taxonID	type	description	language	source
C4255B094E3AFFE6FF551A9AFD86FD82.taxon	description	Figures 3 A – M, 18 A, 19 A	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3AFFE6FF551A9AFD86FD82.taxon	materials_examined	Material examined. Paraíba: Project Algas — 1 spm, 6 ° 55 ′ S 34 ° 48 ′ W, st. 58, 12 m, 04. II. 1981 [UFPB / ECH. 1362]; 4 spms, 7 ° 10 ′ S 34 ° 38 ′ W, st. 37, 25 m, 26. III. 1981 [UFPB / ECH. 1620]; 2 spms, 6 ° 37 ′ S 34 ° 51 ′ W, st. 80, 20 m, 02. VI. 1981 [UFPB / ECH. 1621]; 1 spm, 6 ° 46 ′ S 34 ° 50 ′ W, st. 68, 14 m, 19. V. 1981 [UFPB / ECH. 1624]; 2 spms, 7 ° 12 ′ 5 ″ S 34 ° 36 ′ W, st. 31, 26 m, 01. IV. 1981 [UFPB / ECH. 1626]; 1 spm, 7 ° 21 ′ S 34 ° 38 ′ W, st. 19, 16 m, 13. V. 1981 [UFPB / ECH. 1627]; 1 spm, 7 ° 13 ′ S 34 ° 42 ′ W, st. 33, 20 m, 27. III. 1981 [UFPB / ECH. 1628]; 1 spm, 7 ° 31 ′ S 34 ° 31 ′ W, st. 5, 34 m, 29. I. 1981 [UFPB / ECH. 1636]; 7 spms, 7 ° 34 ′ S 34 ° 45 ′ W, st. 1, 11 m, 21. I. 1981 [UFPB / ECH. 1638]; 1 spm, 7 ° 10 ′ S 34 ° 38 ′ W, st. 37, 25 m [UFPB / ECH. 1640]; 3 spms, 6 ° 37 ′ 5 ″ S 34 ° 46 ′ W, st. 78, 35 m, 29. V. 1981 [UFPB / ECH. 1643]; 1 spm, 7 ° 07 ′ S 34 ° 37 ′ W, st. 38, 27 m, 25. III. 1981 [UFPB / ECH. 1727]; 6 spms, 6 ° 55 ′ S 34 ° 46 ′ 05 ″ W, st. 58, 12 m, 06. II. 1981 [UFPB / ECH. 1728]; 4 spms, 7 ° 04 ′ S 34 ° 38 ′ W, st. 45, 26 m, 17. II. 1981 [UFPB / ECH. 1734]; 8 spms, 6 ° 26 ′ S 34 ° 52 ′ W, st. 93, 26 m, 11. VI. 1981 [UFPB / ECH. 1735]; 3 spms, 7 ° 31 ′ S 34 ° 42 ′ W, st. 7, 16 m, 23. I. 1981 [UFPB / ECH. 1738]; 2 spms, 7 ° 01 ′ S 34 ° 30 ′ W, st. 47, 26 m, 13. II. 1981 [UFPB / ECH. 1739]; 9 spms, 7 ° 15 ′ 05 ″ S 34 ° 36 ′ W, st. 29, 28 m, 02. IV. 1981 [UFPB / ECH. 1755]; 3 spms, 7 ° 12 ′ 5 ″ S 34 ° 39 ′ W, st. 32, 20 m, 01. IV. 1981 [UFPB / ECH. 1756]; 1 spm, 6 ° 50 ′ S 34 ° 47 ′ W, st. 65, 18 m, 12. III. 1981 [UFPB / ECH. 1757]; 8 spms, 7 ° 13 ′ S 34 ° 45 ′ W, st. 34, 10 m, 27. III. 1981 [UFPB / ECH. 1758]; 2 spms, 6 ° 50 ′ S 34 ° 50 ′ W, st. 66, 10 m, 12. III. 1981 [UFPB / ECH. 1759]; 1 spm, 6 ° 55 ′ S 34 ° 43 ′ 05 ″ W, st. 56, 21 m, 11. II. 1981 [UFPB / ECH. 1761]; 2 spms, 7 ° 34 ′ S 34 ° 42 ′ W, st. 2, 20 m, 21. IV. 1981 [UFPB / ECH. 1762]; 1 spm, 7 ° 10 ′ S 34 ° 45 ′ W, st. 35, 10 m, 26. III. 1981 [UFPB / ECH. 1763]; 1 spm, 7 ° 34 ′ S 34 ° 36 ′ W, st. 4, 33 m, 22. I. 1981 [UFPB / ECH. 1764]; 1 spm, 7 ° 34 ′ S 34 ° 39 ′ W, st. 3, 26 m, 22. I. 1981 [UFPB / ECH. 1765]; 3 spms, 6 ° 57 ′ S 34 ° 38 ′ W, st. 54, 30 m, 12. II. 1981 [UFPB / ECH. 1766]; 2 spms, 6 ° 33 ′ S 34 ° 51 ′ W, st. 85, 20 m, 04. VI. 1981 [UFPB / ECH. 1767]; 4 spms, 6 ° 33 ′ S 34 ° 47 ′ W, st. 86, 26 m, 04. VI. 1981 [UFPB / ECH. 1768]; 1 spm, 7 ° 07 ′ S 34 ° 43 ′ W, st. 40, 17 m, 19. III. 1981 [UFPB / ECH. 1769]; 2 spms, 6 ° 55 ′ S 34 ° 40 ′ W, st. 55, 28 m, 11. II. 1981 [UFPB / ECH. 1770]; 2 spms, 7 ° 04 ′ S 34 ° 41 ′ W, st. 44, 22 m, 16. II. 1981 [UFPB / ECH. 1771]; 2 spms, 7 ° 15 ′ 5 ″ S 34 ° 33 ′ W, st. 30, 35 m, 01. IV. 1981 [UFPB / ECH. 1777]; 1 spm, 7 ° 01 ′ S 34 ° 30 ′ W, st. 47, 26 m, 13. II. 1981 [UFPB / ECH. 1785]; 1 spm, 7 ° 28 ′ S 34 ° 37 ′ W, st. 11, 24 m, 05. V. 1981 [UFPB / ECH. 1786]; 1 spm, 6 ° 52 ′ S 34 ° 42 ′ W, st. 61, 20 m, 20. II. 1981 [UFPB / ECH. 1787]; 1 spm, 7 ° 34 ′ S 34 ° 39 ′ W, st. 3, 26 m, 22. I. 1981 [UFPB / ECH. 1788]; 1 spm, 6 ° 57 ′ S 34 ° 41 ′ W, st. 53, 26 m, 12. II. 1981 [UFPB / ECH. 1789]; 1 spm, 6 ° 43 ′ S 34 ° 45 ′ W, st. 71, 22 m, 20. V. 1981 [UFPB / ECH. 1790]; 3 spms, 7 ° 25 ′ S 34 ° 34 ′ W, st. 15, 14 m, 07. V. 1981 [UFPB / ECH. 1810]; 1 spm, 7 ° 21 ′ 9 ″ S 34 ° 31 ′ W; Geomar XXIV, st. 97, 38 m, 30. V. 1985 [UFPB / ECH. 1821]. Pernambuco: 1 spm, Ponta de Serrambi, Ipojuca, 05. IX. 1982 [UFPB / ECH. 1820]; 1 spm, Fernando de Noronha Archipelago, 28. IV. 2001 [UFPB / ECH. 1917]. Alagoas: 2 spms, Saco da Pedra Reef, Maceió, 10. II. 2009 [UFSITAB- 230]; 3 spms, Saco da Pedra Reef, 10. II. 2009 [UFSITAB- 230]; 1 spm, Francês Beach, Marechal Deodoro, 13. XI. 1983 [UFPB / ECH. 1337]; 15 spms, Francês Beach, Marechal Deodoro, 02. II. 1983 [UFPB / ECH. 1813]; 4 spms, Francês Beach, Marechal Deodoro, 19. II. 2011 [UFPB / ECH. 1902]. Bahia: 2 spms, Itapuã Beach, Salvador, 16. II. 2007 [UFSITAB- 75]; 15 spms, Itapuã Beach, 21. XII. 1984 [UFPB / ECH. 1736]; 2 spms, Itapuã Beach, 17. IX. 1982 [UFPB / ECH. 1737]; 2 spms, Pitanga Beach, Lauro de Freitas, 16. IX. 1982 [UFPB / ECH. 1814]; 1 spm, Ponta de Aratuba, Itaparica, 18. X. 1982 [UFPB / ECH. 1815]; 1 spm, Ponta da Coroa Vermelha, Santa Cruz de Cabrália, 15. X. 1982 [UFPB / ECH. 1816]; 3 spms, Arembepe Beach, Camaçari, 16. IX. 1982 [UFPB / ECH. 1817]; 1 spm, Pituba Beach, Salvador, 17. IX. 1982 [UFPB / ECH. 1818].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3AFFE6FF551A9AFD86FD82.taxon	description	Description. Test circular, slightly flattened both orally and aborally (TD = 3.3 to 45.4 mm; TH = 1.4 to 30.2 mm) (Fig. 3 A – C, G, H). Apical system hemicyclic, uniformly tuberculate (Fig. 3 G, J). Periproct pentagonal, covered by tuberculated plates that decrease in size towards the anus (Fig. 3 G, J). Ambulacra narrow and slightly sinuous (Fig. 3 G, H, L). Interambulacra wide, with clear vertical series of primary spines (Fig. 3 G, H, M). Ambulacral plating simple (Fig. 3 L). Interambulacral plates bearing single, perforate and non-crenulate primary tubercle, surrounded by many scrobicular tubercles (Fig. 3 M). Primary spines thick and cylindrical (Fig. 3 A – C), arranged into 10 vertical series, ending in crown of small projections. Secondary spines short and compressed. Oral-most primary spines often banded dirty white and light pinkish. Peristomial membrane densely covered by flat and imbricated small plates arranged in vertical parallel series (Fig. 3 K). Tube foot spicules are rods with several blunt short spines and elongated flattened plates with perforations restricted to middle and one side (Fig. 3 E). Pedicellariae. Tridentate and globiferous pedicellariae spread over the whole test, the latter more numerous around the primary spines and peristome. Fistulate globiferous pedicellariae small, without a neck. Valve oval, with serrated margin, and denticulate foramen (Fig. 3 F). Tridentate pedicellariae short, without a neck. Valve long, narrow, with denticulate margin (Fig. 3 D). Colour. Ground colour of the test is light brown to red-brown. The muscle bases of all secondary spines are brown to red-brown, spine are shafts dirty white, and the distal ends are lightly tinged with light brown (Hendler et al. 1995). Naked test greenish, except for pore zones, which are white.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3AFFE6FF551A9AFD86FD82.taxon	distribution	Distribution. North Carolina, Gulf of Mexico, Mexico, Guatemala, Belize, Cuba, Honduras, Jamaica, Nicaragua, Haiti, Dominican Republic, Costa Rica, Puerto Rico, Panama, Colombia, Venezuela, and Brazil (Hendler et al. 1995; Alvarado et al. 2008; Solís-Marín et al. 2013; Kroh & Mooi 2016). In Brazil from PE, AL, BA, SP, RJ, and SC, including Abrolhos Archipelago, São Pedro and São Paulo Archipelago, Fernando de Noronha, Trindade and Martin Vaz Islands (H. L. Clark 1907; Tommasi 1958 a; Brito 1962; Lima-Verde 1969; Tommasi & Aron 1988; Fernandes et al. 2002; Magalhães et al. 2005; Netto et al. 2005; Manso et al. 2008; Correia & Sovierzoski 2009; Xavier 2010; Martins et al. 2018). This is the first record of this species for the coast of Paraíba. From 0 to 800 m, but most commonly in less than 50 m (Hendler et al. 1995).