. (2012) Echinoderm harvesting in Todosos-Santos Bay, Bahia State, Brazil: the aquarium trade. Sitientibus serie Ciencias Biologicas, 12 (1), 53 - 59. https: // doi. org / 10.13102 / scb 123","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp.","Kordas, R. L., Harley, C. D. G. & O'Connor, M. I. (2011) Community ecology in warming world: the influence of temperature on interspecific interactions in marine systems. Journal of Experimental Biology and Ecology, 400, 218 - 226. https: // doi. org / 10.1016 / j. jembe. 2011.02.029","d'Orbigny, A. D. (1839) Echinodermes et Polypiers. In: Webb, P. B. & Berthelot, S. (Eds.), Histoire Naturelle des Iles Canaries, Zoologie. Vol. 2. Bethune, Paris, pp. 148 - 149.","Clark, A. H. (1916) Six new starfishes from the Gulf of California and adjacent waters. Proceedings of the Biological Society of Washington, 29, 51 - 62.","Sladen, W. P. (1889) Asteroidea. Report of the Scientific Results of H. M. S. Challenger, 30, 1 - 893. https: // doi. org / 10.1111 / j. 1096 - 3642.1882. tb 02281. x","Mah, C. & Foltz, D. (2011) Molecular phylogeny of the Valvatacea (Asteroidea: Echinodermata). Zoological Journal of the Linnean Society, 161 (4), 769 - 788. https: // doi. org / 10.1111 / j. 1096 - 3642.2010.00659. x","Mah, C. L. (2020 a) World Asteroidea Database. Available from: http: // www. marinespecies. org / asteroidea / (accessed 30 April 2020)"]}

