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The class Echinoidea contains among the best-known echinoderms. The group has left the most complete fossil record of this phylum, and contains about 1000 living species, of which 51 are recorded along the Brazilian coast. Although the first record of echinoids in Brazil was made 369 years ago, the knowledge of this fauna remains quite unsatisfactory from a taxonomic and ecological point of view, particularly in the north and northeastern regions of the country. This study provides the first annotated list of echinoids from northeastern Brazil. The studied material largely belongs to collections of the Federal University of Paraíba (CIPY), Federal University of Sergipe (LABIMAR-UFS), Federal University of Bahia (MZUFBA), University of São Paulo (MZUSP), and National Museum of the Federal University of Rio de Janeiro (MNRJ). Thirty-two species from 29 genera, 18 families, and 10 orders were identified. Descriptions of species are provided. Highest diversities of Echinoidea were encountered for the states of Bahia (19 spp.), Alagoas (11 spp.), Paraíba (10 spp.), Ceará (7 spp.), Rio Grande do Norte (7 spp.), and Pernambuco (6 spp.). On the basis of the data analysed, Maranhão (2 spp.), Piauí (2 spp.), and Sergipe (3 spp.) have the lowest diversity. Sandy substrates and depths below 10 m were the least sampled areas over the continental shelf. Although the studied species are common, some taxonomic problems were encountered and discussed. We also provide ecological information and comments on status of the species from the studied region. As a result of this inventory, we were able to provide the first assessment of the echinoid fauna of northeastern Brazil.

Plesiodiadema antillarum (A. Agassiz, 1880) Figure 5 A–G Aspidodiadema antillarum A. Agassiz, 1880: 73 –74. Dermatodiadema antillarum A. Agassiz, 1898: 76. Dermatodiadema (Plesiodiadema) antillarum Mortensen, 1904: 37, pl. 3, fig. 18, pl. 4, figs 8, 16, 29, 33, 35, pl. 5, figs 4, 32. Plesiodiadema antillarum H.L. Clark, 1925: 39.– Tommasi, 1966a: 12, figs 64–65.– Campos & Moura, 2008: 136 (table), 139. Plesiodiadema microtuberculatum Mironov, 1981: 175 –177, figs 1a, b, d, g, and 5, pl. 1, figs 5–6, pl. 2, figs 4, 6, 7 [not Plesiodiadema microtuberculatum A. Agassiz, 1879]. Material examined. Rio Grande do Norte: 2 spms, Potiguar Basin, V.2011, 2094 m [UFPB/ECH.2293]. Description. Test small, high, spherical or ovoid, and fragile (Fig. 5 A– C). Aboral surface slightly flattened. Apical system monocyclic, genital and ocular plates nearly uniform in size. Genital plate almost pentagonal, with large number of small tubercles. Gonopore large and located almost in centre of genital plate. Ocular plate slightly rectangular (with rounded corners), broader than genital plate, with large and some small tubercles. Periproct circular, with circle of periproctal plates of irregular sizes and densely covered by long spines surrounding anus (Fig. 5A). Ambulacra narrow, with tubercles and spines much smaller than in interambulacra (Fig. 5C). Interambulacra with two series of large, strongly crenulate primary tubercles, secondary tubercles small and uniform (Fig. 5C). Ambulacral plates simple (unigeminate). Primary spines very long, slender, glassy, verticillate, curved downwards and extremely fragile (Fig. 5A). Secondary spines similar to primary spines, but shorter, thinner, and straighter (Fig. 5A, B). Tubercles perforate and crenulate (Fig. 5C). Peristome with shallow, rounded buccal notches (Fig. 5B). Ten large and naked buccal plates, covering nearly whole peristome, each with single buccal tube foot (Fig. 5B). Pedicellariae (description based on the figures in Mortensen, 1904). There are small and large forms of ophicephalous pedicellariae (Fig. 5D, E). The large form has pyramidal valves (Fig. 5D), while the small form has short valves, with the distal portion diamond-shaped (with rounded corners) with serrate margin (Fig. 5E). Some