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3AFFE6FF551A9AFD86FD82.taxon	discussion	Remarks. The genus Eucidaris contains five living and three fossil species (Kroh & Mooi 2016). In their phylogenetic study of the genus, Lessios et al. (1999) did not find differences between E. tribuloides from the west Atlantic and E. tribuloides africana, from Africa. Their results suggest that Atlantic Eucidaris belong to a single panmictic unit over the entire western Atlantic (Lessios et al. 1999). No morphological differences were found among the specimens in the present study. However, Bernasconi (1955) observed some variations in the aspect of ambulacral and interambulacral areas and in the ocular plate / periproct ratio in specimens sampled from the Brazilian coast. The size and thickness of primary spines in E. tribuloides can vary with habitat (Hendler et al. 1995). In high hydrodynamic environments, the spines tend to be short and thick, while in low energy environments they are longer and thin. However, this trend was not observed in the specimens examined, since they all had short and thick spines, apart from young individuals, which exhibited slightly longer and thinner spines. Ecological notes. This species lives in small cavities in the reef habitat, under rocks and gravel in the backreef or in seagrass beds (del Valle García et al. 2005). Eucidaris tribuloides is omnivorous; its diet varies according to food availability in the environment (Lawrence 1975; Machado et al. 2008). Pieces of algae and bryozoans, coral fragments, gastropod shells, echinoid spines, sponges, and seagrass have been recorded as food items (Hendler et al. 1995). This species is common in the intertidal zone along the coasts of Alagoas and Bahia, while in the remaining northeastern states it seems to be more restricted to depths greater than 10 m. Few specimens were observed in reef crevices and among macroalgae in the coral reef formations of Tamandaré, State of Pernambuco. On the other hand, the species is abundant on the reefs of Praia do Francês (State of Alagoas), where it occurs on under rocks and co-occurs with crinoids, sponges, and other invertebrates. At this beach, we observed specimens hosting an ectoparasitic gastropod belonging to the family Eulimidae Philippi (Fig. 19 A). Numerous specimens were reported by del Valle García et al. (2005) from strongly polluted areas such as “ Playa del Chivo ” and the coast adjacent to the river “ Quibú ”. Along the Brazilian littoral, specimens are being removed in large quantities, mainly to supply the demands of aquarium enthusiasts (Machado et al. 2008). Martins et al. (2012 a) evaluated echinoderm harvesting in Todos os Santos Bay (BA) for local aquarium trade and listed E. tribuloides as the eighth most collected species for this purpose.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E38FFE1FF551857FBF6FC76.taxon	description	Figure 4 A – G	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E38FFE1FF551857FBF6FC76.taxon	materials_examined	Material examined. 2 spms, 15 ° 18,873 ′ S 38 ° 41,685 ′ W, Oceanographic vessel Thalassa, 11. VI. 2000 [EqMN 2341].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E38FFE1FF551857FBF6FC76.taxon	description	Description. Test thin, flexible, contour circular (TD = 67.4 mm). Preserved test becomes totally flattened (Fig. 4 A, C). Apical system monocyclic, with plating often reduced and genital pores surrounded by number of small platelets (Fig. 4 B). Periproct large (Fig. 4 A, B). Interambulacra about two-thirds wider than ambulacra at ambitus (Fig. 4 A). Primary spines long, thin, straight, and hollow. Aboral surface with some fleshy spines (live specimens). Secondary spines thin and elongate (like needles). Spines from oral surface similar to those from aboral surface, but shorter and with rounded tips. Ambulacral plate trigeminate, with large central element bearing one or two primary tubercles and pair of ambulacral pores, and two demiplates housing one pore-pair each (Fig. 4 E). Interambulacral plates wide, with one to three primary tubercles forming subequal rows (Fig. 4 F). Primary tubercles perforated and non-crenulate (Fig. 4 E, F). Aboral tubercles irregularly arranged. Oral surface flat, covered by numerous and large tubercles with sunken areoles (Fig. 4 C). Peristome large, nearly circular and densely covered by overlapped plates (Fig. 4 C). Pedicellariae. Only tridentate (Fig. 4 G) and ophicephalous pedicellariae are found in P. placenta. The first type occurs in smaller numbers than the second type. Both are very numerous and distributed irregularly over entire test, being particularly abundant on the oral surface towards the edge of the peristome. Colour. In living specimens, the aboral surface is gray with purple spots and oral surface purplish (Bell 1892; Mortensen 1977). The tube feet are purple, and the spines are nearly colourless (Thomson 1874). Specimens become brownish-yellowish in alcohol (Borrero-Pérez et al. 2012).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E38FFE1FF551857FBF6FC76.taxon	distribution	Distribution. Bahamas, North and South Carolina, Virginia, Florida, Gulf of Mexico, Mexico, Honduras, Dominica, Bermudas, Jamaica, Cuba, Haiti, Nicaragua, Puerto Rico, Panama, Colombia, Venezuela, and Brazil (Alvarado 2011; Solís-Marín et al. 2013). In Brazil from BA, ES and RJ (Ventura et al. 2007 a; Campos et al. 2010). From depths of 38 to 4100 m (Mironov 2014; Smithsonian Database), but it is very rare at depths shallower than 500 m (Serafy & Fell 1985).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E38FFE1FF551857FBF6FC76.taxon	discussion	Remarks. Phormosoma contains four species, of which only P. placenta occurs in the Atlantic Ocean. Three subspecies are recognized for P. placenta: P. placenta placenta Thomson, 1872 a typical form in the North and East Atlantic, P. placenta sigsbei A. Agassiz, 1880 in the West Atlantic, and P. placenta africana Mortensen, 1934 in South Africa (Mironov 2014). These subspecies are distinguished mainly on the basis of the number of coronal plates and in the size and disposition of the tubercles on aboral surface. Thomson (1872) published an abstract in which he established the genus Phormosoma and the species P. placenta. However, in this work, the author provided only a short description of the genus and the locality of the material used to designate the species. Thomson (1874) presented a detailed description and provided several illustrations of the taxonomic characters of the species. One of the most conspicuous characters in live specimens of Phomosoma is the presence of aboral fleshy spines, which may partially or totally obscure the dorsal surface of the test or form a “ fence ” along the periphery of the urchin. According to Emson & Young (1998), virtually all the primary spines of P. placenta are to some extent fleshy, as all the oral and lateral spines have a small fleshy enlargement near the tip and the blunt oral walking spines are enlarged by a thick fleshy covering. The main function of these spines most probably is defence (Emson & Young 1998). Ecological notes. Individuals of P. placenta seem to aggregate and this pattern was attributed to their slowmoving deposit-feeding mode of life (Grassle et al. 1975). Pawson (1982) found only mucus-consolidated mud balls in the guts of this species. Tyler & Gage (1984) believe that the species must have a relatively rapid growth rate (up to a TD of 40 to 45 mm). At this point, gonad development begins and there is a slowing of somatic growth, resulting in an accumulation of adults in the 55 to 75 mm TD range. Above 75 mm TD, the population numbers decline, and generally few individuals larger than 85 mm TD are found. Tyler & Gage (1984) recorded specimens with 45 mm TD possessing developing gonads, while specimens between 47 and 55 mm TD showed no evidence of reproductive development. According to Pearse (1969), the lecithotrophic larvae of P. placenta are not demersal and develop at or close to the surface. Young & Cameron (1987) conducted a study on flotation rates of lecithotrophic eggs from P. placenta and pointed out the enhanced physiological tolerance in embryos of deepwater echinothuriids relative to larvae of shelf and coastal echinoderms. According to Benavides-Serrato & Borrero-Pérez (2010), this species is abundant in the Colombian part of the Caribbean.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3FFFE0FF551FD6FE8BFA7E.taxon	materials_examined	Material examined. No specimens available for study.