Oreaster reticulatus (Linnaeus, 1758) Figures 28–29 Asterias reticulata Linnaeus, 1758: 661. Oreaster reticulatus — Tommasi 1970: 10–11, 36, pl. 13, fig. 31; Downey 1973: 60, pl. 24, figs. A–B; Clark & Downey 1992: 293, pl. 72; Hendler et al. 1995: 82, figs. 25–26; Fernandes et al. 2002: 422; Magalhães et al. 2005: 63; Martins & Queiroz 2006: 202–203; Ventura et al. 2007: 238; Manso et al. 2008: 185, fig. 8c–e; Magris & Deìstro 2010: 59, 61; Xavier 2010: 75; Alves & Dias 2010: 157; Benavides-Serrato et al. 2011: 179–180; Miranda et al. 2012: 143–144; Gondim et al. 2014: fig. 8a–g, 12c; Alvarado et al. 2017: S277; Sandino et al. 2017: S294; Souto & Martins 2017: 305, fig. 1C; Agostini & Ozorio 2018: 35; Gurjão & Lotufo 2018: 11; Patrizzi & Dobrovolski 2018: 182; Borrero-Peìrez et al. 2019: 5; Torres & Torres 2019: 413; Cunha et al. 2020: 44, figs. 4E, 8; Magris & Giarrizzo 2020: 3. Material examined (2 specs, 49–98 mm R). BRAZIL, Bahia (12°52’– 13°02’S; 38°40’– 38°37’W)— Itaparica Island, Ponta de Areia beach, 4.vi.1994, 1 spm R 49 mm (UFBA 468). Todos os Santos bay, 16 m, 22.v.1997, 1 spm, R 98 mm (UFBA 367). Comparative material. BRAZIL. Espírito Santo: Trindade Island, Enseada dos Portugueses (20°29’52.3”S; 29°19’15.6”W), 12.5 m, 23.x.2014, 1 spec, R 100 mm (MZUSP 1611); 12.6 m, 6.vii.2015, 1 spec, R 105 mm (MZUSP 1612). São Paulo (23º47’– 23º57’S; 45º23’– 46º20’W)—São Sebastião, 1 spec, R 125 mm (MZUSP 1956); Santos, 76 m, 1.vi.1999, 2 specs, R 130–140 mm (MZUSP 1617). Description (R 98 mm). Body stellate, disc inflated; R/r 1.7 (Fig. 28A–B). Five short arms, distally tapering (Fig. 28E–F). Abactinal plates reticulate, connected by elongated and narrow secondary plates. Abactinal region with granules, large papular pores, and robust tubercles and spines with blunt tips. Madreporite small, subcircular, covered with shallow gyres. Superomarginal plates large, covered by granules and with a tubercle or a short, robust spine. Inferomarginal plates large, confined to actinal surface, covered by granules and with a tubercle; distal plates with a short and robust spine. Actinal surface with tumid plates covered by granules. Interradial actinal area large. Adambulacral plate with 5–6 unequal spines, spines on extremities smallest. One robust subambulacral spine. Oral plates with five pairs of robust, prismatic spines; inner pair shorter and wider than others. Tube feet in two rows, sucking disc with many perforated plates. Bivalve pedicellariae on both surfaces, more numerous in actinal region, never in alveoli. Ontogenetic variation (R 49 mm). R/r 1.9 (Fig. 28C–D). Some of the abactinal primary plates form an elevated circle from which five rays radiate towards tip of arm (Fig. 28C); this pattern was not observed in the larger specimen. Also, the small specimen has five unequal adambulacral spines (Fig. 29H), being the proximal spine the smallest, and the third and fourth spines the largest, and the inner pair of spines in the oral plate is longer than the adjacent spines (Fig. 29I). Finally, the small specimen has fewer abactinal spines than the largest specimen, which are mostly confined to arms and interradial region (vs. throughout abactinal region), and fewer pedicellariae in the actinal region (Fig. 29E). Coloration. Specimens in vivo have beige to orange abactinal surface, beige to yellowish actinal region and sometimes red spots in the margins. Specimens in ethanol are beige to light brown. Distribution. U.S.A. (NC, FL), Gulf of Mexico, Mexico, The The Bahamas, Caribbean Sea, Cuba, Belize, Haiti, Dominican Republic, Puerto Rico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Surinam, Canary Islands, Cape Verde (Verrill 1915; Caso, 1944; Ummels 1963; Walenkamp 1976; Clark & Downey 1992; Hendler et al. 1995; Guzman & Guevara 2002; Entrambasaguas 2008; Hernandéz et al. 2013; Alvarado et al. 2017; Sandino et al. 2017; Borrero-Peìrez et al. 2019; Mah 2020a). BRAZIL: Amapá, Maranhão, Ceará, Paraíba, Pernambuco, Alagoas, Bahia, Trindade Island, Rio de Janeiro, São Paulo, Santa Catarina, Rio Grande do Sul (Rathbun 1879; Verrill 1915; Tommasi 1958, 1970; Brito 1960, 1962, 1968; Lima-Verde 1969; Walenkamp 1976; Fernandes et al. 2002; Magalhães et al. 2005; Ventura et al. 2007; Magris & Deìstro 2010; Xavier 2010; Miranda et al. 2012; Gondim et al. 2014; Souto & Martins 2017; Agostini & Ozorio 2018; Torres & Torres 2019; Cunha et al. 2020). Depth. 0–76 m (Clark & Downey 1992; Cunha et al. 2020). Biological notes. In Bahia, this species is found in protected, shallow waters, often in sandy bottoms with coarse sediment (Manso et al. 2008). Oreaster reticulatus used to be abundant in shallow waters (intertidal up to 5 m), but locals have reported population declines over the last 40 years; currently, this species is rare and found only in deeper regions. In addition to habitat degradation, especially because of urbanization, this population is affected by human exploitation, as O. reticulatus is commonly sold as souvenirs in tourist shops throughout the country. In November 2008, for example, the Brazilian Institute of the Environment and Renewable Natural Resources (IB- AMA) brought in 15– 20 specimens (Fig. 28 E–F) of O. reticulatus that they apprehended, to be identified (unpubl. data). Pinheiro et al. (2018) reported the intense harvesting of O. reticulatus in Espírito Santo, Bahia’s southern neighboring state, with commercial purposes. According to them, specimens are sold by fishermen for US $ 0.50. In Salvador, Bahia, illegally collected specimens have been sold at beaches for US $ 21 (ca. R$120,00) (data from a fisherman interviewed in October 2020; Fig. 29A). This species is also used in religious rituals (Alves & Dias 2010; Souto & Martins 2017), in ornamental aquaria (Martins et al. 2012) and as medicine to treat asthma, cold and tiredness (Alves & Rosa 2007; Alves et al. 2009; Alves & Dias 2010; Alves & Alves 2011; Lima 2018). However, the impact of these activities on O. reticulatus populations has not been studied and it is assumed to be low if compared to the commercial harvesting of this species. Franco et al. (2015) reported that extracts of O. reticulatus contain compounds capable of inhibiting the activity of the bacteria Staphylococcus aureus at low concentrations. Oreaster reticulatus is classified as “Vulnerable” (baseline data indicates that the population size has been reduced by at least 30% as a result of habitat degradation, exploitation and/or introduction of invasive species) by the Ministry of the Environment (MMA, 2018). This assessment was performed before the devastating oil spill recorded in the Brazilian coast in 2019, which may have a strong impact in the coastal populations of O. reticulatus (Magris & Giarrizzo 2020). According to Gurjão & Lotufo (2018), the harvesting of this species in Brazil is currently prohibited. Using models, Patrizzi & Dobrovolski (2018) predicted that the habitable range of O. reticulatus may have a 7–16-fold expansion under higher atmospheric CO 2 concentrations. The effect of this expansion on the local communities is unknown, but it is likely to cause negative trophic impact (Kordas et al. 2011). Holotype. NHMD 76271 [previously as ZMUC AST 104] (Tom Schiøtte, per. comm). Type locality. East Caribbean (as Spanish West Indies) (Tom Schiøtte, per. comm). Remarks. The specimens examined here as comparative material (R 100–140 mm) have 5–7 adambulacral spines (vs. five in the specimen from Bahia, R 49 mm) and the largest specimens (R 130–140 mm) have two subambulacral spines (vs. one in specimens R 49–105 mm). Gondim et al. (2014) found 5–6 adambulacral spines in specimens ranging from R 51–136 mm, but they did not report if the largest specimens had six spines or if there was no trend in this variation. Also, in specimens with R 100–140 mm, the abactinal and actinal spines are equally developed; however, in the specimen described here, the actinal spines are less developed. Finally, the large specimens do not display the pattern (i.e. elevated circle with rays) observed in the abactinal surface of the small specimen. H.L. Clark (1933) noticed that small specimens have a deep olive to green abactinal region and as the size increases, the color changes from yellowish– to deep red. Downey (1973) described intraspecific variation (i.e. variation in shape, in degrees of inflation of the disc, and in coloration) in specimens ranging between R 15–91 mm, however, she did not specify ontogenetic changes. We observed that the presence and abundance of abactinal pedicellariae vary between specimens, but this variation does not seem to be related to growth. Also, Downey (1973) noted that O. reticulatus may have 4–7 arms, but both specimens examined here and the comparative material have five arms. Oreaster reticulatus differs from O. clavatus by having an inflated disc (vs. slightly flattened disc), abactinal plates with tubercles or spines (vs. abactinal plates with granules), and actinal pedicellariae not in alveoli (vs. actinal pedicellariae in alveoli) (Clark & Downey 1992). Order Forcipulatida Perrier, 1884, Published as part of Cunha, Rosana, Martins, Luciana, Menegola, Carla & Souto, Camilla, 2021, Taxonomy of the sea stars (Echinodermata: Asteroidea) from Bahia State, including ontogenetic variation and an illustrated key to the Brazilian species, pp. 1-78 in Zootaxa 4955 (1) on pages 51-55, DOI: 10.11646/zootaxa.4955.1.1, http://zenodo.org/record/4691078, {"references":["Linnaeus, C. (1758) Systema Naturae. Impensis Direct, Laurentii Salvii, Holmiae, 824 pp.","Tommasi, L. R. (1970) Lista dos asteroides recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, Universidade de Sao Paulo, 18, 1 - 61, 38 pls.","Downey, M. E. (1973) Starfishes from the Caribbean and the Gulf of Mexico. Smithsonian Contributions Zoology, 126, 1 - 158. https: // doi. org / 10.5479 / si. 00810282.126","Clark, A. M. & Downey, M. E. (1992) Starfishes of the Atlantic. Chapman and Hall, London, 794 pp. https: // doi. org / 10.5860 / choice. 