small ophicephalous pedicellariae have a large, thin purse of tissue covering the stalk. Triphyllous pedicellariae elongate, with ovoid distal portions (Fig. 5F). Tridentate pedicellariae long, with stalk and neck long. Valve narrow and serrate (Fig. 5G). Colour. When alive this species is light violet or greyish pink, with spines of the same tint (A. Agassiz 1880). It may have a dark purplish red test and spines (Schultz 2011). Distribution. Amphiatlantic (Turner & Graham 2003). In the West Atlantic, from Florida, Gulf of Mexico, Mexico, Bahamas, Cuba, Haiti, Jamaica, Trinidad and Tobago, Puerto Rico, Venezuela, Guyana, and Brazil (Mironov 2006; Laguarda-Figueras et al. 2005b; Alvarado 2011; Solís-Marín et al. 2013). In Brazil, only known from off Maceió (AL) (H.L. Clark 1925). We present herein the first record for the State of Rio Grande do Norte (Potiguar Basin). It occurs from 289 to 5278 m (Mironov 2006; Smithsonian Database). Specimens from Potiguar Basin were collected at 2094 m. Remarks. The genus Plesiodiadema contains seven extant species, of which only P. antillarum occurs in the Atlantic Ocean. Mortensen (1940) stressed that P. antillarum and P. microtuberculatum are very similar, and that the differences between them are not clear. It is possible that they represent a single species (Schultz 2011). However, no study has been conducted to evaluate the taxonomic status of these taxa. According to Schultz (2011), P. antillarum differs from its congeners by having only a few small periproctal plates lying in the centre of the membrane. Moreover, the coronal plates are less numerous than in other species of the genus and the shape of the tridentate pedicellariae is constant in P. antillarum (Mironov 2006). According to H.L. Clark (1925), the specimen of P. antillarum from H.M.S. “Challenger” St. 122 (off eastern Brazil) is very young, and its identity therefore not certain. Clark (1925) remarks that all specimens collected in Brazil were initially erroneously identified as P. microtuberculatum. On the basis of the collected material, we confirm the presence of P. antillarum for the Brazilian continental shelf. We note that Brazilian material has a slightly lower test than the specimens from the Caribbean. A. Agassiz (1883) did not find noteworthy differences among young and adult individuals, and small differences were considered as related to ontogenetic development. Ecological notes. There is little available information on the biology and ecology of P. antillarum. This species feeds on mud and detritus, and does not appear to ingest plant material (Serafy & Fell 1985; Pawson 1982). Emlet (1995) reported that P. antillarum has planktotrophic larvae., Published as part of Gondim, Anne Isabelley, Moura, Rafael Bendayan De, Christoffersen, Martin Lindsey & Dias, Thelma Lúcia Pereira, 2018, Taxonomic guide and historical review of echinoids (Echinodermata: Echinoidea) from northeastern Brazil, pp. 1-72 in Zootaxa 4529 (1) on pages 18-19, DOI: 10.11646/zootaxa.4529.1.1, http://zenodo.org/record/2612564, {"references":["Agassiz, A. (1880) Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Caribbean Sea in 1878 - 79, and along the Atlantic Coast of the United States during summer of 1880, by the U. S. Coast Survey Steamer \" Blake \". IX. Preliminary report on the Echini. Bulletin of the Museum of Comparative Zoology at Harvard College, 8, 69 - 84. Available from: http: // www. biodiversitylibrary. org / item / 95375 page / 81 / mode / 1 up (accessed 31 July 2017)","Agassiz, A. (1898) Reports on the dredging operations off the west coast of Central America to the Galapagos, to the the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U. S. Fish Commission Streamer \" Albatross \", during 1891, Lieut. Commander Z. L. Tanner, U. S. N., Commanding. XXIII. Preliminary report on the Echini. Bulletin of the Museum of Comparative Zoology, 32, 71 - 86. Available from: http: // www. biodiversitylibrary. org / page / 30426278 