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3FFFE0FF551FD6FE8BFA7E.taxon	description	Description (modified from Tommasi 1966 a, Coppard & Campbell 2006 b, and Rodríguez et al. 2013). Test hemispherical, large, and slightly flattened orally and aborally. Apical system hemicyclic (I, IV and V inserted) and depressed. Ocular plates short and pentagonal, with one or two small tubercles. Genital plates wider than long, slightly triangular, with one to three tubercles along their inner edge. Periproct bearing totally black, small anal cone. Ambulacra narrow and slightly swollen, with two rows of primary tubercles. Primary tubercles of ambulacra much smaller than those of interambulacra. Interambulacra wide, with four series of large primary tubercles. In living specimens, a narrow blue line of iridophores occurs down either side of naked median area of the interambulacra and as a ring around the apical disc (or totally black in some specimens). White spots on naked areas of interambulacra easily distinguished at night. Ambulacral plates trigeminate. Primary spines long (around 5 to 6 times the TD), slender, verticillate and hollow internally, blackish or with alternating white and black stripes (in juveniles). Secondary spines similar to primaries, but smaller. Tubercles perforate and crenulate. Peristomial membrane black and covered with many of triphyllous pedicellariae. Buccal plates without spines. Pedicellariae. Only tridentate and triphyllous pedicellariae are present. Tridentate pedicellariae occur in two forms, one with broad valves and a long broad neck on a short stalk, and another form with narrow valves, short neck, and a long stalk. Triphyllous pedicellariae with broad valves rounded distally, with two rows of numerous, small peripheral teeth (Rodríguez et al. 2013). Colour. According to Tommasi (1966 a, 1972), young specimens of D. antillarum have the typical spines with alternating white and black stripes, while adults are whitish, blackish, or greenish. Hendler et al. (1995) further stated that both colour of the test and the spines were typically black, but may have few or many white or gray spines, and some specimens were almost entirely white. In general, black individuals live in clear-water environments, while light-coloured individuals live in less-lighted areas, in turbid waters, rock crevices or deep waters (Hendler et al. 1995).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3FFFE0FF551FD6FE8BFA7E.taxon	distribution	Distribution. Bermudas, Gulf of Mexico, Cuba, Puerto Rico, Costa Rica, Panama, Colombia, Venezuela, and Brazil (Borrero-Pérez et al. 2002; del Valle García et al. 2005; Alvarado 2011). In Brazil, from BA, and RJ, including Trindade and Fernando de Noronha Islands (Tommasi 1966 a; Alves & Cerqueira 2000; del Valle García et al. 2005; Magalhães et al. 2005). From 0 to 400 m, being more common in depths below 50 m (Hendler et al. 1995).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3FFFE0FF551FD6FE8BFA7E.taxon	discussion	Remarks. Presently, eight extant and two fossil taxa of Diadema are known, of which only D. antillarum and D. ascensionis are recorded from the Western Atlantic. D. antillarum differs from D. ascensionis by tridentate pedicellariae with straight valves, and six series of primary tubercles on the interambulacra. Diadema antillarum also differs from D. setosum (Leske, 1778) by not having an orange ring on the anal cone nor genital plates with four or more small tubercles, as in the latter species. Diadema antillarum differs from D. africanum Rodríguez, Hernández, Clemente & Coppard, 2013 by the occurrence of two forms of tridentate pedicellariae, and two to four small tubercles on genital plate. The description of Diadema africanum was based on a population previously identified as D. antillarum-b (see Rodríguez et al. 2013). Considering the close morphological affinity among D. antillarum, D. ascencionis and D. africanum (Rodríguez et al. 2013), taxonomical, biological and ecological information about these species must be interpreted with caution. Oliveira (1951) recorded D. setosum and Diadema sp. at Trindade Island and Brito (1962) cited D. setosum for Fernando de Noronha and Bahia. However, Tommasi (1966 b) believed that these identifications were incorrect since D. setosum is a typically Indo-Pacific species and its occurrence in these areas is unlikely. Thus, it seems more plausible that the specimens identified by Oliveira and Brito were, in fact, D. antillarum or D. ascensionis. Currently, the records of these latter two species are considered valid for the Brazilian coast. The test of this species is so fine and fragile that it remains intact only for a short time after death (Kidwell & Baumiller 1990; Greenstein 1989, 1991, 1993). Ecological notes. This species inhabits reef environments, seagrass beds, mangroves and sandy or rocky bottoms (del Valle García et al. 2005). According to Hendler et al. (1995), the species prefers calm water, actively avoiding heavy wave action. Diadema antillarum is considered omnivorous, feeding preferably on algae (Weil et al. 2005). However, when foraging high-density habitats, it ingests invertebrates such as zoanthids, sponges and coral tissues (Randall et al. 1964), thus affecting the recruitment and mortality rates of young coral and other sessile invertebrates (Sammarco 1980). There are many studies regarding biology and ecology of D. antillarum (e. g. Randall et al. 1964; Lewis 1966; Levitan 1989; Atrill & Kelmo 2007; Blanco et al. 2011; Rodríguez-Barrera et al. 2015).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3EFFE3FF551D23FD95FA91.taxon	description	Figure 18 B	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3EFFE3FF551D23FD95FA91.taxon	description	Lima-Verde, 1969: 10. – Gondim et al., 2013 a: 431 – 435, figs 1 b – d, 1 a – h. – Martins et al., 2018: 526, figs 4 – 5.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3EFFE3FF551D23FD95FA91.taxon	materials_examined	Material examined. Pernambuco: 1 spm, Rata Island, Fernando de Noronha, 13. IV. 1999, 10 m [MZUSP, without voucher]. Alagoas: 2 spms, Francês Beach, Marechal Deodoro, 20. II. 2011 [UFPB / ECH. 1910]; 4 spms, Francês Beach, 29. I. 1983 [UFPB / ECH. 1922]. Bahia: 1 spm, Itapuã Beach, Salvador, 29. XI. 2006 [UFBA 00151].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3EFFE3FF551D23FD95FA91.taxon	description	Description (modified from Gondim et al. 2013 a). Test large, circular, oral region flat. Apical system depressed and hemicyclic. Ocular plates with one or two small tubercles. Genital plates with only one tubercle or with no tubercles at all. Periproct with small, black anal cone. Ambulacra swollen and narrow. Interambulacra large. Ambulacral plates trigeminate, with one primary tubercle. Interambulacral plates with one or three primary tubercles. Primary spines long, thin, hollow, axial cavities in tips of spines filled with dense, non-reticular tissue. Secondary spines similar to primaries, but smaller. Tubercles perforate and crenulate. Peristome twice as large as periproct. Pedicellariae. Triphyllous and tridentate pedicellariae distributed all over test surface, being more abundant on the aboral surface. Claviform pedicellariae short and distributed around primary spines. Tridentate pedicellariae with narrow and strongly curved valves. Triphyllous pedicellariae with long stalk and neck. Valves long and narrow, with proximal region narrowest and broadening towards the tip (Gondim et al. 2013 a). Colour. Specimens are black, with blue lines in the interambulacra and around the apical system (usually appearing completely black). White spots in central zones of naked areas of interambulacrum. Naked test white.