30 - 2659","Hendler, G., Muller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea Stars, Sea Urchins and Allies: Echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp.","Fernandes, M. L. B., Tommasi, L. R. & Lima, E. J. B. (2002) Filo Echinodermata de Pernambuco. In: Tabarelli, M. & Silva, J. M. C. (Eds.), Diagnostico da Biodiversidade de Pernambuco. Vol. 2. Massangana, Recife, pp. 405 - 427.","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 9 (1), 61 - 65. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Martins, I. X. & Queiroz, A. C. M. (2006) Equinodermos do litoral do estado do Ceara. In: Matthews-Cascon, H. & Lotufo, T. M. C. (Eds.), Biota marinha da costa oeste do Ceara. Ministerio do Meio Ambiente, Serie Biodiversidade, Brasilia, pp. 200 - 219.","Ventura, C. R. R., Verissimo, I., Nobre, C. C. & Zama, P. C. (2007) Filo Echinodermata. In: Lavrado, H. P. & Viana, M. S. (Eds.), Atlas de invertebrados marinhos da regiao central da Zona Economica Exclusiva brasileira. Parte 1. Museu Nacional do Rio de Janeiro, Serie Livros 25. Museu Nacional do Rio de Janeiro, Rio de Janeiro, pp. 258.","Manso, C. L. C., Alves, O. F. S. & Martins, L. R. (2008) Echinodermata da Baia de Todos os Santos e da Baia de Aratu (Bahia, Brasil). Biota Neotropica, 8 (3), 179 - 196. https: // doi. org / 10.1590 / S 1676 - 06032008000300017","Xavier, L. A. R. (2010) Inventario dos equinodermos do Estado de Santa Catarina, Brasil. Brazilian Journal of Aquatic Science and Technology, 14, 73 - 78. https: // doi. org / 10.14210 / bjast. v 14 n 2. p 73 - 78","Alves, R. N. & Dias, T. L. P. (2010) Uso de invertebrados na medicina popular no Brasil e suas implicacoes para conservacao. Tropical Conservation Science, 3 (2), 159 - 174. https: // doi. org / 10.1177 / 194008291000300204","Benavides-Serrato, M., Borrero-Perez, G. & Diaz-Sanchez, C. (2011) Equinodermos del caribe colombiano: Crinodea, Asteroidea y Ophiuroidea. Vol. 1. Marquillas, Santa Marta, 384 pp.","Miranda, A. L. S., Lima, M. L. F., Sovierzoski, H. H. & Correia, M. D. (2012) Inventory of the Echinodermata collection from the Universidade Federal de Alagoas. Biota Neotropica, 12 (2), 135 - 146. https: // doi. org / 10.1590 / S 1676 - 06032012000200014","Gondim, A., Christoffersen, M. & Dias, T. (2014) Taxonomic guide and historical review of starfishes in northeastern Brazil (Echinodermata, Asteroidea). ZooKeys, 449, 1 - 56. https: // doi. org / 10.3897 / zookeys. 449.6813","Alvarado, J. J., Chacon-Monge, J. L., Solis-Marin, F. A., Pineda-Enriquez, T., Caballero-Ochoa, A. A., Rivera, S. S. & Chaves, R. R. (2017) Equinodermos del Museo de Zoologia de la Universidad de Costa Rica. Revista de Biologia Tropical, Universidad de Costa Rica, 65 (1), S 272 - S 287. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31695","Sandino, O. B., Solis-Marin, F. A., Caballero-Ochoa, A. A., Conejeros-Vargas, C. A., Marquez-Borras, F., Ayala-Aguilera, A. P. & Laguarda-Figueras, A. (2017) Equinodermos de Nicaragua: nuevos registros del Pacifico y Caribe Sur. Revista de Biologia Tropical, 65, 288 - 298. https: // doi. org / 10.15517 / rbt. v 65 i 1 - 1.31696","Souto, C. & Martins, L. (2017) Os equinodermos. In: Nunes, J. M. C. & Matos, M. R. B. (Eds.), Litoral Norte da Bahia: caracterizacao ambiental, biodiversidade e conservacao. Vol. 1. EDUFBA, Salvador, pp. 303 - 315.","Agostini, V. O. & Ozorio, C. P. (2018) Caracterizacao da macrofauna de substratos consolidados do Litoral Norte do Rio Grande do Sul, Brasil. Arquivos de Ciencias do Mar, 51 (1), 26 - 48. https: // doi. org / 10.32360 / acmar. v 51 i 1.31293","Gurjao, L. M. & Lotufo, T. M. C. (2018) Native species exploited by marine aquarium trade in Brazil. Biota Neotropica, 18 (3), e 20170387. https: // doi. org / 10.1590 / 1676 - 0611 - bn- 2017 - 0387","Patrizzi, N. & Dobrovolski, R. (2018) Integrating climate change and human impacts into marine spatial planning: a case study of threatened starfish species in Brazil. Ocean & Coastal Management, 161, 177 - 188. https: // doi. org / 10.1016 / j. ocecoaman. 2018.05.003","Torres, V. S. & Torres, F. S. S. (2019) Inventario preliminar dos invertebrados marinhos observados na praia da Pinheira, Palhoca, SC, Brasil. UNISANTA Bioscience, 8 (4), 408 - 422.","Cunha, R., Tavares, M. & Mendonca, J. B. (2020) Asteroidea (Echinodermata) from shallow-waters of the remote oceanic archipelago Trindade and Martin Vaz, southeastern Atlantic, with taxonomic and zoogeographical notes. Zootaxa, 4742 (1), 31 - 56. https: // doi. org / 10.11646 / zootaxa. 4742.1.2","Magris, R. A & Giarrizzo, T. (2020) Mysterious oil spill in the Atlantic Ocean threatens marine biodiversity and local people in Brazil. Marine Pollution Bulletin, 153, 1 - 3. https: // doi. org / 10.1016 / j. marpolbul. 2020.110961","Verrill, A. E. (1915) Report on the Starfishes of the West Indies, Florida, and Brazil, Including Those Obtained by the Bahama Expedition from the University of Iowa in 1893. Bulletin of the State of University of Iowa, 7, 1 - 232, 29 pls. https: // doi. org / 10.5962 / bhl. title. 12035","Caso, M. E. (1944) Estudio sobre asteridos de Mexico. Algunas especies interesantes de asteridos litorales. Anales del Instituto de Biologia Universidad Nacional Autonoma de Mexico, 15, 237 - 257.","Ummels, F. (1963) Asteroids from the Netherlands Antilles and other Carribbean localities (Oreasteridae, Ophidiasteridae, Asterinidae and Luidiidae). Studies on the Fauna of Curacao and Other Caribbean Islands, 15, 72 - 101, 3 figs., 9 pls.","Walenkamp. J. H. C. (1976) The asteroids of the coastal waters of Surinam. Zoologische Verhandelingen, 147, 1 - 91, 18 pls.","Guzman, H. M. & Guevara, C. A. (2002) Annual reproductive cycle, spatial distribution, abundance, and size structure of Oreast- er reticulatus (Echinodermata: Asteroidea) in Bocas del Toro, Panama. Marine Biology, 141, 1077 - 1084. https: // doi. org / 10.1007 / s 00227 - 002 - 0898 - 2","Entrambasaguas, L. (2008) Estudio fauniistico y ecoloigico de los equinodermos del archipieilago de Cabo Verde. Tesis doctoral no publicada, Universidad de Murcia, Murcia, 315 pp.","Mah, C. L. (2020 a) World Asteroidea Database. Available from: http: // www. marinespecies. org / asteroidea / (accessed 30 April 2020)","Rathbun, R. (1879) A list of the Brazilian echinoderms, with notes on their distribution. Transactions of the Connecticut Academy of Arts and Sciences, 5 (3), 139 - 158. https: // doi. org / 10.5962 / bhl. title. 16126","Tommasi, L. R. (1958) Os equinodermos do litoral de Sao Paulo. II. Diadematidae, Schizasteridae, Brissidae, Cidaroidea (Echinoidea) e Asteroidea do Bentos Costeiro. Contribuicoes avulsas do Instituto Oceanografico, 2, 1 - 27, 6 pls.","Brito, I. M. (1960) Asteroides dos estados do Rio de Janeiro e de Sao Paulo. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 5, 1 - 13.","Brito, I. M. (1962) Ensaio de catailogo dos equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zooloigicos, 13, 1 - 10.","Brito, I. M. (1968) Asteroiides e equinoiides do Estado da Guanabara e adjace ˆ ncias. Boletim do Museu Nacional, 260, 1 - 51.","Lima-Verde, J. S. (1969) Primeira contribuicao ao inventario dos equinodermas do Nordeste brasileiro. Arquivos de Ciencias do Mar, 9 (1), 9 - 13.","Pinheiro, F. C. F., Pinheiro, H. C., Costa, T. J. F, Teixeira, J. B., Gasparini, J. L., Joyeux, J. C. & Martins, A. S. (2018) Harvest of endangered marine invertebrates in a priority area for conservation in Brazil. Nature Conservation Research, 3 (4), 78 - 81. https: // doi. org / 10.24189 / ncr. 2018.050","Martins, L., Souto, C., Magalhaes, W. F., Alves, O. F. S., Rosa, I. L. & Sampaio, C. L. S. (2012) Echinoderm harvesting in Todosos-Santos Bay, Bahia State, Brazil: the aquarium trade. Sitientibus serie Ciencias Biologicas, 12 (1), 53 - 59. https: // doi. org / 10.13102 / scb 123","Alves, R. R. N. & Rosa, I. L. (2007) Biodiversity, traditional medicine and public health: where do they meet? Journal of Ethnobiology and Ethnomedicine, 3, 1 - 9. https: // doi. org / 10.1186 / 1746 - 4269 - 3 - 14","Alves, R. N., Neto, N. L., Brooks, S. E. & Albuquerque, U. P. (2009) Commercialization of animal-derived remedies as complementary medicine in the semi-arid region of Northeastern Brazil. Journal of Ethnopharmacology, 124, 600 - 608. https: // doi. org / 10.1016 / j. jep. 2009.04.049","Alves R. R. & Alves H. N. (2011) The faunal drugstore: animal-based remedies used in traditional medicines in Latin America. Journal of Ethnobiology and Ethnomedicine, 10, 1746 - 4269. https: // doi. org / 10.1186 / 1746 - 4269 - 7 - 9","Lima, R. J. P. (2018) Uso de Zooterapicos na Zona Rural do Municipio de Jacana, Semiarido Potiguar. Unpublished Honors Thesis, Instituto Federal de Educacao, Ciencia e Tecnologia da Paraiba, Joao Pessoa, Paraiba, 34 pp.","Franco, O. P., Patino, G. S. & Ortiz, A. A. (2015) Actividad antibacteriana y antifungica de la estrella de mar Oreaster reticulatus (Valvatida: Oreasteridae) y de los erizos de mar Mellita quinquiesperforata (Clypeasteroida: Mellitidae) y Diadema antillarum (Diadematoida: Diadematidae) del Caribe Colombiano. Revista de Biologia Tropical, 63, 329 - 337.","MMA (Ministerio do Meio Ambiente) (2018) Livro Vermelho da Fauna Brasileira Ameacada de Extincao: Invertebrados. IC- MBio / MMA, Brasilia, 727 pp.","Kordas, R. L., Harley, C. D. G. & O'Connor, M. I. (2011) Community ecology in warming world: the influence of temperature on interspecific interactions in marine systems. Journal of Experimental Biology and Ecology, 400, 218 - 226. https: // doi. org / 10.1016 / j. jembe. 2011.02.029","Clark, H. L. (1933) A handbook of the littoral echinoderms of Porto Rico and the other West Indian Islands. Scientific Survey of Puerto Rico, 16, 1 - 147.","Perrier, E. (1884) Memoire sur les etoiles de mer recueillies dans la mer des Antilles et le golfe du Mexique: durant les expeditions de dragace faites sous la direction de M. Alexandre Agassiz. Archives Museum National d'histoire Naturelle Paris, 6 (2), 127 - 276. https: // doi. org / 10.5962 / bhl. title. 82184"]}