page / 185 / mode / 1 up (accessed 14 July 2017)","Mortensen, T. (1904) The Danish Expedition to Siam 1899 - 1900. III. Echinoidea (1). Kongelige Danske Videnskabelige Selskabs. Skrifter, Series 7, 1, 1 - 124. Available from: http: // www. biodiversitylibrary. org / page / 7323340 page / 21 / mode / 1 up (accessed 14 July 2017)","Clark, H. L. (1925) A catalogue of the recent sea-urchins (Echinoidea) in the Collection of the British Museum. Trustees of the British Museum, London. The Oxford University Press, London, 250 pp.","Tommasi, L. R. (1966 a) Lista dos Equinoides recentes do Brasil. Contribuicoes do Instituto Oceanografico, Serie Oceanografia Biologica, 11, 1 - 50.","Campos, L. S. & Moura, R. B. (2008) Macrostructure and evolution of the digestive system in Echinoida (Echinodermata). Zoomorphology, 127, 135 - 141. https: // doi. org / 10.1007 / s 00435 - 008 - 0058 - 4","Mironov, A. N. (1981) Deep sea echinoids of the genus Plesiodiadema (Echinoidea, Aspidodiadematidae). Trudy Instituta Oceanologii, 115, 174 - 182.","Agassiz, A. (1879) Preliminary report on the Echini of the Exploring Expedition of H. M. S \" Challenger \". Proceedings of the American Academy of Arts and Sciences, New Series, 6, 190 - 212. Available from: https: // archive. org / stream / proceedingsofame 14 amer page / 199 / mode / 1 up (accessed 31 July 2017)","Schultz, H. A. G. (2011) Sea urchins III, Worldwide regular deep-water species. Heinke & Peter Schultz Partner Scientific Publications, Hemdingen, 478 pp. [pp. 861 - 1338 pp.]","Turner, R. L. & Graham, B. D. (2003) Calocidaris micans (Cidaridae) and Pseudoboletia maculata (Toxopneustidae): additions to the sea urchin fauna (Echinodermata: Echinoidea) of the Gulf of Mexico. Proceedings of the Biological Society of Washington, 116 (1), 61 - 81. Available from: http: // biostor. org / reference / 76237 (accessed 10 July 2017)","Mironov, A. N. (2006) Echinoids from seamounts of the north-eastern Atlantic; onshore-offshore gradients in species distribution. In: Mironov, A. N., Gebruk, A. V. & Southward, A. J. (Eds.), Biogeography of the North Atlantic seamounts. KMK Science Press Ltd., Moscow, pp. 96 - 133.","Laguarda-Figueras, A., Gutierrez-Castro, A. I., Solis-Marin, F. A., Duran-Gonzalez, A. & Torres-Veja, J. (2005 b) Equinoideos (Echinodermata: Echinoidea) del Golfo de Mexico. Revista de Biologia Tropical, 53 (Supplement 3), 69 - 108.","Alvarado, J. J. (2011) Echinoderm diversity in the Caribbean Sea. Marine Biodiversity, 41, 261 - 285. https: // doi. org / 10.1007 / s 12526 - 010 - 0053 - 0","Solis-Marin, F. A., Alvarado, J. J., Abreu-Perez, M., Aguilera, O., Alio, J., Bacallado-Aranega, J. J., Barraza, E., Benavides- Serrato, M., Benitez-Villalobos, F., Betancourt-Fernandez, L., Borges, M., Brandt, M., Brogger, M. I., Borrero-Perez, G. H., Buitron-Sanchez, B. E., Campos, L. S., Cantera, J., Clemente, S., Cohen-Renjifo, M., Coppard, S., Costa-Lotufo, L. V., del Valle-Garcia, R., Diaz, Y., Diaz de Vivar, M. E., Diaz-Martinez, J. P., Duran-Gonzalez, A., Epherra, L., Escolar, M., Francisco, V., Freire, C. A., Garcia-Arraras, E., Gil, D. G., Guarderas, P., Hadel, V. F., Hearn, A., Hernandez, J. C., Hernandez-Delgado, E. A., Herrera-Moreno, A., Herrero-Perezrul, M. D., Hooker, Y., Honey-Escandon, M. B. I., Lodeiros, C., Luzuriaga, M., Manso, C. L. C., Martin, A., Martinez, M. I., Martinez, S., Moro-Abad, L., Mutschke, E., Navarro, J. C., Neira, R., Noriega, N., Palleiro-Nayar, J. S., Perez, A. F., Perez-Ruzafa, A., Prieto-Rios, E., Reyes, J., Rodriguez, R., Rubilar, T., Sancho-Mejia, T., Sangil, C., Silva, J. R. M. C., Sonnenholzner, J. I., Ventura, C. R., Tablado, A., Tavares, Y., Tiago, C. G., Tuya, F. & Williams, S. M. (2013) Appendix. In: Alvarado, J. J. & Solis-Marin, F. A. (Eds.), Echinoderm Research and Diversity in Latin America. Springer, Berlin / Heidelberg, pp. 543 - 654.","Mortensen, T. H. (1940) A Monograph of the Echinoidea. Aulodonta, with additions to Vol. II (Lepidocentra and Stirodonta). Vol III. 1. C. A. Reitzel, Copenhagen, 370 pp.","Agassiz, A. (1883) Report on the Echini. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877 - 78) in the Caribbean Sea (1878 - 79), and along the Atlantic coast of the United States (1880), by the U. S. Coast survey steamer \" Blake \". XXIV. Part I. Cambridge. Memoirs of the Museum of Comparative Zoology at Harvard College, 10 (1), 1 - 94. Available from: http: // www. biodiversitylibrary. org / page / 30719676 page / 104 / mode / 1 up (accessed 21 October 2016)","Serafy, D. K. & Fell, F. J. (1985) Marine flora and fauna of the Northeastern United States. Echinodermata: Echinoidea. NOAA Technical Report National Marine Fisheries Service, 33, 1 - 27.","Pawson, D. L. (1982) Papers from the Echinoderm Conference. 8. Deep-sea echinoderms in the Tongue of the Ocean, Bahamas Islands: a survey, using the research submersible Alvin. Australian Museum Memoir, 16, 129 - 145. https: // doi. org / 10.3853 / j. 0067 - 1967.16.1982.362","Emlet, R. B. (1995) Developmental mode and species geographic range in regular sea urchins (Echinodermata: Echinoidea). Evolution, 49 (3), 476 - 489. https: // doi. org / 10.1111 / j. 1558 - 5646.1995. tb 02280. x"]}

Family Aspidodiadematidae Duncan, 1889 Genus Plesiodiadema Pomel, 1883, Published as part of Gondim, Anne Isabelley, Moura, Rafael Bendayan De, Christoffersen, Martin Lindsey & Dias, Thelma L��cia Pereira, 2018, Taxonomic guide and historical review of echinoids (Echinodermata: Echinoidea) from northeastern Brazil, pp. 1-72 in Zootaxa 4529 (1) on page 18, DOI: 10.11646/zootaxa.4529.1.1, http://zenodo.org/record/2612564, {"references":["Duncan, P. M. (1889) A revision of the genera and great groups of the Echinoidea. Journal of the Linnean Society of London, Zoology, 23, 1 - 311. https: // doi. org / 10.1111 / j. 1096 - 3642.1889. tb 01431. x","Pomel, A. (1883) Classification methodique et Genera des Echinides vivantes et fossiles. Theses presentees a la Faculte des Sciences de Paris pour obtenir le Grade de Docteur es Sciences Naturelles 503. Adolphe Jourdan, Alger, 131 pp. Available from: http: // www. biodiversitylibrary. org / item / 43038 (accessed 23 May 2011)"]}

Aspidodiadema africanum Mortensen, 1939 Fig. 24 A���C. Aspidodiadema africanum n. sp.: Mortensen, 1939: 548. Aspidodiadema nicobaricum: H.L. Clark, 1923 *: 371���372; H.L. Clark, 1925a: 39. Aspidodiadema africanum: Clark & Courtman-Stock, 1976: 223, Schultz, 2011: 1118, Fig. 1964. Material examined. MBC-A 022216. Identification. Test small, sub-globular. Tubercles perforate and crenulate. Primary spines slender, brittle, hollow, or with a partially-filled core, serrated. Peristome encircled by club-shaped spines, and buccal plates. Preserved specimen light brown to purplish with white spines. Global maximum size. Maximum test diameter 16 mm. Global distribution. Endemic to South coast of South Africa, at 720���820 m depth (Mortensen, 1939; Clark & Courtman-Stock 1976). Remarks. Represented by just one specimen (MBC-A 022216) in the Iziko collection., Published as part of Filander, Zoleka & Griffiths, Charles, 2017, Illustrated guide to the echinoid (Echinodermata: Echinoidea) fauna of South Africa, pp. 1-72 in Zootaxa 4296 (1) on pages 26-27, DOI: 10.11646/zootaxa.4296.1.1, http://zenodo.org/record/843325, {"references":["Mortensen, T. (1939) New Echinoida (Aulodonta). Preliminary Notice. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i KObenhavn, 103, 547 - 550.","Clark, H. L. (1923) The echinoderm fauna of South Africa. Annals of the South African Museum, 13 (7), 221 - 435.","Clark, H. L. (1925 a) A Catalogue of the Recent Sea-urchins (Echinoidea) in the Collection of the British Museum (Natural History). Trustees of the British Museum, London, 250 pp.","Clark, A. & Courtman-Stock, J. (1976) The Echinoderms of Southern Africa. Publication no. 766. British Museum of Natural History, London, 277 pp.","Schultz, H. (2011) Sea-Urchins III, Worldwide Regular Deep Water Species. 1 st Edition. Scientific Publications, Hemdingen, 1338 pp."]}