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3EFFE3FF551D23FD95FA91.taxon	distribution	Distribution. Gulf of Mexico, Jamaica, Puerto Rico, Panama, Barbados, and Brazil, including Ascension and St. Helena Islands (Tommasi 1966 a; Coppard & Campbell 2006 a). In Brazil from AL, and BA, including Fernando de Noronha, Atol das Rocas, Trindade, and Martin Vaz Islands (Brito 1968, 1971; Lima-Verde 1969; Fernandes et al. 2002; Martins et al. 2018). From intertidal to 400 m depth (Tommasi 1966 a). In the present study, specimens from Alagoas were collected about 2 m deep.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3EFFE3FF551D23FD95FA91.taxon	discussion	Remarks. Diadema ascensionis differs from D. antillarum by the strongly curved valves in the tridentate pedicellariae. Since its original description, the taxonomic status of D. ascensionis has been debated, having been considered a synonym or a subspecies of D. antillarum. Presently it is considered a valid species, but molecular data (Lessios 2001; Lessios et al. 2001) conflict with morphology. However, several studies have found differences to support its designation as a distinct species (Coppard & Campbell 2004, 2006 a, 2006 b; Gondim et al. 2013 a; Rodríguez et al. 2013). Specimens studied herein, from reefs at Francês Beach (Alagoas), had all the characters that permit its distinction from D. antillarum. However, the specimens from this locality were previously identified as D. antillarum. To distinguish these two species correctly, an accurate analysis of the pedicellariae and of the structure of the spines is necessary. We analyzed both adult and young individuals. In distinction to other sea urchins, D. ascensionis has a hemicyclic apical system in all developmental stages, with the possible exception of post-larvae (not observed by us). Ecological notes. In the coastal reefs of Francês Beach, State of Alagoas, D. ascensionis was found in areas with sandy substrates or among rocks of the reef formations, in zones of clear and well-illuminated water. Both isolated and clusters of individuals were observed, dense populations occurring up to 2 m depth (Gondim et al., 2013 a). Martins et al. (2018) observed a gregarious habit in specimens from Trindade and Martin Vaz Islands, where specimens were usually found in reef crevices during the day and moving around areas with a high percentage of green algae cover.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3DFFFCFF551EFEFA02FE35.taxon	materials_examined	Material examined. No specimens available for study.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3DFFFCFF551EFEFA02FE35.taxon	description	Description (modified from A. H. Clark 1934, Mortensen 1940 and Hendler et al. 1995). Test large, very low, fairly flexible and slightly pentagonal in dorsal view. Apical system monocyclic. Genital plates large, triangular, projecting strongly interradially. Ocular plates small and rectangular. Periproct covered marginally by small and variably shaped plates. Anal cone without plates. Ambulacra slightly swollen. Ambulacral plating trigeminate, with pairs of pores arranged in arches. Ambulacral plates closest to peristome without primary tubercles. Interambulacra wide, broadest at middle height of test, with extensive, naked, Y-shaped areas on aboral surface. Primary spines long, thin, fragile, verticillate, banded with reddish-brown and whitish-yellow stripes, and filled with calcite meshwork. Secondary spines similar to primary spines, but weaker and thinner. Tubercles perforate and crenulate. Number of tubercles increases towards peristome. Peristome covered by small, fragile, broadened plates. Pedicellariae (modified from Mortensen 1940). Claviform pedicellariae well-developed. Tridentate pedicellariae small, with very narrow valves. Triphyllous pedicellariae as in A. pulvinata (Lamarck, 1816) and A. radiata (Leske, 1778). For more detail, see Mortensen (1940: 206, pl. LXX, figs 4 – 6). Colour. Naked areas of test golden yellow. Brown ambulacra bordered by single rows of brilliant, iridescent blue spots. The inflated anal cone is bluish white, with a brown peripheral ring and a dark brown anus. The spines are banded in reddish brown and yellowish white (Hendler et al. 1995). Juvenile can be pink-reddish with green spines banded by red stripes (Humann & Deloach 2002).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3DFFFCFF551EFEFA02FE35.taxon	distribution	Distribution. North Carolina, South Carolina, Mexico, Gulf of Mexico, Barbados, Cuba, Dominican Republic, Jamaica, Puerto Rico, Belize, Panama, Colombia, Venezuela, Surinam, and Brazil (Espinosa et al. 1997; del Valle García et al. 2005; Laguarda-Figueras et al. 2005 a; Alvarado et al. 2008; Solís-Marín et al. 2013; Rodríguez-Barreras 2014). In Brazil from PE, BA, and SC (Magalhães et al. 2005; Lima & Fernandes 2009). From 2 to 89 m depth (Solís-Marín et al. 2013; Rodríguez-Barreras 2014).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E3DFFFCFF551EFEFA02FE35.taxon	discussion	Remarks. A. H. Clark (1934) described A. magnifica on the basis of four specimens collected 88 m south of Dry Tortugas. Presently, two species are known from the coast of Brazil [A. magnifica and A. nuptialis Tommasi, 1958 b]. The first is widely distributed, from North Carolina to southern Brazil, while records of the second are restricted to São Paulo and Santa Catarina. Tommasi (unpublished data) considers A. magnifica and A. nuptialis synonyms. We reach the same conclusion on the basis of comparisons between published descriptions. However, a more detailed comparison involving type material is necessary. According to A. H. Clark (1934), juveniles of A. magnifica resemble A. pulvinata more closely than the adults. Ecological notes. This species, called the " king urchin " in Brazil, lives in sheltered habitats with sandy substrates, fragments of calcareous algae, gravel and calcareous outcrops. It has also been observed in muddy substrates and in meadows of Halophila decipiens Ostenfeld, 1902 (del Valle García et al. 2005). Astropyga magnifica is rarely observed in situ in Brazil, although it seems to have a patchy distribution and to be gregarious in some Caribbean localities (Hendler et al. 1995). Kier & Grant (1965) observed it moving across the sand flats in groups of two to five, at a speed of approximately 3 feet per minute, concluding that it is highly mobile. Lessios (1990) considered egg size and recorded mean values of 104.04 µm for the longest axis and 5.59 × 10 5 µm 3 for egg volume. Mortensen (1940) and De Ridder & Lawrence (1982) recorded shell fragments in the stomach contents. Commensal species, mainly shrimps, have been recorded (Kier & Grant 1965). Williams et al. (1986) reported an event of mass mortality of this species in the shallow-waters on the northwest coast of Puerto Rico in the winter of 1984 – 1985. Since then, no similar event has been reported in the literature. Between the years of 1996 and 2005, 226 specimens were captured along the littoral of the State of Bahia by aquarium hobbyists (Martins et al. 2012 a).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E22FFFFFF551999FC17FD16.taxon	description	Figure 5 A – G	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E22FFFFFF551999FC17FD16.taxon	materials_examined	Material examined. Rio Grande do Norte: 2 spms, Potiguar Basin, V. 2011, 2094 m [UFPB / ECH. 2293].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E22FFFFFF551999FC17FD16.taxon	description	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. 