Cassidulus caribaearum Lamarck, 1801 (Figs. 4���6) Cassidulus cariboearum Lamarck, 1801: 348 ���349. Rhyncholampas caribaearum ��� A. Agassiz, 1869:270 ���271. Rhynchopygus caribaearum ��� A. Agassiz, 1872 ���1874:153, 343, pl. 15. Rhyncholampas cariboearum ��� H.L. Clark, 1917:106, pl. 144. Cassidulus caribaearum ��� Mortensen, 1948a: 205 ���210, pls. 2, 11; Gray, 1855:34; Mooi, 1990b: 80. Cassidulus cariboearum ��� Kier, 1962: 176 ���178, pl. 26. Type material. Neotype (designated herein): Caribbean Sea, British Virgin Islands, Anegada, Loblolly Bay, 0.5��� 1 m, collected by R. Mooi and M. Telford, 01.IV.1986, 26.68 mm TL (CASIZ 222205). Neotype choice. In the absence of specimens from the type locality, we chose an adult specimen from one of the few well-studied populations of C. caribaearum. The specimen agrees with the original description of the species and is in great condition (i.e., test is intact and still has spines and pedicellariae). Comparative material examined. Cassidulus caribaearum Bahamas, TL 17 mm (NHM-UK 87.6.27.7); British Virgin Islands, Anegada, TL 9���30 mm (CASIZ 112632, 112633, 112637, 112638, 112683 A); and French Antilles, TL 8���20 mm (UF 11786���11788, 11797���11798, 11825, 11892, 11933). Description. Test oval (TW ca. 85% of TL), lateral edges straight, anterior and posterior margins round; greatest height at apical disc, test height decreases sharply from apical disc to periproct; transverse cross section triangular; oral region concave along the midline of the anterior-posterior axis. Test measurements (TL * TW * TH in mm): 26.68 * 22.65 * 11.71 (Fig. 4A���B, D���E). Apical disc anterior, monobasal, ca. 11% of TL, flat, with four elongated gonopores on disc edge; anterior gonopores closer together, posterior gonopores further apart (Fig. 4A; 4F, not from N); hydropores abundant and spread across plate. Anterior ocular plates between their adjacent gonopores; posterior ocular plates near distal edge of gonopores 1 and 4. Petalodium system about 74% of TL. Posterior petals longer than others; anterior petals wider than posterior petals. Anterior petals broad in the middle and narrow distally (not closing), and with bowed columns of respiratory podia. Posterior petals roughly with same width throughout; columns of respiratory podia curved outwards towards test edge. Poriferous zone of petals narrow, pores slightly conjugated; outer pores slightly elongated, inner pores round and smaller than outer. Columns a and b of posterior petals differ by 3���5 pore-pairs (number of respiratory podia: petal I, 27/30; petal V, 27/32), of anterior paired petals differ by 3���4 pore-pairs (number of respiratory podia: petal II, 20/23; petal IV, 20/24), anterior petal is equal (petal III, 25/25 respiratory podia). Primary tubercles present in poriferous zone; 2, sometimes 3 primary tubercles per petal plate. No occluded plates in petals. Ambulacra beyond petals increase 55% in relation to end-petal W; unipores in plates beyond petals: aboral plates wider than long, pores on suture, between adradial edge and middle of plate. Phyllodes with unipores, with only one column of pores on each half (number of phyllopores: phyllodes 1 and 5, 4/5; phyllodes 2 and 4, 6/6; phyllode 3, 6/5) (Fig. 4C). Columns mostly straight, largest W adorally and phyllodes narrow down distally. Buccal pores same size as phyllopores. Four to 6 sphaeridia (109���116 ��m L; Fig. 5L) in large and enclosed pits near buccal pores. Peristome anterior (34% TL from anterior region), slightly transverse (L 68% of W), pentagonal. Mouth opening in center of peristomial membrane. Bourrelets slightly developed as mounds mostly towards the inside of the peristome. Naked zone well-developed throughout midline of test on interambulacrum 5 and ambulacrum III, wider anteriorly than posteriorly (W 22% of TW). Deep pits present in oral region, mostly on naked zone. One primary tubercle present on naked zone, adjacent to phyllode 1. Periproct marginal and transverse (L 70% of W), beyond posterior petals; aboral plates form a prominent lip, oral and lateral plates do not bend inside the periproct (Fig. 4E; 4I ���J, not from N). Periproctal membrane with 2 rows of 5���6 medium- to large-sized plates; smaller plates scattered in aboral region near anus (Fig. 4I ���J, not from N). Anus opening aborally, on center of periproctal membrane and surrounded by elongated papillae. Primary tubercles perforate and slightly crenulate. Oral primary tubercles with mamelon displaced in the opposite direction of the spine (usually anteriorly from center of bosses) and 2.25x as large as aboral tubercles. Aboral spines short, straight, apex thicker than base, shaft serrated (Fig. 5C); bourrelet spines curved, shaft serrated, base short (Fig. 5D); oral spines long, straight, thicker at the base than at the apex, shaft mostly smooth (Fig. 5E; not from N); spines on lip above periproct thin, straight, intermediate in size between oral and aboral spines, shaft slightly serrated on both extremities. Miliary spines abundant, short, straight, thickness uniform, tip crown-shaped, shaft serrated (Fig. 5F, not from N). Periproctal and peristomial membranes with small ossicles. Triphyllous pedicellariae small and apparently abundant near aboral ambulacra. Valves short and broad (45���56 ��m L; Fig. 6A���B) with delicate teeth along whole blade margin. Ophicephalous pedicellariae small and rare. Valves (135���181 ��m L; Fig. 6C���D) with coarse teeth along whole blade margin (open row, U-shaped). Hinge broad, handles conspicuous. Two types of tridentate pedicellariae found. Large type found near posterior petals and periproct; valves (302���407 ��m L; Fig. 6E���H) broad and robust; blade with thin teeth along whole margin, teeth on distal margin numerous and very long; region between blade and basal region wide, basal region tall; stalk (494���553 ��m L; Fig. 5G) robust, proximal region thicker than distal region and with pointy upward projections (Fig. 5H), no neck present. Small type found around apical disc where embryos and young (Fig. 5A���B) are; valves (228���302 ��m L; Fig. 6I ���L) broad; blade with coarse teeth along whole margin, teeth on distal margin larger; region between blade and basal region narrow, basal region short and spineless; stalk (610���767 [broken] ��m L; Fig. 5I) long and thin, base thicker than apex, stereom thin and intricate (Fig. 5J), neck present. Additional information based on comparative material and literature. Posterior region of apical disc slightly bulged towards interambulacrum. Oral plates beyond petals about 1.15���1.60x longer than wide, pores near the distal edge of plate suture. Plates on phyllodes sometimes unequal in size and shape; pyrinoid plates not present. Phyllopores usually aligned in a uniform column. Ambulacral basicoronal plates pierced by buccal pore and one per ambulacrum also pierced by a phyllopore in the sequence a, b, a, b, b from phyllode I to V (Fig. 4H). Columns a and b of interambulacrum 5 with 6���7 plates between basicoronal plate and base of periproct and 3 (rarely 4) until adapical region of periproct (i.e., periproct is framed by 4 [rarely 3] plates on each side). Interambulacral basicoronal plates 1 and 4 narrower than others, 5 the broadest, 2 and 3 intermediate in size. Second and third oral plates on interambulacrum 5 are much longer than wide. Periproct framed adorally by plates 5.a.6 (or 5.a.7) and 5.b.7, and adapically by plates 5.a.10 (or 5.a.9) and 5.b.10. Gastric caecum highly reduced (Ziegler et al., 2008). Anterior stomach located in ambulacrum III, where it joins the short esophagus, and in interambulacrum 3; a cluster of 4���6 smooth, finger-like pouches at the junction of esophagus and anterior stomach, and further subdivided into two smaller clusters directed laterally towards interambulacra 2 and 3 (Ziegler et al., 2010). Color in vivo: white (Kier, 1975). Intraspecific variability. Besides the difference in the number of plates on oral interambulacrum 5, other differences were observed between specimens. For instance, the difference in the number of respiratory podia between the columns a and b of anterior paired petals varies from 2���4 pore-pairs. Also, the number of phyllopores varies from 4���5 in phyllodes I, III and V, and from 5���7 in phyllodes II and IV; and although most specimens have deep pits in the naked region, a few others have shallow and small pits. Bathymetric and geographic distribution. Cassidulus caribaearum has been recorded in Belize, the British Virgin Islands, Jamaica, Mexico, Panama and Puerto Rico (Lamarck, 1801; Gladfelter, 1978; Kier, 1975; Tzompantzi et al., 1999; Alvarado et al., 2008; Rodr��guez-Barreras et al., 2012), Bahamas and French Antilles (extended herein). The NHM-UK database holds occurrences from Antigua and Barbuda. Occurrence of the species in the Dominican Republic has been indicated by Alvarado (2011), but we have not been able to find the original reference for the occurrence nor a record in museum databases. This species occurs in shallow water, up to 18 m of depth (Mooi, 1990b; extended herein). A. Agassiz���s (1872 ��� 1874, p. 343) record of specimens living at 106 fathoms (ca. 194 m) of depth is probably wrong. Natural history notes. Even though C. caribaearum was described more than 200 years ago and is fairly welldistributed in shallow Caribbean waters, information about this species is still scarce. The specimens analyzed herein were collected from 10���100 mm beneath the surface of the sediment, which was composed of clean calcareous sand. This specific population migrates with changes in season and the density of individuals varies between 5���25 individuals/m2 (Gladfelter, 1978). Some cassiduloids are known to brood their young (Mortensen, 1948a), sometimes in brooding pouches or among the aboral spines, as is the case for C. caribaearum. The neotype described herein has several embryos and young in the aboral region (see dark yellow patch around apical disc on Fig. 4A). Gladfelter (1978) suggested that fertilization in this species may be internal and he also did not rule out the possibility of parthenogenesis; both hypotheses remain to be tested. Sexual dimorphism in gonopore size is not as apparent as in other cassiduloids (e.g., Neolampas rostellata A. Agassiz, 1869) but because of the presence of embryos on the test, the neotype is most likely a female. Sexual maturity was observed in individuals larger than 10 mm in TL. See Gladfelter (1978) and Telford & Mooi (1996) for additional information on the natural history of C. caribaearum., Published as part of Souto, Camilla & Martins, Luciana, 2018, Synchrotron micro-CT scanning leads to the discovery of a new genus of morphologically conserved echinoid (Echinodermata: Cassiduloida) in Zootaxa 4457 (1), DOI: 10.11646/zootaxa.4457.1.3, http://zenodo.org/record/1457567, {"references":["Lamarck, J. B. P. A. de M. de (1801) Systeme des animaux sans vertebres, ou tableau general des classes, des ordres et des genres de ces animaux; presentant leurs caracteres essentiels et leur distribution, d'apres la consideration de leurs rapports naturels et de leur organisation, et suivant l'arrangement etabli dans les galeries du Museum d'hist. naturelle, parmi leurs depouilles conservees; precede du discours d'ouverture de cours de zoologie donne dans le Museum national d'histoire naturelle l'an 8 de la Republique. Chez l'auteur & Deterville, Paris, 432 pp.","Agassiz, A. (1869) Preliminary report on the echini and star-fishes dredged in deep water between Cuba and the Florida Reef. Bulletin of the Museum of Comparative Zoology, 1 (9), 253 - 308.","Agassiz, A. (1872 - 1874) Revision of the echini. Memoirs of the Museum of Comparative Zoology, 3, 383 - 762.","Clark, H. L. (1917) Hawaiian and other Pacific echini. The Echinoneidae, Nucleoltitidae, Urechinidae, Echinocorythidae, Calymnidae, Pourtalesiidae, Palaeostomatidae, Aeropsidae, Palaeopneustidae, Hemiasteridae, and Spatangidae. Memoirs of the Museum of Comparative Zoology, 46 (2), 91 - 283.","Mortensen, T. (1948 a) A Monograph of the Echinoidea. IU. 1. Holectypoida, Cassiduloida. C. A. Reitzel, Copenhagen, 363 pp.","Gray, J. E. (1855) Catalogue of the Recent Echinida, or sea eggs, in the collection of the British Museum. Part I. Echinida Irregularia. Order of the Trustees, London, 69 pp.","Mooi, R. (1990 b) Living cassiduloids (Echinodermata: Echinoidea): a key and annotated list. Proceedings of the Biological Society of Washington, 130 (1), 63 - 85.","Kier, P. M. (1962) Revision of the cassiduloid echinoids. Smithsonian Miscellaneous Collections, 144, 1 - 262.","Ziegler, A., Faber, C., Mueller, S. & Bartolomaeus, T. (2008) Systematic comparison and reconstruction of sea urchin (Echinoidea) internal anatomy: a novel approach using magnetic resonance imaging. BMC Biology, 6, 33. https: // doi. org / 10.1186 / 1741 - 7007 - 6 - 33","Ziegler, A., Mooi, R., Rolet, G. & De Ridder, C. (2010) Origin and evolutionary plasticity of the gastric caecum in sea urchins (Echinodermata: Echinoidea). BMC Evolutionary Biology, 10, 313. https: // doi. org / 10.1186 / 1471 - 2148 - 10 - 313","Kier, P. M. (1975) The echinoids of Carrie Bow Cay, Belize. Smithsonian Contributions to Zoology, 206, 1 - 45. https: // doi. org / 10.5479 / si. 00810282.206","Gladfelter, W. B. (1978) General ecology of the cassiduloid urchin Cassidulus caribbearum. Marine Biology, 47, 149 - 160. https: // doi. org / 10.1007 / BF 00395636","Tzompantzi, D. B., Solis-Marin, F. A., Laguarda-Figueras, A., Abreu-Perez, M. & Duran-Gonzalez, A. (1999) Echinoids (Echinodermata: Echinoidea) from the Mexican Caribbean: Puerto Morelos, Quintana Roo, Mexico. Avicennia, 10 / 11, 43 - 72.","Alvarado, J. J., Solis-Marin, F. A. & Ahearn, C. (2008) Equinodermos (Echinodermata) del Caribe Centroamericano. Revista de Biologia Tropical, 56 (Supplement 3), 37 - 55.","Rodriguez-Barreras, R., Sabat, A. M., Benavides-Serrato, M. & Bontemps, D. R. (2012) A new record for Puerto Rico of the irregular echinoid Cassidulus caribaearum. Marine Biodiversity Records, 5, e 85.","Alvarado, J. J. (2011) Echinoderm diversity in the Caribbean Sea. Marine Biodiversity, 41 (2), 261 - 285. https: // doi. org / 10.1007 / s 12526 - 010 - 0053 - 0","Telford, M. & Mooi, R. (1996) Podial particle picking in Cassidulus caribaearum (Echinodermata: Echinoidea) and the phylogeny of sea urchin feeding mechanisms. Biological Bulletin, 191, 209 - 223. https: // doi. org / 10.2307 / 1542924"]}