Here we use histological sections and, to a limited extent, scanning electron microscopy to ascertain whether siphons or siphonal grooves parallel the stomach of five sea urchin species representing five higher taxa in the Echinoidea. We find a siphonal groove in Centrostephanus coronatus of the Diadematidae and in Caenopedina diomedeae of the Pedinidae; we find a siphon in Tromikosoma panamense of the Echinothurioida and in Strongylocentrotus purpuratus of the Camarodonta; and we find a hemisiphon in Aspidodiadema hawaiiense of the Aspidodiadematidae (it is currently unsettled whether this last group belongs in the Pedinoida or the Diadematoida). Several recent accounts of echinoid gut anatomy have claimed that species in the Echinothurioida have a siphonal groove instead of a siphon and that species in the Diadematidae and Pedinidae have siphons instead of siphonal grooves. The present work shows that these claims are mistakes and confirms that Holland and Ghiselin (1970) correctly described the distribution of siphons and siphonal grooves in the Echinoidea.

The sea-urchins (class Echinoidea) of South Africa have received little taxonomic attention since the last comprehensive regional monographic guide, which was published in 1976 and is thus now severely outdated. That account was also poorly illustrated, making it difficult to use. Recent work by the authors has also added some 19 additional species to the known fauna, which now stands at 70 species. The aims of this study are thus to revise and update the South African echinoid fauna list, and to present a new, easy to use and fully-illustrated guide to all known regional species. The analysis is based mainly on examination of samples from the Iziko South African Museum collections, although additional records from other museum collections, the published literature, photographic images submitted to the EchinoMap Virtual Museum Database and any other reliable records are also included. A short account of each species is given, including relevant synonymy and literature, and brief notes on identification, size, and global distribution. Photographs that best represent the diagnostic features of each species and a map showing its known records within the region accompany each species account. A binary key to species is also provided.

The complete larval development of an echinoid in the family Aspidodiadematidae is described for the first time from in vitro cultures of Aspidodiadema jacobyi, a bathyal species from the Bahamian Slope. Over a period of 5 months, embryos grew from small (98-micron) eggs to very large (3071-micron) and complex planktotrophic echinopluteus larvae. The fully developed larva has five pairs of red-pigmented arms (preoral, anterolateral, postoral, posterodorsal, and posterolateral); fenestrated triangular plates at the bases of fenestrated postoral and posterodorsal arms; a complex dorsal arch; posterodorsal vibratile lobes; a ring of cilia around the region of the preoral and anterolateral arms; and a long, unpaired posterior process containing a fenestrated rod. The presence of a posterior process and posterodorsal arms makes the larva of Aspidodiadema jacobyi much more similar to larvae of irregular urchins in the order Spatangoidea than to other families of the order Diadematoida, to which the family is normally assigned. This unexpected larval form lends support to a recommendation that the Aspidodiadematidae should be either elevated to ordinal status as a sister group of the order Diadematoida, or split off as a sister group of the other families within the order. In either case, if we accept the parsimonious hypothesis that the aboral process and posterodorsal arms were derived only once in the evolutionary history of euechinoids, then the larval data suggest that the Aspidodiadematidae may be very near the node where the irregular and regular euechinoids first diverged.