5 A). Ambulacra narrow, with tubercles and spines much smaller than in interambulacra (Fig. 5 C). Interambulacra with two series of large, strongly crenulate primary tubercles, secondary tubercles small and uniform (Fig. 5 C). Ambulacral plates simple (unigeminate). Primary spines very long, slender, glassy, verticillate, curved downwards and extremely fragile (Fig. 5 A). Secondary spines similar to primary spines, but shorter, thinner, and straighter (Fig. 5 A, B). Tubercles perforate and crenulate (Fig. 5 C). Peristome with shallow, rounded buccal notches (Fig. 5 B). Ten large and naked buccal plates, covering nearly whole peristome, each with single buccal tube foot (Fig. 5 B). Pedicellariae (description based on the figures in Mortensen, 1904). There are small and large forms of ophicephalous pedicellariae (Fig. 5 D, E). The large form has pyramidal valves (Fig. 5 D), while the small form has short valves, with the distal portion diamond-shaped (with rounded corners) with serrate margin (Fig. 5 E). Some small ophicephalous pedicellariae have a large, thin purse of tissue covering the stalk. Triphyllous pedicellariae elongate, with ovoid distal portions (Fig. 5 F). Tridentate pedicellariae long, with stalk and neck long. Valve narrow and serrate (Fig. 5 G). 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).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E22FFFFFF551999FC17FD16.taxon	distribution	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. 2005 b; 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.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E22FFFFFF551999FC17FD16.taxon	discussion	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.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E21FFF9FF5518FBFDC9F861.taxon	description	Figures 6 A – M, 18 C, 19 B	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E21FFF9FF5518FBFDC9F861.taxon	materials_examined	Material examined. Rio Grande do Norte: 4 spms, Ponta de Cajaúna, Tibau do Sul, 19. VIII. 1982 [UFPB / ECH. 22]; 11 spms, Pipa Beach, Tibau do Sul, 19. VIII. 1982 [UFPB / ECH. 25]; 7 spms, Barra de Maxaramguape, Ceará-Mirim, 21. VIII. 1982 [UFPB / ECH. 26]; 4 spms, Cabo do Cabopari, Baía Formosa, 18. VIII. 1982 [UFPB / ECH. 27]; 1 spm, Cabo de São Roque, 22. VIII. 1982 [UFPB / ECH. 1611]; 2 spms, Pirangi do Sul, Nísia Floresta, 22. VIII. 1982 [UFPB / ECH. 1642]; 3 spms, Zumbi Beach, Rio do Fogo, 3 – 6 m, 27. XI. 2009 [MZUSP, without voucher]; 2 spms, 5 ° 19,908 ′ S 35 ° 21,607 ′ W, Zumbi Beach, 27. XI. 2009 [MZUSP, without voucher]. Paraíba: 2 spms, Ponta do Seixas, João Pessoa, 8. V. 1977 [UFPB / ECH. 01]; 3 spms, Cabo Branco Beach, João Pessoa, 12. V. 1980 [UFPB / ECH. 02]; 2 spms, Ponta do Seixas, João Pessoa, 12. II. 1982 [UFPB / ECH. 03]; 3 spms, Cabo Branco Beach, 03. VII. 1985 [UFPB / ECH. 21]; 5 spms, Cabo Branco Beach, 12. III. 1981 [UFPB / ECH. 351]; 1 spm, Ponta de Campina, Cabedelo, 24. VIII. 2006 [UFPB / ECH. 352]; 3 spms, Cabo Branco Beach, 04. XI. 2006 [UFPB / ECH. 354]; 1 spm, Cabo Branco Beach, 09. IX. 2006 [UFPB / ECH. 355]; 1 spm, 6 ° 55 ' S 34 ° 47 ′ W, 10 m, 03. VII. 2006 [UFPB / ECH. 573]; 1 spm, Cabo Branco Beach, 26. IX. 2007 [UFPB / ECH. 702]; 5 spms, Coqueirinho Beach, Conde, 27. X. 2007 [UFPB / ECH. 777]; 2 spms, Seixas Reef, 22. XII. 2007 [UFPB / ECH. 1018]; 4 spms, Baía da Traição Beach, 24. XI. 2007 [UFPB / ECH. 1020]; 1 spms, Barra de Camaratuba Beach, Mataraca, 23. I. 2008 [UFPB / ECH. 1146]; 2 spms, Areia Vermelha Reef, Cabedelo, 22. II. 2008 [UFPB / ECH. 1148]; 2 spms, Baía da Traição Beach, 04. V. 2008 [UFPB / ECH. 1181]; 2 spms, Barra de Camaratuba Beach, Mataraca, 06. V. 2008 [UFPB / ECH. 1182]; 3 spms, Areia Vermelha Reef, Cabedelo, 06. IV. 2008 [UFPB / ECH. 1212]; 1 spm, Cabo Branco Beach, 28. X. 2007 [UFPB / ECH. 1315]; 5 spms, Cabo Branco Beach, 16. XII. 1997 [UFPB / ECH. 1317]; 3 spms, Cabo Branco Beach, 25. X. 2007 [UFPB / ECH. 1319]; 2 spms, Cabo Branco Beach, 21. III. 2000 [UFPB / ECH. 1320]; 2 spms, Ponta do Seixas, 25. V. 1998 UFPB / ECH. 1322]; 4 spms, Cabo Branco Beach, 07. VI. 2001 [UFPB / ECH. 1323]; 3 spms, Santa Catarina Beach, Cabedelo, 18. V. 2007 [UFPB / ECH. 1324]; 3 spms, Tambaba Beach, Conde, 28. IX. 2007 [UFPB / ECH. 1325]; 12 spms, Cabo Branco Beach, 0 9. III. 2005 [UFPB / ECH. 1326]; 1 spm., Cabo Branco Beach, 25. X. 2007 [UFPB / ECH. 1328]; 3 spms, Ponta do Seixas, João Pessoa, 03. XI. 1982 [UFPB / ECH. 1329]; 2 spms, Cabo Branco Beach, 19. IX. 2005 [UFPB / ECH. 1331]; 3 spms, Cabo Branco Beach, 09. II. 2001 [UFPB / ECH. 1335]; 18 spms, Cabo Branco Beach, 01. XII. 1975 [UFPB / ECH. 1336]; 9 spms, Galé Reef, Pitimbú, 12. XII. 2008 [UFPB / ECH. 1455]; 2 spms, Barra de Mamanguape Reef, Rio Tinto, 10. I. 2009 [UFPB / ECH. 1456]; 2 spms, Areia Vermelha Reef, Cabedelo, 06. IV. 2008 [UFPB / ECH. 1472]; 3 spms, Cabo Branco Beach, 13. XI. 2008 [UFPB / ECH. 1473]; 1 spm, Pier de Cabedelo, Cabedelo, 05. V. 2008 [UFPB / ECH. 1476]; 5 spms, Seixas Reef, João Pessoa, 12. I. 2009 [UFPB / ECH. 1477]; 2 spms, Maceiozinho Beach, Jacumã, 06. V. 2008 [UFPB / ECH. 1482]; 2 spms, Tabatinga Beach, Conde, 10. II. 2009 [UFPB / ECH. 1512]; 3 spms, Coqueirinho Beach, Conde, 03. VI. 2008 [UFPB / ECH. 1513]; 1 spm, Tambaba, Beach Conde, 10. II. 2009 [UFPB / ECH. 1514]; 2 spms, Barra de Mamanguape Reef, Rio Tinto, 11. II. 2009 [UFPB / ECH. 1515]; 1 spm, Cabo Branco Beach, 03. XI. 1982 [UFPB / ECH. 1649]; 1 spm, Pitimbú Reef, Pitimbú, 16. IX. 2006 [UFPB / ECH. 1656]; 2 spms, Pitimbú Reefs, Pitumbú, 16. IX. 2005 [UFPB / ECH. 1675]; 1 spm, Cabo Branco Beach, 16. III. 2002 [UFPB / ECH. 1701]; 1 spm, Cabo Branco Beach, 29. V. 2009 [UFPB / ECH. 1702]; 1 spm, Cabo Branco Beach, 09. IV. 2005 [UFPB / ECH. 1729]; 2 spms, Tambaba Beach, Conde, 04. VI. 2008 [UFPB / ECH. 1730]. Pernambuco: 13 spms, rocky shore between the Gabu Beach and Cabo de Santo Agostinho, 04. IX. 1982 [UFPB / ECH. 04]; 6 spms, Itarema Beach, Cabo, 03. IX. 1982 [UFPB / ECH. 05]; 6 spms, Ponta de Pedras, 30. X. 1982 [UFPB / ECH. 11]; 3 spms, Candeias Reef, Jaboatão, 02. XI. 1982 [UFPB / ECH. 12]; 1 spm, Piedade Reef, Jaboatão, 2. XI. 1982 [UFPB / ECH. 13]; 3 spms, Gabu Reef, Cabo, 03. IX. 1982 [UFPB / ECH. 29]; 1 spm, São José da Coroa Grande, 06. IX. 1982 [UFPB / ECH. 1316]; 5 spms, Pau Amarelo Reef, Paulista, 01. XI. 1982 [UFPB / ECH. 1327]; 1 spm, Catuama, Goiana, 31. X. 1982 [UFPB / ECH. 1332]; 2 spms, Gamela Sirinhaén, 05. IX. 1982 [UFPB / ECH. 1659]; 1 spm, Ponta de Serrambi Beach, Ipojuca, 05. IX. 1982 [UFPB / ECH. 1670]. Alagoas: 1 spm, Pajuçara Beach, Maceió, 21. IX. 1982 [UFPB / ECH. 06]; 3 spms, Francês Beach, Marechal Deodoro, 19. X. 1982 [UFPB / ECH. 10]; 3 spms, Riacho Doce Beach, 31. I. 1983 [UFPB / ECH. 16]; 3 spms, Ponta Verde Beach, Maceió, 20. X. 1982 [UFPB / ECH. 28]; 1 spm, Ponta Verde Beach, 31. I. 1983 [UFPB / ECH. 1330]; 1 spm, Paripueira Beach, 01. II. 1983 [UFPB / ECH. 1653]; 2 spms, Francês Beach, Marechal Deodoro, 29. I. 1983 [UFPB / ECH. 1655]; 1 spm, Pontal do Cururipe, Curuipe, 26. I. 1983 [UFPB / ECH. 1733]; 1 spm., Saco da Pedra Reef, Maceió, 10. II. 2009 [UFSITAB- 206]; 1 spm, Saco da Pedra Reef, 10. II. 2009 [UFSITAB- 208]; 1 spm, Saco da Pedra Reef, 10. II. 2009 [UFSITAB- 250]. Bahia: 4 spms, Ponta da Coroa Vermelha, Santa Cruz da Cabrália, 15. X. 1982 [UFPB / ECH. 08]; 1 spm, Ponta Grande Reef, Porto Seguro, 15. X. 1982 [UFPB / ECH. 09]; 2 spms, South of the Redonda Island, Abrolhos, 01. II. 1982 [UFPB / ECH. 15]; 5 spms, Ponta de Imbaçuaba, Cumuruxatiba, 13. X. 1982 [UFPB / ECH. 23]; 1 spm, reef between Cumuruxatiba and Ponta de Ibaçuaba, Prado, 28. IX. 1982 [UFPB / ECH. 30]; 4 spms, south-west of Siriba Islands, Abrolhos, 30. XI. 1982 [UFPB / ECH. 31]; 4 spms, Imbaçuaba Beach, Prado, 14. X. 1982 [UFPB / ECH. 33]; 1 spm, Pituba Beach, Salvador, 17. IX. 1982 [UFPB / ECH. 1318]; 1 spm, Pitanga Beach, Lauro Freitas, 16. IX. 1982 [UFPB / ECH. 1333]; 1 spm, Coroa Beach, Itaparica, 18. IX. 1982 [UFPB / ECH. 1334]; 1 spm, Pituba Beach, Salvador, 17. IX. 1982 [UFPB / ECH. 1634]; 1 spm, Penha Beach, Vera Cruz, Itaparica, 08. IX. 1982 [UFPB / ECH. 1654]; 3 spms, Arembepe Beach, Camaçari, 16. IX. 1982 [UFPB / ECH. 1658]; 1 spm, Itaparica Beach, 22. XII. 1984 [UFPB / ECH. 1674].