Cassidulus briareus sp. nov. (Figs. 1���2, 10) Type material. Holotype: Australia (Neuholland), collected by Salmin (probably Carl Ludwig Salmin), 18.36 mm TL (MP 1267 Holotype MNHWU). Paratype: same collection data as H, 18.20 mm TL (MP 1267 Paratype MNHWU). Comparative material of other species examined. Cassidulus caribaearum ? Lamarck, 1801: possibly Caribbean Sea (MNHN-IE-2013-10590 [Syntype?]); Anegada, British Virgin Islands, TL 18���30 mm (CASIZ 112633, 112638, 112683 A���B). Cassidulus infidus Mortensen, 1948b: Salvador���BA, Brazil, TL 7.5���13 mm (SMNH 4859 [H], UFBA 314, 757). Cassidulus mitis Krau, 1954: Rio de Janeiro���RJ, Brazil, TL 26���46 mm (CASIZ 116110, MNRJ 3673���3674, ZUEC 11���12). Diagnosis. Test arched in cross section, greatest height at apical disc. Anterior region of test very inflated. Paired petals with unequal columns of respiratory podia. Naked zone along oral midline developed and pitted. Phyllodes without occluded plates. Peristome and periproct transverse. Three interambulacral plates on each side of periproct. Basicoronal plate 5 narrow and elongated. Etymology. Named after Briareus, the Greek God of violent sea storms, in reference to the turbulent taxonomic history related to the specimens described herein (see taxonomic history below). Description. Test oval (TW 79% of TL), lateral edges straight, anterior and posterior margins round; anterior and posterior regions of aboral region inflated with greatest height at apical disc, transverse cross section domed; oral region concave along the midline of the anterior-posterior axis. Test measurements (TL * TW * TH in mm): 18.36 * 14.52 * 10.4 (H) (Fig. 1A���B) and 18.2 * ca. 14.3 * ca. 10.4 (P) (Fig. 1D���E). Apical disc anterior, monobasal, ca. 9.5% of TL, flat, with four gonopores on disc edge; hydropores abundant and spread across plate (Fig. 2A). Anterior ocular plates between their adjacent gonopores; posterior ocular plates large and slightly posterior to gonopores 1 and 4; posterior region of apical disc slightly bulging towards interambulacrum. Petalodium about 75% of TL. Petals roughly with same L and W, broad in the middle and narrow distally, but not closing (Fig. 2B���D); inner and outer columns of respiratory podia bowed; poriferous zone narrow, pores slightly conjugated; outer pores elongated, inner pores round and smaller than outer. Columns a and b of posterior petals differ by 2 pore-pairs (number of respiratory podia in P: petal I, 21/23; petal V, 24/22), of anterior paired petals differ by 1���2 pore-pairs (number of respiratory podia in H: petal II, 21/20; petal IV, 18/20), anterior petal is equal (H: petal III, 21/21 respiratory podia). Primary tubercles present in poriferous zone; 3, sometimes 4 primary tubercles per petal plate. No occluded plates in petals. Ambulacra beyond petals increase 60���90% in relation to end-petal W; unipores in plates beyond petals: aboral plates wider than long, pores on suture, between adradial edge and middle of plate (Fig. 2E), and oral plates about 2x longer than wide, pores on middle of plate suture (Fig. 2I). Phyllodes with unipores, with only one column of pores on each half (number of phyllopores per half: anterior phyllode 5���6, paired phyllodes 7���9) (Fig. 2G���L). Plates sometimes unequal in size and shape; pyrinoid plate present on phyllode II, between plates 5 and 6 (Fig. 2G, J). Columns slightly bowed proximally and W narrows down distally. Pores usually aligned in a uniform column. Buccal pores same size as phyllopores. Ambulacral basicoronal plates pierced by buccal pore and one per ambulacrum also pierced by a phyllopore in the sequence a, a, b, a, b from phyllode I to V (Fig. 2G). Four to 5 sphaeridia in large and enclosed pits near buccal pores (Fig. 2H). Peristome anterior (34% TL from anterior region), deep (basicoronal plates extend further towards the inside of the test), slightly transverse (L 78���81% of W), pentagonal on the outside and subpentagonal on the inside (Fig. 1C). Mouth opening in center of peristomial membrane. Bourrelets slightly developed as mounds mostly towards the inside of the peristome (Fig. 10A���B). Cross-section of bourrelet with ca. 8 spines (Fig. 10B). Columns a and b of interambulacrum 5 with 7 and 8 plates between basicoronal plate and base of periproct, respectively, and 2 until adapical region of periproct (i.e., periproct is framed by 3 plates on each side). Interambulacral basicoronal plates 1 and 4 very narrow (much reduced and occluded in one specimen), 2 and 3 broadest, 5 intermediate in size (Fig. 2G). Second and third oral plates on interambulacrum 5 are much longer than wide. Naked zone well-developed throughout midline of test on interambulacrum 5 and ambulacrum III (W 22��� 25% of TW) (Fig. 1B). Deep pits present in oral region, mostly on naked zone. Periproct marginal and transverse (L 46% of W), beyond posterior petals; aboral plates form a prominent lip, oral and lateral plates do not bend inside the periproct (Fig. 1E). Periproct framed adorally by plates 5.a.7 and 5.b.8, and adapically by plates 5.a.9 and 5.b.10. Periproctal membrane with 2 rows of 4 large plates; smaller plates scattered in aboral region near anus (Fig. 2F). Anus opening aborally, on center of periproctal membrane. Primary tubercles perforate and slightly crenulate. Oral primary tubercles with mamelon displaced in the opposite direction of the spine (usually anteriorly from center of bosses) and ca. 2.5x as large as aboral tubercles. Bourrelet spines curved and with thick tip, oral spines long and straight, aboral spines short and straight, spines on periproct thin, straight, and intermediate in size between oral and aboral spines. Miliary tubercles all over the test. Few tridentate pedicellariae on stalks (valves 220���225 mm L) on periproct. Because very little soft tissue was preserved on available specimens, ophicephalous pedicellariae and tube foot ossicles were not observed. Remarks. Cassidulus briareus sp. nov. differs from its congeners (i.e., C. caribaearum, C. infidus and C. mitis) (Table 2) by having the anterior region of the test very inflated (vs. gradual height increase towards apical disc); a narrow and elongated basicoronal plate 5 (vs. squarish shape); a subpentagonal to triangular peristome, from the inside of the test (vs. subpentagonal to pentagonal); and a reduced number of plates framing the periproct (3 vs. 4 in the others [but rarely 3 plates on b column]). In addition, it differs from C. caribaearum by having an arched test in cross section of the adult (vs. triangular shape), shorter periproct, and more developed phyllodes with 2���3 more phyllopores than C. caribaearum; from C. mitis by having a uniform row of pores in the phyllodes (vs. disorganized row in which some pores are displaced forming an apparent inner series), and from C. infidus by having the greatest height at the apical disc (vs. posterior to the apical disc). Cassidulus briareus sp. nov. and its historical context within the genus Cassidulus. The specimens described herein were formerly identified as Cassidulus australis Blv. To our knowledge, Blainville did not describe any living species of Cassidulus and in his papers, he acknowledged Lamarck as author of C. australis (see Blainville, 1830, p. 192; 1834, p. 210). This species was described by Lamarck (1816) but later considered to be invalid (Gray, 1855, p. 34; A. Agassiz, 1872 ���1874, p. 153; Mortensen, 1948a, p. 209���210). Here we provide information on the taxonomic history of the genus Cassidulus that was pertinent in the decision to describe the new species. Lamarck (1801, p. 348���349) described C. caribaearum together with the genus Cassidulus, without recording the repository of the holotype or the type locality of the species. The latter was assumed to be the Caribbean, given the name of the species and also because Lamarck mentioned Spanish Town (near Kingston, Jamaica) in a later publication (Lamarck, 1816, p. 35). He (1801, p. 349) also made reference to Brugui��re et al. (1827, pl. 143, fig. 8��� 10), which includes illustrations of the specimen that, although of poor quality and lacking detail, captured the triangular shape of the test in cross section and the shape of the periproct and peristome. In 1816 (p. 35), Lamarck described C. australis, based on specimens supposedly collected by Charles Alexandre Lesueur and Fran��ois P��ron during the Baudin Expedition (1801���1803) in Shark���s Bay, Western Australia (as ���baie des Chiens marins, Nouvelle-Hollande ���). This description was slightly different from the description of C. caribaearum; however, Lamarck mentioned that C. australis also occurred in the Caribbean and included in his synonymy list Brugui��re et al. (1827, pl. 143, fig. 8���10), which means this species and C. caribaearum are synonyms. Again, he failed to record where the specimen was deposited. Shortly afterwards, Lamouroux et al. (1824, p. 174) synonymized C. caribaearum with C. australis and redescribed the species. They also suggested re-naming the species C. richardi, in honor of the person who had collected the type of C. caribaearum, and mentioned that it was not clear that the specimen collected by P��ron and Lesueur was the same as the one described by Lamarck (1816). Their description stated that the specimen was ���rather convex above���, which matches the shape of Cassidulus briareus sp. nov., but the test shape in the illustration is more similar to that of C. caribaearum (triangular shape seen in large specimens). All subsequent publications involving the genus Cassidulus (including the ones by Lamarck) accepted the synonymy proposed by Lamouroux et al. (1824), although any of the three names (C. caribaearum, C. australis and C. richardi) was used for the species (e.g., Eichwald, 1829; Blainville, 1830; 1834; Lamarck, 1837; 1840; Des Moulins, 1835 ���1837; A. Agassiz, 1869; Mortensen, 1948b; Mooi, 1990b). The occurrence of the species in Australia was often disregarded even though Lamarck kept including this locality in his papers. L. Agassiz & Desor (1847, p. 157) finally indicated that there was a specimen of C. australis (possibly the holotype) in the MNHN; and Mortensen (1948b, p. 209���210) suggested that the occurrence of the species in Australia was probably a result of ���erroneous labeling���. His reasoning was that if the species occurred in Australia, more specimens would have been collected during the intervening 150 years; logic that makes sense for many taxa. This species, however, appears to be less common than expected. Vadon et al. (1984) mentioned that they did not find the type of C. caribaearum in the MNHN and that Lamarck probably described the same specimen twice. The label retained the name used last, i.e., C. australis. However, Lamarck (1816) clearly mentioned that they were specimens from different localities. In addition, Lamarck���s descriptions of C. caribaearum and of C. australis were slightly different, although both were somewhat imprecise. Possible type specimen of C. caribaearum at the MNHN. Following up on this debate, we believe that the vial identified as C. caribaearum in the MNHN (Fig. 3, MNHN-IE-2013-10590 [Syntype?]; previously as EcEs 5040) might contain the type of one of the species described by Lamarck or perhaps even of both species. Two labels in the vial provide different identifications: one is identified as C. australis and the other (label apparently younger and written by A. Agassiz) is identified as Rhyncholampas caribaearum, with C. australis and Nucleolites richardi as synonyms. Also, both labels refer to the ���Voyage de P��ron & Lesueur 1801���, which indicates the material was from Australia rather than the Caribbean (MNHN, 2017). This vial contains broken pieces of two specimens as indicated by the fact that there are six complete phyllodes (one entire set and an additional broken phyllode). The following traits characterize them as Cassidulus: petal unequal in length (only one present, not sure if anterior or posterior; Fig. 3C, G), peristome transverse and subpentagonal (W 3.39 mm, L 2.18 mm), buccal pores present, sphaeridia in enclosed pits (Fig. 3D), phyllodes with a single column of phyllopores per half (Fig. 3D���F), naked zone developed and deeply pitted, mamelons of oral primary tubercles displaced from center. Additional characteristics are: posterior phyllodes and phyllode III with 5���6 pores, anterior paired phyllodes with 6���7 pores, plates beyond phyllodes longer than wide. Basicoronal plate 5 seems more elongated than in specimens of C. caribaearum preserved in the CASIZ collection (CASIZ 112633, 112683 A) but not as narrow as in Cassidulus briareus sp. nov. (Fig. 3A), and the number of phyllopores also overlaps both species. Therefore, identifying the broken pieces based on these traits is not reliable, nor would be any assertion that they belong to one or two species. Also, the illustrations in Brugui��re et al. (1827, pl. 143, fig. 8���10) are very poor and only capture the overall shape of the specimen, making it impossible to determine if they correspond to the broken specimens in the vial at the MNHN. Decision to describe a new species. According to the synonymy list provided by Lamarck (1816), C. australis is a subjective junior synonym of C. caribaearum. In addition, since C. australis was poorly described, it is impossible to determine if any subsequently collected Australian Cassidulus specimens are conspecific. The type specimen is probably lost, unidentifiable, or lacking diagnostic features, leading to much confusion in the literature regarding its validity. Therefore, it is not possible to determine that the material collected by Salmin is the same as Lamarck's C. australis, and we elected to describe a new species while at the same time stabilizing the nomenclature within the genus Cassidulus. Prediction of the type locality of C. briareus sp. nov. C.L. Salmin was a trader in Hamburg, Germany, who collected and sold specimens during the 1860s���1870s without taking much care in recording details regarding sampling locality (Holthuis, 2002). The only record we found in the literature documenting a collection of Australian echinoderms made by him is of holothuroids, collected in Cape York, Queensland (Samyn et al., 2013). However, museum registries also indicate that he has collected marine fish in Victoria, suggesting that the specimens described herein could have been collected in other places along the Australian coast. If ��� C. australis ��� was indeed collected by P��ron and Lesueur in Australia, it is possible that C. briareus sp. nov. lived in the same region (i.e., Shark���s Bay, Western Australia). Decision to designate a neotype for Cassidulus caribaearum Lamarck. As the type material of this species cannot be reliably identified and the illustrations provided by Brugui��re et al. (1827, pl. 143, fig. 8���10) do not contain diagnostic characteristics, a neotype designation for C. caribaearum is necessary to objectively define this species name (ICZN 1999, articles 75.1 and 75.3.4), and also to clarify the taxonomic status of the higher taxonomic ranks in which this species serves as the type species., Published as part of Souto, Camilla & Martins, Luciana, 2018, Synchrotron micro-CT scanning leads to the discovery of a new genus of morphologically conserved echinoid (Echinodermata: Cassiduloida) in Zootaxa 4457 (1), DOI: 10.11646/zootaxa.4457.1.3, http://zenodo.org/record/1457567, {"references":["Lamarck, J. B. P. A. de M. de (1801) Systeme des animaux sans vertebres, ou tableau general des classes, des ordres et des genres de ces animaux; presentant leurs caracteres essentiels et leur distribution, d'apres la consideration de leurs rapports naturels et de leur organisation, et suivant l'arrangement etabli dans les galeries du Museum d'hist. naturelle, parmi leurs depouilles conservees; precede du discours d'ouverture de cours de zoologie donne dans le Museum national d'histoire naturelle l'an 8 de la Republique. Chez l'auteur & Deterville, Paris, 432 pp.","Mortensen, T. (1948 b) New Echinoida (Cassiduloida; Clypeasteroida). Uidenskabelige Meddelelser fra Dansk Naturhistorisk Forening, 111, 67 - 72.","Krau, L. (1954) Nova especie de ourico do mar: Cassidulus mitis, ordem Cassiduloida, Echinoidea, capturado na Baia de Sepetiba. Memorias do Instituto Oswaldo Cruz, 52 (2), 455 - 475. https: // doi. org / 10.1590 / S 0074 - 02761954000200013","Blainville, H. M. de (1830) Zoophytes. In: Dictionnaire des sciences naturelles, dans lequel on traitre methodiquement des differens etres de la nature, consideres soit en eux-memes, d'apres l'etat actuel de nos connoissances, soit relativement a l'utlite qu'en peuvent retirer la medicine, l'agriculture, le commerce et les arts. Edited by F. G. Levrault. Uol. 60. Le Normant, Paris, 631 pp.","Lamarck, J. B. P. A. de M. de (1816) Histoire naturelle des animaux sans vertebres. Uol. 3. Verdiere, Paris, 586 pp.","Gray, J. E. (1855) Catalogue of the Recent Echinida, or sea eggs, in the collection of the British Museum. Part I. Echinida Irregularia. Order of the Trustees, London, 69 pp.","Agassiz, A. (1872 - 1874) Revision of the echini. Memoirs of the Museum of Comparative Zoology, 3, 383 - 762.","Mortensen, T. (1948 a) A Monograph of the Echinoidea. IU. 1. Holectypoida, Cassiduloida. C. A. Reitzel, Copenhagen, 363 pp.","Bruguiere, J. - G., Bory de Saint-Vincent, J. B. G. & Muller, O. F. (1827) Tableau encyclopedique et methodique des trois regnes de la nature: vers, coquilles, mollusques et polypiers. Uol. 2. Chez Mme veuve Agasse, Paris, 219 pls. [pls. 96 - 314] https: // doi. org / 10.5962 / bhl. title. 63299","Lamouroux, M. M., Saint-Vincent, B. & Deslongchamps, E. (1824) Encyclopedie methodique: histoire naturelle des zoophytes, ou animaux rayonnes. Uol. 2. Chez Mme veuve Agasse, Paris, 819 pp.","Eichwald, E. (1829) Zoologia specialis quam expositis animalibus tum vivis, tum fossilibus potissimum Rossiae in universum et Poloniae in specie. Uol. 1. J. Zawadzki, Vilnius, 314 pp.","Blainville, H. M. de (1834) Manuel d'Actinologie ou de Zoophytologie. F. G. Levrault, Strasbourg, 694 pp.","Lamarck, J. B. P. A. de M. de (1837) Histoire naturelle des animaux sans vertebres. Uol. 1. Meline, Cans et Compagnie, Bruxelles, 670 pp.","Lamarck, J. B. P. A. de M. de (1840) Histoire naturelle des animaux sans vertebres. Uol. 3. J. B. Bailliere, Paris, 770 pp.","Des Moulins, C. (1835 - 1837) Etudes sur les echinides. Actes de la Societe Linneenne de Bordeaux, 7 / 9, 1 - 518.","Agassiz, A. (1869) Preliminary report on the echini and star-fishes dredged in deep water between Cuba and the Florida Reef. Bulletin of the Museum of Comparative Zoology, 1 (9), 253 - 308.","Mooi, R. (1990 b) Living cassiduloids (Echinodermata: Echinoidea): a key and annotated list. Proceedings of the Biological Society of Washington, 130 (1), 63 - 85.","Agassiz, L. & Desor, E. (1847) Catalogue raisonne des especes, des genres et des familles d'echinides. Annales des Sciences Naturelles, Zoologie, 3 (7), 129 - 168.","Vadon, C., De Ridder, C., Guille, A. & Jangoux, M. (1984) Les types d'echinides actuels (echinodermes) du Museum National d'Histoire Naturelle de Paris. Bulletin du Museum National d'Histoire Naturelle, 6 (4), 1 - 38.","MNHN (2017) Museum national d'Histoire naturelle, Paris (France). Collection: Echinoderms (IE). Specimen MNHN-IE- 2013 - 10590. Accessed from: http: // coldb. mnhn. fr / catalognumber / mnhn / ie / 2013 - 10590 (accessed 17 July 2018)","Holthuis, L. B. (2002) The Indo-Pacific scyllarine lobsters (Crustacea, Decapoda, Scyllaridae). Zoosystema, 24, 499 - 683.","Samyn, Y., Smirnov, A. & Massin, C. (2013) Carl Gottfried Semper (1832 - 1893) and the location of his type specimens of sea cucumbers. Archives of Natural History, 40 (2), 324 - 339. https: // doi. org / 10.3366 / anh. 2013.0179","ICZN (1999) International Code of Zoological Nomenclature. Fourth Edition. The International Trust for Zoological Nomenclature, London, 306 pp."]}