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E21FFF9FF5518FBFDC9F861.taxon	description	Description. Test elliptical, oral surface concave (TD = 5.8 to 129 mm; TH = 2.5 to 45.5 mm) (Fig. 6 A, B, I, J). Apical system hemicyclic (Fig. 6 K). Plates of periproct and apical system densely covered by small spines (Fig. 6 K). Ocular and genital plates with many tubercles of various sizes (Fig. 6 K). Ambulacral plate polygeminate (echinoid type), with six pairs of pores arranged in arc, one primary tubercle and few secondary tubercles (Fig. 6 L). Interambulacral plate with one primary tubercle and many secondary tubercles (Fig. 6 M). Primary and secondary tubercles of ambulacra and interambulacra non-crenulate and non-perforate (Fig. 6 I, L, M). Primary spines of variable length, thick at base and thin at tip (Fig. 6 A). Secondary spines thin and short. Peristome large, with few buccal plates densely covered by purple ophicephalous pedicellariae (Fig. 6 B). Tube feet with C-shaped spicules and ossicles forming rosette in terminal discs (Fig. 6 G, H). Spicules of tube feet often organized into longitudinal rows. Pedicellariae. Globiferous, ophicephalous, tridentate and triphyllous pedicellariae present over entire test. Globiferous pedicellariae of various sizes, with a long stalk, without a neck, and with three pairs of well-developed venom glands, in which C-shaped spicules are found. Valves with lateral tooth paired or unpaired and a single end tooth (Fig. 6 C, D). Ophicephalous pedicellariae with a long stalk and short neck. Valves with small marginal teeth that are serrate (like shark teeth) (Fig. 6 E). Tridentate pedicellariae short, with stalk and neck long, in which Cshaped spicules similar to those of the tube feet are found. Valves narrow, long, with margins slightly serrate (Fig. 6 F). Triphyllous pedicellariae with a long neck, about the same length as stalk. Valves short and broad, being narrower at the base and widening towards the tip. Colour. Echinometra lucunter can be black, gray, green, red, or even bluish, displaying a high degree of colour polymorphism (Lessios 1998). The spines are black but may have a greenish tinge at the base and a purple tinge at the apex. Naked test white.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E21FFF9FF5518FBFDC9F861.taxon	distribution	Distribution. Mexico, Cuba, Belize, Guatemala, Honduras, Nicaragua, Jamaica, Haiti, Dominican Republic, Caribbean Sea, Costa Rica, Panama, Puerto Rico, Madagascar, Mozambique, Colombia, Venezuela, and Brazil (del Valle García et al. 2008; Alvarado et al. 2008; Kroh & Mooi 2016). In Brazil from CE, PB, PE, AL, BA, ES, RJ, SP, SC, including Abrolhos, Fernando, Fernando de Noronha, Trindade and Martin Vaz Islands (Rathbun 1879; H. L. Clark 1925; Brito 1962, 1968; Tommasi 1966 a; Lima-Verde 1969; Fernandes et al. 2002; Magalhães et al. 2005; Gondim et al. 2008, Lima & Fernandes 2009; Xavier 2010; Miranda et al. 2012; Martins et al. 2018). Herein we first record the species for the State of Rio Grande do Norte. The species is known to occur between 0 and 45 m (Hendler et al. 1995), but it is almost exclusively a shallow-water species, restricted to less than 1 m of depth (Lessios 1998).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E21FFF9FF5518FBFDC9F861.taxon	discussion	Remarks. Presently eight species of Echinometra are known worldwide (Bronstein & Loya 2013), two of which occur in the tropical part of the Western Atlantic [E. lucunter and E. viridis A. Agassiz, 1863]. Only E. lucunter occurs along the Brazilian coastline. Echinometra lucunter differs from E. viridis in not having ambulacral plates with five pairs of pores and a reddish brown test with yellowish spines. In this study, we observed small morphological differences among adults and young individuals, such as the dicyclic apical system with few tubercles in young individuals and hemicyclic with many tubercles in adults. The length and thickness of spines were also quite variable, from long and thin to short, thick, and blunt. Apparently, these variations were not related to environment (low or high energy habitats) nor to developmental stage, but further studies are necessary to confirm this observation. Young specimens tend to have spines that are thin and long. Lewis & Storey (1984) observed that individuals living in high energy habitats tend to have thicker, smaller, and more flattened tests than specimens from low energy habitats. Ecological notes. This species inhabits the intertidal and subtidal zones of reef environments but is common in the surf zone, where it lives inside crevices excavated by the action of their spines and teeth on the rock substrate (Hendler et al. 1995) (Fig. 19 B). According to Espinoza et al. (2008), this species can be also be found in sandy substrates and banks of seagrass (e. g., Thalassia testudinum K. D. Koenig, 1805). Echinometra lucunter is nocturnal and lives almost permanently inside of its burrow. Only a few individuals leave their burrows at night to feed. The species feeds primarily on drifting algae (McPherson 1969), but may also prey on marine invertebrates such as sponges and cnidarians (McClintock et al. 1982; McClanahan & Muthiga 2007). Echinometra is considered a " gardener " species, feeding on the algae that grow inside of its burrow, moving back and forth along the burrow to allow algae to regenerate as it feeds on new growth (McClanahan & Muthiga 2007; Mooi, personal communication). Griinbaum et al. (1978) reported an agonistic behaviour in individuals of this species. During our field surveys, we observed some specimens of E. lucunter exhibiting " covering behaviour ". Echinometra lucunter is the most common species in the intertidal zone of the NE coast of Brazil, forming high-density populations in these areas (Fig. 19 B). Due to its habit of excavating rocks and forming dense populations, this species plays an important role in the bioerosion of reef environments (Hendler et al. 1995) (Fig. 20 B). Echinometra lucunter, as a specialized herbivore, also has an important role in the reef food web, where it acts in controlling macroalgal cover. It is also the most important prey of the gastropod Cassis tuberosa (Linnaeus, 1758) throughout the Western Atlantic (Dias et al. 2017). According to Martins et al. (2012 a), E. lucunter was among the sea urchin species captured for the aquarium trade on the coast of Salvador (BA). In Cabo de Santo Agostinho (PE), there is a traditional event called " Festa da Ouriçada " (Sea Urchin Festival), which occurs every December 13 th since 1960, in devotion to Saint Luzia. Local fishermen capture sea urchins in large numbers to consume their gonads among local families (Santos & Andrade 2013).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E24FFF4FF551A15FCBDFE6E.taxon	description	Figures 7 A – O, 18 D, 19 C – D	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E24FFF4FF551A15FCBDFE6E.taxon	materials_examined	Material examined. Paraíba: 1 spm, Cabo Branco Beach, João Pessoa, 25. IX. 2007 [UFPB / ECH. 1158]; 1 spm, Cabo Branco Beach, 21. III. 2000 [UFPB / ECH. 1321]; 1 spm, Cabo Branco Beach, 20. XI. 1980 [UFPB / ECH. 1364]; 1 spm, Galé Reef, Pitimbú, 12. XII. 2008 [UFPB / ECH. 1461]; 2 spms, Cabo Branco Beach, 29. V. 2009 [UFPB / ECH. 1552]; 1 spm, 7 ° 07 ′ 00 ″ S 34 ° 43 ′ 54 ″ W, 14. III. 2006 [UFPB / ECH. 1718]; 1 spm, Bessa Beach, João Pessoa, 20. VIII. 2005 [UFPB / ECH. 1719]; 1 spm, 6 ° 55 ′ S 34 ° 47 ′ W, 03. VII. 2006, 10 m [UFPB / ECH. 574]; 1 spm, Ponta do Seixas, João Pessoa, 12. IV. 2006 [UFPB / ECH. 1811]; Project Algas — 5 spms, 6 ° 50 ′ S 34 ° 50 ′ W, st. 66, 10 m 12. III. 1981 [UFPB / ECH. 1363]; 1 spm, 6 ° 46 ′ S 34 ° 53 ′ W, st. 67, 10 m, 13. III. 1981 [UFPB / ECH. 1631]; 2 spms, 7 ° 10 ′ S 34 ° 38 ′ W, st. 37, 35 m, 26. III. 1981 [UFPB / ECH. 1639]; 3 spms, 7 ° 01 ′ S 34 ° 41 ′ W, st. 49, 16 m, 06. II. 1981 [UFPB / ECH. 1644]; 1 spm, 7 ° 01 ′ S 34 ° 47 ′ 05 ″ W, st. 50, 11 m, 05. II. 1981 [UFPB / ECH. 1703]; 1 spm, 7 ° 01 ′ S 34 ° 41 ′ 05 ″ W, st. 48, 24 m, 13. II. 1981 [UFPB / ECH. 1704]; 1 spm, 7 ° 28 ′ S 34 ° 40 ′ W, st. 10, 14 m, 05. V. 1981 [UFPB / ECH. 1705]; 5 spms, 6 ° 29 ′ S 34 ° 57 ′ W, st. 90, 12 m, 10. VI. 1981 [UFPB / ECH. 1706]; 4 spms, 6 ° 33 ′ S 34 ° 57 ′ W, st. 83, 12 m, 03. VI. 1981 [UFPB / ECH. 1707]; 3 spms, 7 ° 10 ′ S 34 ° 38 ′ W, st. 37, 25 m, 26. III. 1981 [UFPB / ECH. 1708]; 1 spm, 6 ° 55 ′ S 34 ° 46 ′ 05 ″ W, st. 57, 18 m, 06. II. 1981 [UFPB / ECH. 1709]; 1 spm, 7 ° 10 ′ S 34 ° 45 ′ W, st. 35, 10 m, 26. III. 1981 [UFPB / ECH. 1710]; 1 spm, 7 ° 15 ′ 05 ″ S 34 ° 42 ′ W, st. 27, 16 m, 13. IV. 1981 [UFPB / ECH. 1711]; 2 spms, 6 ° 55 ′ S 34 ° 48 ′ W, st. 58, 12 m, 04. II. 1981 [UFPB / ECH. 1712]; 1 spm, 6 ° 58 ′ S 34 ° 46 ′ 05 ″ W, st. 51, 14 m, 05. II. 1981 [UFPB / ECH. 1713]; 2 spms, 7 ° 34 ′ S 34 ° 42 ′ W, st. 2, 20 m, 21. IV. 1981 [UFPB / ECH. 1714]; 1 spm, 7 ° 13 ′ S 34 ° 42 ′ W, st. 33, 20 m, 27. III. 1981 [UFPB / ECH. 1715]; 1 spm, 6 ° 36 ′ S 34 ° 43 ′ W, st. 79, 22 m, 29. V. 1981 [UFPB / ECH. 1717]; 1 spm, 6 ° 50 ′ S 34 ° 50 ′ W, st. 66, 10 m, 12. III. 1981 [UFPB / ECH. 1760]; 1 spm, 6 ° 52 ′ S 34 ° 48 ′ W, st. 59, 10 m, 04. II. 1981 [UFPB / ECH. 1776]; 1 spm, 7 ° 13 ′ S 34 ° 42 ′ W, st. 33, 20 m, 27. III. 1981 [UFPB / ECH. 1812]. Pernambuco: 4 spms, Itaparica Island, V. 1980 [UFPB / ECH. 24]. Alagoas: 3 spms, Paripueira Beach, 01. II. 1983 [UFPB / ECH. 17]; 1 spm, Mangue Beach, 03. II. 1983 [UFPB / ECH. 18]; 3 spms, Ponta Verde Reef, Maceió, 26. VI. 2007 [UFSITAB- 78]. Bahia: 7 spms, Ponta da Coroa Vermelha, Santa Cruz da Cabrália, 15. X. 1982 [UFPB / ECH. 07]; 4 spms, coral reef between Cumuruxatiba and Imbaçuaba Beach, Prado, 28. XI. 1982 [UFPB / ECH. 14]; 2 spms, Itaparica Beach, Salvador, 22. XII. 1984 [UFPB / ECH. 19]; 4 spms, Coroa Vermelha, Santa Cruz da Cabrália, 20. XII. 1984 [UFPB / ECH. 20]; 3 spms, Imbaçuaba, Prado, 14. X. 1982 [UFPB / ECH. 32]; 4 spms, Cabuçu Beach, Santo Amaro, 19. IX. 1982 [UFPB / ECH. 1720]; 1 spm, Itaparica Beach, Salvador, 22. XII. 1984 [UFPB / ECH. 1721]; 3 spms, Maré Island, Salvador, 01. II. 2002 [UFSITAB- 119]; 1 spm, Santa Cruz da Cabrália, 6 – 7 m [MZUSP, without voucher].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E24FFF4FF551A15FCBDFE6E.taxon	description	Description. Test hemispherical, slightly pentagonal with flat oral side (TD = 4.4 to 81.7 mm; TH = 1.8 to 38.6 mm) (Fig. 7 A, B, I, J). Apical system hemicyclic, ocular plates I and V insert (Fig. 7 L). Genital plates with two or three tubercles (Fig. 7 L). Ocular plates with five to eight small tubercles (Fig. 7 L). Ambulacra broad at ambitus, almost as broad as interambulacra (Fig. 7 I, J). Ambulacral plate trigeminate (echinoid type), with one primary tubercle and three pairs of pores arranged in vertical row (Fig. 7 N). Interambulacral plate with one to five primary tubercles (Fig. 7 O); number of tubercles tend to increase toward oral region. Primary spines short, with tapering tips, and arranged into clear vertical rows in ambulacra and interambulacra. Secondary spines smaller and narrower than primary spines. Tubercles non-crenulate and non-perforate (Fig. 7 I, J, N, O). Peristomial membrane densely covered by peristomial plates (Fig. 7 N), with many ophicephalous and triphyllous pedicellariae. Pedicellariae. Globiferous pedicellariae with a long stalk, without a neck, and with many C-shaped and dumbbell-shaped spicules. Valves long, with a long end tooth (Fig. 7 C, D). White ophicephalous pedicellariae over entire test (Fig. 7 A). Ophicephalous pedicellariae with long stalk and neck, with a small number of C-shaped and dumbbell-shaped spicules (Fig. 7 H). Valves short, oval, with margin denticulate and slightly serrate (Fig. 7 E). Tridentate pedicellariae short, with a long neck and C-shaped and dumbbell-shaped spicules. Valves long and narrow, with margin slightly denticulate, these teeth being short, the longest occurring near base of the blade (Fig. 7 E). Triphyllous pedicellariae with large C-shaped spicules (Fig. 7 H) and short, broad, perforated blade, perforations on external margin longer and narrower (Fig. 7 G). Colour. Corona greenish, with white ambulacra and spines with purplish tips (Fig. 7 A, B). Globiferous pedicellariae large, white, visible to the naked eye (Fig. 7 A). Test and spines colour variable and used to identify subspecies of L. variegatus (Serafy 1973). Hendler et al. (1995) indicated that colour of the test and spines may vary within a single population. We did not observe this variation on populations from the NE Brazil.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E24FFF4FF551A15FCBDFE6E.taxon	distribution	Distribution. Mexico, Gulf of Mexico, Cuba, Jamaica, Haiti, Belize, Guatemala, Honduras, Dominican Republic, Puerto Rico, Caribbean Sea, Costa Rica, Panama, Colombia, Venezuela, and Brazil (Alvarado et al. 2008; del Valle García et al. 2008; Kroh & Mooi 2016). In Brazil, from CE, PB, PE, AL, BA, RJ, SP, SC, RS, including Abrolhos and Fernando de Noronha Islands (Rathbun 1879; H. L. Clark 1925; Tommasi 1966 a, 1972; Brito 1968; Lima-Verde 1969; Tommasi & Aron 1988; Manso 1989; Fernandes et al. 2002; Magalhães et al. 2005; Netto et al. 2005; Gondim et al. 2008; Lima & Fernandes 2009; Oliveira et al. 2010; Xavier 2010). Herein we establish a new record of L. variegatus for the State of Rio Grande do Norte. The species occurs from 0 to 250 m, being more common above depths of 50 m (Hendler et al. 1995).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E24FFF4FF551A15FCBDFE6E.taxon	discussion	Remarks. Lytechinus is currently known from three fossil and eight living species, four of which come from the tropical West Atlantic (L. variegatus; L. callipeplus H. L. Clark, 1912; L. euerces H. L. Clark, 1912; and L. williamsi Chesher, 1968). Only L. variegatus and the subspecies L. variegatus carolinus A. Agassiz, 1863 are known from Brazilian waters. Lytechinus variegatus differs from L. callipeplus by having tridentate and triphyllous pedicellariae and by its peristomial membrane, which is covered by whitish plates. Lytechinus euerces differs from L. variegatus by having test and spines white, and short primary spines. Lytechinus williamsi differs from L. variegatus by having a beige test and large, purple globiferous pedicellariae. Mortensen (1943 a) separated L. variegatus into allopatric subspecies. Tommasi (1966 a) provided a taxonomic key for these subspecies and considered the peristomial membrane, spines, and test colour important characters. Serafy (1973) studied morphological variation in L. variegatus from the West Atlantic and concluded that three (L. variegatus atlanticus, L. variegatus carolinus, and L. variegatus variegatus) of the four subspecies proposed by Mortensen (1933) were valid, raising L. variegatus pallidus H. L. Clark, 1925 to species category. On the other hand, Hendler et al. (1995) highlighted the need for further studies to support validity of these subspecies. A study carried out by Zigler & Lessios (2004), showed that the L. variegatus variegatus mtDNA is distinct from that of L. variegatus carolinus from the North American seaboard and the Gulf of Mexico, whereas their bindins are very similar. Here, young and adult specimens of L. variegatus were examined, and only variation in the apical system arrangement was observed, being dicyclic in the young and hemicyclic in adults. Bernasconi (1955) examined specimens from São Sebastião (SP) and observed variations in the number of secondary tubercles in the interambulacral plates, which can reach up to 2 – 3 in the adapical series and 2 in the adradial series at the ambitus. Ecological notes. This species is preferentially found in calm waters, especially in banks of seagrass and on rocky and sandy substrates (del Valle García et al. 2005). According to Lessios (1998), L. variegatus is normally abundant in Thalassia beds in the Caribbean. Lima-Verde (1968) observed this species in calm waters with rocky or sandy substrates in the coastal areas of Ceará and Pernambuco, Brazil. Recently, this species was observed inhabiting hypersaline estuaries along the semiarid littoral of Rio Grande do Norte, being associated with sand-mud bottoms with a salinity of 42. In general, it is herbivorous (Lawrence 1975), but its food