Genus Euthyonidiella Heding & Panning 1954 Type species. Euthyonidiella kyushuensis Heding & Panning 1954; by original designation.

Cucumaria solangeae sp. nov. (Figures 7���8) Type material. Holotype: Pituba beach, Salvador, BA, Brazil (13 ��00��� S; 38 �� 27 ��� W): intertidal, 1 February 1994, 4 cm length (UFBA 650). Paratypes: same locality and date as Holotype, 8 spms 2���4 cm long (UFBA 1750; MZUSP 286���290; ZUEC HOL 21��� 22). Diagnosis. Body fusiform. Tube feet arranged in double rows restricted to the ambulacral region. Ten tentacles, ventral-most pair smaller. Calcareous ring compact and simple, lacking posterior processes; plates laterally united at the base. Deposits: body wall with knobbed buttons; introvert and tentacles with rods; tube feet with endplate and supporting rods; anal region with plates. Etymology. Named after Dr. Solange Peixinho (in memorian), in recognition of her dedication and contribution to the taxonomy of marine invertebrates, and for collecting the specimens described herein. Description. Body fusiform, tapering to the posterior end (Fig. 7 A), maximum length 4 cm. Tube feet arranged in double rows restricted to the ambulacral region, although a few scattered tube feet may also be present. Ten tentacles, ventral-most pair smaller (Fig. 7 B). Anal region with five thin papillae (Fig. 7 C) and calcareous teeth. Calcareous ring compact and simple, lacking posterior processes (Fig. 7 D���E). Radial plates broad at the base and tapering at the anterior region (we tried to remove the tissue to look if there really is a bifurcation at the apex but the radials were too fragile and disintegrated). Interradial plates curved at the base and tapering at the anterior region. Radial and interradial plates of same height and laterally united at the base. Coloration in ethanol: body light brown, tentacles dark purple, tube feet brown. Details of the internal morphology could not be observed because of the poor conservation of the specimens. Deposits: body wall with four-holed (rarely five-holed) buttons (70���120 ��m) (Fig. 8 A). Introvert and tentacles with arched rods with one perforation at each extremity and two in the middle (150���180 ��m) (Fig. 8 B). Tube feet with endplate (200���400 ��m) and supporting rods with small perforations at the extremities and an enlarged central region with a large perforation (Fig. 8 C) (140���160 ��m). Anal region with two-layered perforated plates (300���400 ��m) (Fig. 8 D), (300���400 ��m), elongated and knobbed perforated plates (Fig. 8 E), and perforated plates with irregular margins (400���600 ��m) (Fig. 8 F). Type locality. Pituba beach, Salvador ��� BA, Brazil. Biological notes. Specimens collected under rocks. Remarks. Cucumaria solangeae sp. nov. is the first record of the genus in Brazil, and differs from the five other valid Atlantic species [C. frondosa (Gunnerus 1767), C. georgiana Lampert 1886, C. arcuata (Herouard 1921), C. parassimilis Deichmann 1930, and C. acuta Massin 1992] by the presence of knobbed buttons in its body wall. Some previously described Cucumaria species with knobbed buttons have been transferred to other genera; however, we place C. solangeae sp. nov. in this genus because of the combination of characters (i.e. arrangement of tube feet, morphology of the calcareous ring, shape of plates, absence of cups on body wall). Other congeners with knobbed buttons are Cucumaria crax Deichmann 1941 and Cucumaria flamma Sol��s- Mar��n & Laguarda-Figueras 1999, from the Pacific coast of Mexico; however, their buttons are rudimentary (i.e. reduced knobbed edges) and these species have tentacles of the same size (vs. ventral-most two smaller in C. solangeae sp. nov.). The species Cucumaria compressa (Perrier 1898) from the northeastern Atlantic and Cucumaria vicaria Sluiter 1910 from the northwestern Atlantic have cups in the body wall and should not be classified within the subfamily Cucumariinae. According to the classification proposed by Smirnov (2012), these species should be Colochirinae., Published as part of Martins, Luciana & Souto, Camilla, 2015, Taxonomic review of four western Atlantic dendrochirotids (Holothuroidea) with the description of a new Brazilian cucumariid species and designation of neotypes, pp. 362-374 in Zootaxa 3919 (2) on pages 370-373, DOI: 10.11646/zootaxa.3919.2.8, http://zenodo.org/record/243332, {"references":["Gunnerus, J. E. (1767) Beskrifning pa trenne Norrska Sjo-Krak, Sjo-Pungar kallade. Kungliga Svenska vetenskapsakademiens Handlingar, 28, 114 - 124.","Lampert, K. (1886) Die Holothurien von Sud-Georgien, nach der Ausbeute der deutschen Polarstation in 1882 und 1883. Jahrbuch der wissenschaftlichen Anstalten zu Hamburg, 3, 11 - 21.","Herouard, E. (1921) Holothuries nouvelles des campagnes du yacht Princess-Alice. Bulletin de L'institut Oceanographique, 239, 1 - 9.","Deichmann, E. (1930) The holothurians of the western part of the Atlantic Ocean. Bulletin of the Museum of Comparative Zoology at Harvard College, 71 (3), 1 - 226.","Massin, C. (1992) Three new species of Dendrochirotida (Holothuroidea, Echinodermata) from the Weddell Sea (Antarctica). Bulletin de l'Institut Royal des Sciences Naturelles de Belgique: Biologie, 62, 179 - 191.","Deichmann, E. (1941) The Holothuroidea collected by the velero III during the years 1932 to 1938 - Part I. Dendrochirota. The University of Southern California Publications, 8 (3), 1 - 194.","Solis-Marin, F. A. & Laguarda-Figueras, A. (1999) Cucumaria flamma, a new species of sea cucumber from the central eastern Pacific (Echinodermata: Holothuroidea). Proceedings of the Biological Society of Washington, 112 (4), 778 - 786.","Perrier, R. (1898) Sur les Holothuries recueillies par le Travailleur et le Talisman. Comptes Rendus Academie Sciences, 126, 1664 - 1666.","Sluiter, C. P. (1910) Westindische Holothurien. Zoologische Jahrbucher, 11 (Supplement), 331 - 342.","Smirnov, A. V. (2012) System of the Class Holothuroidea. Paleontological Journal, 46 (8), 793 - 832. http: // dx. doi. org / 10.1134 / S 0031030112080126"]}

Genus Thandarum Martinez & Brogger 2012 Type species. Thandarum hernandezi; by original designation., Published as part of Martins, Luciana & Souto, Camilla, 2015, Taxonomic review of four western Atlantic dendrochirotids (Holothuroidea) with the description of a new Brazilian cucumariid species and designation of neotypes, pp. 362-374 in Zootaxa 3919 (2) on page 368, DOI: 10.11646/zootaxa.3919.2.8, http://zenodo.org/record/243332, {"references":["Martinez, M. I. & Brogger, M. I. (2012) Thandarum hernandezi, a new genus and new species of sea cucumber family Sclerodactylidae (Echinodermata: Holothuroidea: Dendrochirotida) from the Southwestern Atlantic Ocean. Zootaxa, 3304, 63 - 68."]}

Thandarum manoelina (Tommasi 1971) comb. nov. (Figures 5���6) Cucumaria manoelina Tommasi 1971: 2, figs. 3���7. Cucumaria pulcherrima ��� Tommasi 1969: 141, pl. 11, fig. 13. Type material. Neotype: designated herein, Guarajuba, Cama��ari, BA, Brazil (12 �� 47 ��� S; 38 ��08��� W): 23 m, 1 July 2007, 1 spm 0.5 cm long (UFBA 1746). [Holotype originally at the Tommasi collection at the Universidade de S��o Paulo; type locality: Ilha Grande ��� RJ, Brazil, now lost]. Other material examined. Guarajuba, Cama��ari, BA, Brazil (12 �� 47 ��� S; 38 ��08��� W): 23 m, 1 July 2007, 1 spm 0.5 cm long (UFBA 01844); 23 m, 15 August 2010, 15 spms 0.3���0.7 cm long (UFBA 1747); 23 m, 15 July 2011, 2 spms 0.5 cm long (UFBA 1748). Ba��a de Todos-os-Santos, Salvador, BA, Brazil (12 �� 50 ��� S; 38 �� 45 ��� W): 12 m, 9 March 2010, 1 spm 3 cm long (UFBA 1749). Neotype diagnosis: Body U-shaped; tube feet arranged in two rows restricted to the ambulacral region. Ten tentacles, ventral-most pair smaller. Calcareous ring compact; radial and interradial plates joined only at their base; radials with posterior processes subdivided. Deposits: body wall with four-pillared tables and irregular buttons; introvert with rosettes; tentacles with rods; and tube feet with rods and star-shaped endplate. Neotype description. Body U-shaped (Fig. 5 A). Tube feet arranged in double rows restricted to the ambulacra. Ten tentacles, ventral-most pair smaller. Anal region with five groups of three papillae. Calcareous ring compact and delicate, radial and interradials united only at their base (Fig. 5 B). Radials thin anteriorly and thick at the base, with thick and subdivided posterior processes (made up of 3���4 pieces) (Fig. 5 C). Interradials triangularshaped and strongly concave. Radial/interradial length ratio approximately 1: 3. The internal morphology of the specimens could not be observed but, according to Tommasi (1971), this species bears one Polian vesicle. Body color yellowish in vivo and white in ethanol. Deposits: body wall with four-pillared tables (30���60 ��m) and buttons (70���120 ��m). Table discs with irregular and smooth margins, and a strong central concavity bordered by numerous irregular perforations; spire high and ending in many teeth (�� 20) (Fig. 6 A���C). Buttons with irregular shape and many perforations (Fig. 6 D���E). Introvert with rosettes (20���50 ��m). Tentacles with typical rods (70���90 ��m) (Fig. 6 F). Tube feet with typical rods (80���100 ��m) (Fig. 6 G) and a star-shaped endplate entirely perforated (100���120 ��m) (Fig. 6 H). Distribution. Brazil (Bahia and Rio de Janeiro); depth: 12���50 m (Tommasi 1971; present paper). Biological notes. This species has been found buried in silt, sand (Tommasi 1971), medium sand and gravel (present paper). Remarks. The genus Cucumaria belongs to the family Cucumariidae and, therefore, has a simple calcareous ring, lacking posterior processes. The holotype of Cucumaria manoelina was originally described by Tommasi (1971) as possessing a calcareous ring with posterior processes, a typical ring of the Sclerodactylidae or Phyllophoridae. The morphology of the calcareous ring indicates that C. manoelina should be classified in the subfamily Sclerothyoninae (family Sclerodactylidae) since the radials are compact (i.e. without subdivisions), radials and interradials are united only at their base, and the radials are 3���4 times larger than the interradials. Three Sclerothyoninae genera have been described; among them, the diagnosis of the genus Thandarum includes the species C. manoelina. In this regard, we propose the new combination Thandarum manoelina (Tommasi 1971). Thandarum was recently described to the coast of Argentina (Buenos Aires, Martinez & Brogger 2012) and T. manoelina extends the distribution of this genus to the tropical Brazilian waters (up to Bahia State). Thandarum manoelina differs from Thandarum hernandezi Martinez & Brogger 2012 by the presence of smaller tables (40���60 ��m vs. 260 ��m���same-sized spms) with irregular discs (vs. circular margins) and numerous perforations (vs. few perforations). Thandarum manoelina differs from the other Sclerothyoninae (Sclerothyone velligera [Ludwig & Heding 1935], Sclerothyone unicolumnus Thandar 2008 and Temparena chuni [Ludwig & Heding 1935]���all from South Africa) by the presence of four-pillared body wall tables and rosettes in the introvert. Thandarum manoelina is the first occurrence of the subfamily Sclerothyoninae in the Brazilian coast., Published as part of Martins, Luciana & Souto, Camilla, 2015, Taxonomic review of four western Atlantic dendrochirotids (Holothuroidea) with the description of a new Brazilian cucumariid species and designation of neotypes, pp. 362-374 in Zootaxa 3919 (2) on pages 368-370, DOI: 10.11646/zootaxa.3919.2.8, http://zenodo.org/record/243332, {"references":["Tommasi, L. R. (1971) Equinodermes do Brasil. I. Sobre algumas especies novas e outras pouco conhecidas para o Brasil. Boletim do Instituto Oceanografico, Universidade de Sao Paulo, 302, 1 - 21.","Tommasi, L. R. (1969) Lista dos Holothuroidea recentes do Brasil. Contribuicoes Avulsas do Instituto Oceanografico, 15, 1 - 29.","Martinez, M. I. & Brogger, M. I. (2012) Thandarum hernandezi, a new genus and new species of sea cucumber family Sclerodactylidae (Echinodermata: Holothuroidea: Dendrochirotida) from the Southwestern Atlantic Ocean. Zootaxa, 3304, 63 - 68.","Ludwig, H. L. & Heding, S. G. (1935) Die Holothurien der deutschen Tiefsee-Expedition. I. Fußlose und dendrochirote Formen. Wissenschaftliche Ergebnisse der Deutschen Tiefsee - Expedition auf dem Dampfer Valdivia 1898 - 1899, 24 (2), 123 - 241.","Thandar, A. S. (2008) Additions to the holothuroid fauna of the southern African temperate faunistic provinces, with descriptions of new species. Zootaxa, 1697, 1 - 57."]}

Euthyonidiella occidentalis (Ludwig 1875) comb. nov. (Figures 1���4) Thyonidium occidentale Ludwig 1875: 119 ���120. Euthyonidiella dentata Cherbonnier 1961: 611 ���613, fig. 1. Thyone constituta ��� Sluiter 1910: 340, fig. F. Phyllophorus occidentalis ��� Deichmann 1930: 148, pl. 18. Phyllophorus (Urodemella) occidentalis ��� Heding & Panning 1954: 164, fig. 76. Type material. Neotype: designated herein, Itapu�� beach, Salvador, BA, Brazil (12 �� 57 ��� S; 38 �� 21 ��� W): intertidal, 12 January 2014, 1 spm 8 cm long (MZUSP 1139). [Holotype originally at the ZMB; type locality: Suriname, now lost (Dr. Carsten Lueter, pers. comm.)]. Other material examined. Panama, Caribbean Sea: 1973, 5 spms 2���5 cm long (USNM E 22300). Itaparica Island, BA, Brazil (12 �� 54 ��� S; 38 �� 37 ��� W): 7 February 1993, 1 spm 3 cm long (EQMN 1719). Itapu�� beach, Salvador, BA, Brazil (12 �� 57 ��� S; 38 �� 21 ��� W): intertidal, 12 January 2014, 4 spms 3���5 cm long (MZUSP 1067). Amaralina beach, Salvador, BA, Brazil (13 ��00��� S; 38 �� 28 ��� W): intertidal, 8 April 2008, 3 spms 2���6 cm long (UFBA 635); intertidal, 19 April 2011, 14 spms 2���10 cm long (UFBA 1335); intertidal, 21 April 2011, 19 spms 1���5 cm long (UFBA 1341); intertidal, 1 May 2011, 1 spm 5 cm long (UFBA 1379). Porto Seguro, BA, Brazil (16 �� 25 ��� W; 39 ��02��� S): 27 August 1980, 11 spms 2���5 cm long (EQMN 607). Cumuruxatiba, BA, Brazil (17 �� 56 ��� S; 39 �� 12 ��� W): 14 October 1982, 2 spms 2.5 cm long (UFPB 1876). Abrolhos, BA, Brazil (18 ��03��� S; 38 �� 41 ��� W): 29 October 1997, 7 spms 1���5 cm long (EQMN 1878); 31 May 1998, 1 spm 3.5 cm long (EQMN 1879). Piloto beach, Vit��ria, ES, Brazil (19 �� 49 ��� S; 40 ��02��� W): 28 January 1979, 1 spm 3 cm long (EQMN 726); 20 July 1981, 3 spms 2���5 cm long (EQMN 726). Arma����o beach, B��zios, RJ, Brazil (22 �� 45 ��� S; 41 �� 51 ��� W): May 1983, 1 spm 4 cm long (EQMN 891). Ferradura beach, B��zios, RJ, Brazil (22 �� 46 ��� S; 41 �� 52 ��� W): 12 May 1999, 1 spm 4 cm long (EQMN 1923). Comparative material examined. Euthyonidiella dentata ��� holotype slides deposited in the MNHN (edentata dorsal hol 001; edentata dorsal hol 002; edentata dorsal hol 003; edentatatenthol 001; edentatatenthol 002; edentatatenthol 003; edentata ventral hol 001; edentata ventral hol 002; edentata ventral hol 003). Neotype diagnosis. Body U-shaped; twenty tentacles distributed in two circles (10 + 10), the inner circle has smallest tentacles. Calcareous ring compact; radials with short and undivided posterior processes. Deposits in body wall are tables with two pillars low spire, and dentate margins; introvert with rosettes and tables; tentacles with rods and rosettes; tube feet with rods and endplate. Neotype description. Body U-shaped (Fig. 1 A, C, D). Tube feet scattered over entire body. 20 tentacles arranged in 2 circles (10 + 10): 5 large pairs in outer circle alternating with 5 small pairs in inner circle (Fig. 1 E, F). Anal region bears five groups with three papillae each. Calcareous ring compact; radials with a deep anterior notch, and short and undivided posterior processes; interradials slightly shorter than the radials, with a broad base that overlap with the radials, and very short posterior bifurcation (Figs. 1 B, 2). Two Polian vesicles 4 mm long (Fig. 3 A, C), madreporite filamentous, stone canal 4.8 mm long (Fig. 3 A, B). Deposits: body wall tables (40���60 ��m) with dentate margins, 4���8 perforations (Fig. 4 A), and a reduced spire with two short pillars ending in four spines; introvert with rosettes (30���60 ��m) (Fig. 4 B) and tables with several perforations (Fig. 4 C, D); tentacles with rudimentary rods (35���75 ��m) (Fig. E) and rosettes (Fig. 4 B); tube feet with rods (Fig. 4 E) and endplate (150���300 ��m) (Fig. 4 F). Body color light brown (in vivo and preserved specimen). Distribution. U.S.A. (Florida), Caribbean Sea (Antigua, Aruba, Barbados, Puerto Rico, Trinidad and Tobago), Suriname and Brazil (up to Rio de Janeiro); depth: intertidal region up to 2 m (Hendler et al. 1995; present paper). Biological notes. Specimens of E. occidentalis comb. nov. live burrowed in soft sediment, under rocks and associated with seagrass (Hendler et al. 1995; present paper). Genus change. We noticed that Euthyonidiella occidentalis comb. nov. was distinct from the members of the family Phyllophoridae by possessing a calcareous ring compact, which in turn characterizes the family Sclerodactylidae. Within this family, this species should be placed within the subfamily Cladolabinae, which is characterized by the presence of 15���20 tentacles (vs. 10 tentacles in Sclerodactylinae and Sclerothyoninae); and within this subfamily, this species fits perfectly the diagnosis of the genus Euthyonidiella, by possessing 20 tentacles (vs. 15 in Clarkiella), short spired tables in the body wall (vs. no tables in Afrocucumis; and tall spired tables in Cladolabes), and regular rods and plates (vs. thorny rods and spinuos plates in Ohshimella). Pawson & Miller (1992, p. 483) discussed inconsistencies regarding the tentacles��� arrangement and the morphology of the calcareous ring within the subgenus Urodemella; nevertheless, they preferred to describe the species Phyllophorus (Urodemella) arenicola Pawson & Miller 1992 in this subgenus because of its similarity to E. occidentalis comb. nov. Therefore, we believe that P. (U.) arenicola should also be transferred to the genus Euthyonidiella, and be accepted as Euthyonidiella arenicola comb. nov. A revision of the families Phyllophoridae and Sclerodactylidae is needed since the classification of these dendrochirotids was built on divergent opinions and an agreement has not yet been reached. Regarding the calcareous ring, there are clearly two extreme morphologies that define the family Phyllophoridae and the subfamily Sclerodactylinae: plates subdivided in a mosaic of pieces vs. plates compact (see Thandar 1989, fig. 1 A, B). However, the calcareous ring of Phyllophorus (i.e. plates subdivided in large pieces) and of Sclerothyoninae (radial processes long and subdivided in small pieces), for instance, fall in between those two extremes. Moreover, Heding & Panning (1954) described the subfamily Cladolabinae and divided it into two groups based on the presence of tables in the body wall and on the presence of subdivisions in the calcareous ring. Given the great amount of tables in the body wall of these taxa it would be probably hard to miss it; however, the radial processes of their calcareous ring are generally short and fragile, breaking easily when manipulated. For instance, some processes broke down into smaller pieces recalling subdivided posterior processes (Fig. 2 A) when we removed the tentacles from the calcareous ring or after bleaching it. In addition, the subfamily Cladolabinae and the genus Phyllophorus possess an uncommon number and arrangement of tentacles (i.e. 10 + 10, 15 + 5, 15 + 5 + 5), distinguishing these taxa from the other dendrochirotids, which are mainly 10 +0 and 8 + 2. In this regard, phylogenetic studies based on morphological data could help to redefine the dendrochirotid clades by assessing phylogenetically informative traits. In addition, molecular data can provide a great set of characters in the absence of unambiguous morphological data, especially within the genera. However, regardless of the classification used, we believe the genus Euthyonidiella best fit both species (i.e. E. occidentalis comb. nov., and E. arenicola comb. nov.). Euthyonidiella occidentalis and Euthyonidiella dentata. After analyzing slides of the holotype of E. dentata and its original description (in Cherbonnier 1961), we concluded that there is no difference between this species and E. occidentalis c omb. nov. Euthyonidiella dentata is known only from its holotype, collected in Bahia (Brazil), and was never mentioned again, probably because the specimens collected were identified as ��� P. (U.) occidentalis ���. Therefore, we suggest that E. dentata should be designated as a junior synonym of E. occidentalis comb. nov. Comparison with other species. Considering the modifications proposed herein, ten species of Euthyonidiella are recorded: E. occidentalis comb. nov., E. aculeata (Ludwig 1894), E. trita (Sluiter 1910), E. destichata (Deichmann 1930), and E. arenicola comb. nov. [West Atlantic]; E. dubia Cherbonnier 1958 [East Atlantic]; E. tungshanensis (Yang 1937) and E. kyushuensis Heding & Panning 1954 [West Pacific]; E. zacae (Deichmann 1938) [East Pacific], and E. ambigua (Heding 1942) [Mediterranean]. Euthyonidiella occidentalis comb. nov. differs from E. trita, E. ambigua, E. kyushuensis and E. dubia by the arrangement of the tentacles in two circles with ten tentacles each (vs. 15 + 5, 10 + 5 + 5, 14, 10 + 5 + 5, respectively), from E. aculeata, E. zacae, E. destichata, E. kyushuensis and E. dubia by the presence of tables with dentate margins (vs. oval or circular shape), from E. tungshanensis by the presence of tables with low spire (vs. high spire), and from E. arenicola comb. nov. by the morphology of the tables, which have pointy projections in their discs (vs. blunt projections) and pillars fusing together at the base (vs. pillars linked by a horizontal ridge)., Published as part of Martins, Luciana & Souto, Camilla, 2015, Taxonomic review of four western Atlantic dendrochirotids (Holothuroidea) with the description of a new Brazilian cucumariid species and designation of neotypes, pp. 362-374 in Zootaxa 3919 (2) on pages 363-368, DOI: 10.11646/zootaxa.3919.2.8, http://zenodo.org/record/243332, {"references":["Ludwig, H. L. (1875) Beitragezur Kenntniss der Holothurien. Arbeitenausdem Zoologischen Zootomischen Institut in Wurzburg, 2 (2), 77 - 120.","Cherbonnier, G. (1961) Deux Nouvelles especes d'holothuries Dendrochirotes des cotes Bresilienes. Bulletin do Museum National d'histoire Naturelle, 33 (6), 611 - 615.","Sluiter, C. P. (1910) Westindische Holothurien. Zoologische Jahrbucher, 11 (Supplement), 331 - 342.","Deichmann, E. (1930) The holothurians of the western part of the Atlantic Ocean. Bulletin of the Museum of Comparative Zoology at Harvard College, 71 (3), 1 - 226.","Heding, S. G. & Panning, A. (1954) Phyllophoridae: eine Bearbeitung der Polytentaculaten Dendrochiroten Holothurien des Zoologischen Museums in Kopenhagen. Spolia Zoologische Musei Hauniensis, 13, 7 - 209.","Hendler, G., Miller, J. E., Pawson, D. L. & Kier, P. M. (1995) Sea stars, sea urchins and allies: echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp.","Pawson, D. L. & Miller, J. E. (1992) Phyllophorus (Urodemella) arenicola, a new sublittoral sea cucumber from the southeastern United States (Echinodermata: Holothuroidea). Proceedings of the Biological Society of Washington, 105 (3), 483 - 489.","Thandar, A. S. (1989) The Sclerodactylid holothurians of southern Africa, with the erection of one new subfamily and two new genera (Echinodermata: Holothuroidea). South African Journal of Zoology, 24 (4), 290 - 304.","Ludwig, H. (1894) Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U. S. Fish Commission Steamer Albatross, during 1891, Lieut. Commander Z. L. Tanner, U. S. N., commanding, xii. The Holothurioidea. Memoirs of the Museum of Comparative Zoology, 17 (3), 1 - 183. [Harvard]","Cherbonnier, G. (1958) Holothuries des cotes de Sierra-Leone. 4 e note. Bulletin Museum National Histoire Naturelle Paris, Serie 2, 30 (3), 294 - 299.","Yang, P. F. (1937) Report on the holothurians from the Fukien Coast. Bulletin Marine Biology, Amoy, China, 2, 1 - 46.","Deichmann, E. (1938) Eastern Pacific Expeditions of the New York Zoological Society. XVI. Holothurians from the western coasts of Lower California and Central America, and from the Galapagos Islands. Zoologica, 23, 361 - 387. [New York]","Heding, S. G. (1942) Uber Cucumella triplex und zwei neue Holothurien der Deutschen Tiefsee-Expedition. Zoologischer Anzeiger, 137, 217 - 220."]}

Genus Thandarum Martinez & Brogger 2012 Type species. Thandarum hernandezi; by original designation.

Genus Cucumaria Blainville 1830 Type species. Cucumaria frondosa (Gunnerus 1767); by subsequent designation (Panning 1949: 416)., Published as part of Martins, Luciana & Souto, Camilla, 2015, Taxonomic review of four western Atlantic dendrochirotids (Holothuroidea) with the description of a new Brazilian cucumariid species and designation of neotypes, pp. 362-374 in Zootaxa 3919 (2) on page 370, DOI: 10.11646/zootaxa.3919.2.8, http://zenodo.org/record/243332, {"references":["de Blainville, H. M. D. (1830) Zoophytes. In: Levrault, F. G. (Ed.), Dictionnaire des sciences naturelles, dans lequel on traitre methodiquement des differens etres de la nature, consideres soit en eux-memes, d'apres l'etat actuel de nos connoissances, soit relativement a l'utlite qu'en peuvent retirer la edicine, l'agriculture, le commerce et les arts. Tome 60. Le Normat, Paris, pp. 548.","Gunnerus, J. E. (1767) Beskrifning pa trenne Norrska Sjo-Krak, Sjo-Pungar kallade. Kungliga Svenska vetenskapsakademiens Handlingar, 28, 114 - 124.","Panning, A. (1949) Versucheiner Neuordnung der familie Cucumariidae (Holothuroidea: Dendrochirotida). Zoologische Jahrbucher Abtheilungfur Systematik, 78 (4), 404 - 470."]}

Genus Euthyonidiella Heding & Panning 1954 Type species. Euthyonidiella kyushuensis Heding & Panning 1954; by original designation., Published as part of Martins, Luciana & Souto, Camilla, 2015, Taxonomic review of four western Atlantic dendrochirotids (Holothuroidea) with the description of a new Brazilian cucumariid species and designation of neotypes, pp. 362-374 in Zootaxa 3919 (2) on page 363, DOI: 10.11646/zootaxa.3919.2.8, http://zenodo.org/record/243332, {"references":["Heding, S. G. & Panning, A. (1954) Phyllophoridae: eine Bearbeitung der Polytentaculaten Dendrochiroten Holothurien des Zoologischen Museums in Kopenhagen. Spolia Zoologische Musei Hauniensis, 13, 7 - 209."]}

Martins, Luciana, Souto, Camilla (2015): Taxonomic review of four western Atlantic dendrochirotids (Holothuroidea) with the description of a new Brazilian cucumariid species and designation of neotypes. Zootaxa 3919 (2): 362-374, DOI: http://dx.doi.org/10.11646/zootaxa.3919.2.8

Coronatum baiensis sp. nov. Martins & Souto (Figs. 1–4) Type Locality. Amaralina beach, Salvador, BA, Brazil. Holotype. Amaralina beach, Salvador, BA, Brazil (13 °00’ 96 ” S; 38 ° 28 ’ 16 ” W), 0 m, 2 May 2011, coll. L. Martins, C. Souto & U. Lopes, 2.6 cm long (UFBA – 1677). Paratypes. Same locality, date and collector as Holotype, 58 specimens, maximum total length up to 4.3 cm (48, UFBA – 1678; 5, MZUSP 00241; 5 ZUEC HOL 01–05). Additional material. Itapuã beach, Salvador, BA, Brazil (12 ° 57 ’ 46 ” S; 38 ° 21 ’ 36 ” W), 0 m, 21 April 2011, coll. L. Martins, C. Souto & U. Lopes, 35 specimens (UFBA – 1679). Diagnosis. Tentacles 10, ventral-most two much reduced. Tube feet scattered on body wall. Body wall ossicles are two-pillared tables with knobbed margins and low spire ending in numerous teeth, tube feet with two-pillared supporting tables, and introvert contains tables and rosettes. Etymology. The species epithet is derived from the name of the State where it was collected (Bahia State). Description of holotype. Medium-sized body, rough to the touch, barrel-shaped and sometimes slightly upturned at both ends. Tube feet gray, non-retractile and scattered throughout the body (Fig. 1 A). Ten dendritic tentacles, ventral-most pair smaller (Fig. 1 B–C). Five groups of three anal papillae around the anus (Fig. 1 E); under these, five anal teeth, each forming a triangle pointing outwards (Fig. 1 F). Calcareous ring (Fig. 1 D) poorly calcified, plates laterally united and radials slightly higher than interradials. Radials compact, bifid anteriorly and with subdivided posterior paired processes. Interradials spear-shaped and pointed anteriorly. Internal morphology: Gonads found in specimens over 1.5 cm in length and arranged in a single tuft attached to the middle portion of the body (Fig. 2 A–B). One Polian vesicle (Fig. 2 C–D), stone canal filamentous and short (Fig. 2 C, E), madreporite bean-shaped (Fig. 2 E). Deposits: Body wall tables with oval disc (100–130 µm diameter), thick and knobbed margins, 2 central and 12–30 peripheral perforations, and low spire of two pillars ending in over 20 short round teeth (Fig. 3 A–B). Tentacles with two types of rods: a curved rod with two callosities in the center and ± 25 oval perforations in both extremities which tend to decrease in size towards the end (140–240 µm) (Fig. 3 C); and an elongated rod rosettelike (190–240 µm) (Fig. 3 D). Introvert with rosettes (30–50 µm) (Fig. 3 E) and two-pillared tables of two types: one with circular disc with many perforations and a medium-sized spire ending in a few denticles (Fig. 3 F–G); and a second type with knobbed edges and a low spire ending in a few denticles (Fig. 3 H). Anal region with tables similar to the body wall tables. Tube feet with supporting tables (150–180 µm) composed of two short spires and ending in numerous small teeth (± 16) (Fig. 3 I), and end plates (Fig. 3 J). Color in life: most specimens are white, brown or gray, with dark spots scattered around the body; tentacles dark brown, introvert white and tube feet white with brown spots. Color not affected when preserved in ethanol. Distribution. Salvador, BA, Brazil. Bathymetric range: intertidal region. Biological Notes. Occur under rocks, inside crevices or buried with tentacles extended towards the surface. It can also be found in association with cucumariids, such as Thyonidium seguroensis (Deichmann, 1930). When touched in the field, Coronatum baiensis sp. nov. readily eviscerates internal structures through the introvert; therefore, they are often found without the calcareous ring, although the tentacles are intact.

Coronatum baiensis sp. nov. Martins & Souto (Figs. 1���4) Type Locality. Amaralina beach, Salvador, BA, Brazil. Holotype. Amaralina beach, Salvador, BA, Brazil (13 ��00��� 96 ��� S; 38 �� 28 ��� 16 ��� W), 0 m, 2 May 2011, coll. L. Martins, C. Souto & U. Lopes, 2.6 cm long (UFBA ��� 1677). Paratypes. Same locality, date and collector as Holotype, 58 specimens, maximum total length up to 4.3 cm (48, UFBA ��� 1678; 5, MZUSP 00241; 5 ZUEC HOL 01���05). Additional material. Itapu�� beach, Salvador, BA, Brazil (12 �� 57 ��� 46 ��� S; 38 �� 21 ��� 36 ��� W), 0 m, 21 April 2011, coll. L. Martins, C. Souto & U. Lopes, 35 specimens (UFBA ��� 1679). Diagnosis. Tentacles 10, ventral-most two much reduced. Tube feet scattered on body wall. Body wall ossicles are two-pillared tables with knobbed margins and low spire ending in numerous teeth, tube feet with two-pillared supporting tables, and introvert contains tables and rosettes. Etymology. The species epithet is derived from the name of the State where it was collected (Bahia State). Description of holotype. Medium-sized body, rough to the touch, barrel-shaped and sometimes slightly upturned at both ends. Tube feet gray, non-retractile and scattered throughout the body (Fig. 1 A). Ten dendritic tentacles, ventral-most pair smaller (Fig. 1 B���C). Five groups of three anal papillae around the anus (Fig. 1 E); under these, five anal teeth, each forming a triangle pointing outwards (Fig. 1 F). Calcareous ring (Fig. 1 D) poorly calcified, plates laterally united and radials slightly higher than interradials. Radials compact, bifid anteriorly and with subdivided posterior paired processes. Interradials spear-shaped and pointed anteriorly. Internal morphology: Gonads found in specimens over 1.5 cm in length and arranged in a single tuft attached to the middle portion of the body (Fig. 2 A���B). One Polian vesicle (Fig. 2 C���D), stone canal filamentous and short (Fig. 2 C, E), madreporite bean-shaped (Fig. 2 E). Deposits: Body wall tables with oval disc (100���130 ��m diameter), thick and knobbed margins, 2 central and 12���30 peripheral perforations, and low spire of two pillars ending in over 20 short round teeth (Fig. 3 A���B). Tentacles with two types of rods: a curved rod with two callosities in the center and �� 25 oval perforations in both extremities which tend to decrease in size towards the end (140���240 ��m) (Fig. 3 C); and an elongated rod rosettelike (190���240 ��m) (Fig. 3 D). Introvert with rosettes (30���50 ��m) (Fig. 3 E) and two-pillared tables of two types: one with circular disc with many perforations and a medium-sized spire ending in a few denticles (Fig. 3 F���G); and a second type with knobbed edges and a low spire ending in a few denticles (Fig. 3 H). Anal region with tables similar to the body wall tables. Tube feet with supporting tables (150���180 ��m) composed of two short spires and ending in numerous small teeth (�� 16) (Fig. 3 I), and end plates (Fig. 3 J). Color in life: most specimens are white, brown or gray, with dark spots scattered around the body; tentacles dark brown, introvert white and tube feet white with brown spots. Color not affected when preserved in ethanol. Distribution. Salvador, BA, Brazil. Bathymetric range: intertidal region. Biological Notes. Occur under rocks, inside crevices or buried with tentacles extended towards the surface. It can also be found in association with cucumariids, such as Thyonidium seguroensis (Deichmann, 1930). When touched in the field, Coronatum baiensis sp. nov. readily eviscerates internal structures through the introvert; therefore, they are often found without the calcareous ring, although the tentacles are intact., Published as part of Martins, Luciana, Souto, Camilla & Menegola, Carla, 2012, A new genus and new species of Sclerodactylidae (Holothuroidea: Dendrochirotida) from the south-western Atlantic coast, pp. 54-62 in Zootaxa 3506 on pages 59-60, DOI: 10.5281/zenodo.214409, {"references":["Deichmann, E. (1930) The holothurians of the western part of the Atlantic Ocean. Bulletin of the Museum of Comparative Zoology at Harward College, 71 (3), 1 - 226."]}

Coronatum gen. nov. Martins & Souto. Diagnosis. Tentacles 10, ventral-most two much reduced. Tube feet scattered on body wall, never restricted to the ambulacra. Calcareous ring compact with short posterior processes. Body wall ossicles are two-pillared tables, tube feet with two-pillared supporting tables, and introvert contains tables and rosettes. Etymology. The name of the genus is derived from the Latin words corona (crown) in reference to the numerous small teeth as shown on top of the body wall tables. The gender is neuter. Type species. Coronatum baiensis sp. nov. Martins & Souto. Remarks. The new genus was described to accommodate only the type species described herein. The twelve Sclerodactylinae genera can be organized into four distinct groups according to the set of ossicles within the body wall: 1) cups (Apentamera, Deichmannae, Eugeliella), 2) knobbed buttons (Eupentacta, Neothyone, Neopentamera, Pachythyone, Pseudothyone), 3) plates (Athyone), and 4) tables (Havelockia, Sclerodactyla, Coronatum gen. nov.). Therefore, Coronatum gen. nov. differs from the other groups by having tables in the body wall. Coronatum gen. nov. differs from Havelockia by having rosettes in the introvert, and from Sclerodactyla by the presence of four-pillared tables in the latter genus., Published as part of Martins, Luciana, Souto, Camilla & Menegola, Carla, 2012, A new genus and new species of Sclerodactylidae (Holothuroidea: Dendrochirotida) from the south-western Atlantic coast, pp. 54-62 in Zootaxa 3506 on page 55, DOI: 10.5281/zenodo.214409

Martins, Luciana, Souto, Camilla, Menegola, Carla (2012): A new genus and new species of Sclerodactylidae (Holothuroidea: Dendrochirotida) from the south-western Atlantic coast. Zootaxa 3506: 54-62, DOI: 10.5281/zenodo.214409

Souto, Camilla, Manso, Cynthia L. C., Martins, Luciana (2011): Rediscovery and redescription of Cassidulus infidus (Echinoidea: Cassidulidae) from Northeastern Brazil. Zootaxa 3095: 39-48, DOI: 10.5281/zenodo.279091