preference is probably conditioned by the defensive chemistry produced by certain species of algae (Souza et al. 2008). Inside its stomach, fragments of bryozoans, Foraminifera, diatoms, and several benthic marine invertebrates have been recorded (Tommasi 1964). Like other species of Toxopneustidae, L. variegatus shows a " covering behaviour " using fragments of shells, gravel, and algae (Hendler et al. 1995) (Fig. 19 C). The same behavior was recorded for specimens from hypersaline habitats. Although there is much controversy about the meaning of this behaviour, it can provide protection against predators or physical factors such as light and temperature, thus preventing body dehydration (Verling et al. 2004). In some areas along northeastern Brazil, L. variegatus displays population outbreaks in shallow, warm waters. At the reefs of Tamandaré, located in the Environmental Protection Area known as the Coral Coast (State of Pernambuco), dense populations of this species are commonly observed during the dry period, between September and February. Martins et al. (2012 a) documented the aquarium fishery of this species along the coast of Salvador (BA). It is also one of the most important prey of the king helmet, Cassis tuberosa, in Brazilian waters (Dias et al. 2017) (Fig. 19 D).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E2AFFF7FF55195EFDADFB7B.taxon	description	Figures 8 A – N, 18 E	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E2AFFF7FF55195EFDADFB7B.taxon	materials_examined	Material examined. Paraíba: 1 spm, Bessa Beach, João Pessoa, 31. V. 1985 [UFPB / ECH. 1361]; Project Algas — 3 spms, 6 ° 39 ′ S 34 ° 49 ′ W, st. 77, 20 m, 28. V. 1981 [UFPB / ECH. 1630]; 1 spm, 7 ° 31 ′ S 34 ° 31 ′ W, st. 5, 34 m, 22. I. 1981 [UFPB / ECH. 1635]; 1 spm, 7 ° 34 ′ S 34 ° 45 ′ W, st. 1, 11 m, 21. I. 1981 [UFPB / ECH. 1637]; 1 spm, 7 ° 28 ′ S 34 ° 40 ′ W, st. 10, 14 m, 5. V. 1981 [UFPB / ECH. 1641]; 3 spms, 6 ° 29 ′ S 34 ° 51 ′ W, st. 88, 22 m, 5. VI. 1981 [UFPB / ECH. 1724]. Alagoas: 1 spm, Francês Beach, Marechal Deodoro, 29. I. 1983 [UFPB / ECH. 1725]; 1 spm, Francês Beach, Marechal Deodoro, 09. II. 2011 [UFPB / ECH. 1916]. Bahia: 2 spms, Itaparica Island, 22. XII. 1985 [UFPB / ECH. 1722]; 1 spm, Itapuã Beach, 21. XII. 1984 [UFPB / ECH. 1723]; 1 spm, Arembepe Beach, Camaçari, 16. IX. 1982 [UFPB / ECH. 1726].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E2AFFF7FF55195EFDADFB7B.taxon	description	Description. Test hemispheric or approaching pentagonal, with oral side flat (TD = 8 to 95.3 mm; TH = 5.6 to 57 mm) (Fig. 8 A, B, J, K). Apical system hemicyclic (Fig. 8 J, L). Plates of apical system bearing many small tubercles (Fig. 8 J, L). Ambulacra and interambulacra approximately same width (Fig. 8 J, K). Ambulacral plates trigeminate, with pores arranged in three vertical series (Fig. 8 M). Interambulacral plate with three to four primary tubercles (Fig. 8 N). Primary tubercles forming three to four vertical series on the ambulacra and four or five horizontal series on each plate of ambitus (Fig. 8 J, K, M, N). Primary spines small, whitish, with several longitudinal striae. Secondary spines similar to primary spines, but smaller. Tubercles non-crenulate and imperforate (Fig. 8 J, K, M, N). Peristomial membrane with few plates, numerous ophicephalous pedicellariae surrounding mouth (close to buccal feet), and few, sparsely distributed triphyllous and tridentate pedicellariae (Fig. 8 B, K). Buccal plates with many ophicephalous pedicellariae (Fig. 8 B). Tube feet with C-shaped spicules (Fig. 8 H). Pedicellariae: Globiferous pedicellariae small and lacking a neck, abundant especially on the apical system and in interambulacra. There are three glands on the stalk and three associated with the valves. Valves long, proximal region broad, blade narrow with a long end tooth (Fig. 8 C, D). C-shaped and dumbbell-shaped spicules found on all pedicellariae, being more numerous on the glands and in the areas close to the end tooth. Ophicephalous pedicellariae with short stalk, long neck (twice as long as stalk), and dumbbell-shaped spicules. Valves elongated, with broad areas for insertion of adductor muscle, and denticulate and serrate margin (Fig. 8 E). Triphyllous pedicellariae with a long neck (almost as long as stalk) and short head, dumbbell-shaped spicules within entire pedicellaria. Valves short, proximal region short and narrower than distal portion, which is broader and longer (Fig. 8 G). Tridentate pedicellariae with neck and stalk of about same size, bearing dumbbell-shaped spicules. Valves short, with base and lamina of the same size, blade serrate and denticulate at the tip (Fig. 8 F). Colour. Test appears uniformly dark brown, including the basal muscles of primary spines (Fig. 8 A, B). Short primary and secondary spines white to dirty white (Hendler et al. 1995) (Fig. 8 A, B). Naked test completely white (Fig. 8 J, K).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E2AFFF7FF55195EFDADFB7B.taxon	distribution	Distribution. Gulf of Mexico, Mexico, Belize, Guatemala, Honduras, Cuba, Haiti, Dominican Republic, Puerto Rico, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, and Brazil (Alvarado 2011; del Valle García et al. 2008; Kroh & Mooi 2016). In Brazil from RN, PE, AL, BA, SP, RJ, and SC, including Fernando de Noronha and Trindade Islands and Abrolhos Archipelago (Rathbun 1879; Bernasconi 1956; Tommasi 1966 a; Lima-Verde 1969; Brito 1971; Tommasi & Aron 1988; Hendler et al. 1995; Fernandes et al. 2002; Magalhães et al. 2005; Lima & Fernandes 2009; Xavier 2010; Martins et al. 2018). In this study, we record T. ventricosus for the State of Paraíba. From 0 to 55 m depth (Hendler et al. 1995).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E2AFFF7FF55195EFDADFB7B.taxon	discussion	Remarks. Tripneustes contains four extant species (Bronstein et al. 2017), of which only T. ventricosus is recorded from the Brazilian littoral. Tripneustes ventricosus differs from its congener T. depressus A. Agassiz, 1863 by having a purple test, only two types of pedicellariae (ophicephalous and tridentate) and less numerous primary spines. Caso (1974 a, b) provided important contributions on the morphology of T. ventricosus and distinguishing characters with regards to T. depressus. According to Hendler et al. (1995), T. ventricosus may reach a TD of 150 mm. In the present study, both young and adult specimens were analysed, but we failed to observe morphological variations, except for, apical system, which was dicyclic in young individuals and hemicyclic in adults. Tripneustes ventricosus is sexually dimorphic, with a long and tubular genital papilla in males, and short and conical genital papilla in females (Turner et al. 2010). Ecological notes. This species lives in areas on sandy substrates with seagrass, and in cracks or in gravel in reef environments (Hendler et al. 1995). In Cuba, it was found in high densities near reefs, or in reef lagoons with or without marine phanerogams (mainly Thalassia testudinum and Syringodium sp.) (del Valle García et al. 2005). It feeds mainly on algae and detritus (Ventura et al. 2007 a). Tripneustes ventricosus, like L. variegatus, displays a “ covering behaviour ” with shell fragments, gravel, and algae (Hendler et al. 1995). Along the littoral of northeastern Brazil, the species is usually not numerous, although population outbreaks have been recorded for some localities, such as the beaches of the Fernando de Noronha Archipelago (State of Pernambuco, Brazil). At Francês Beach (State of Alagoas), the species occurs in protected areas and in crevices and caves, sharing space with crinoids, hydroids, and sponges.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E29FFF6FF551EDFFBC8F9D4.taxon	description	Figure 9 A – D	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E29FFF6FF551EDFFBC8F9D4.taxon	description	Echinoneus orbicularis Desor in L. Agassiz & Desor, 1847: 144. Echinoneus ventricosus L. Agassiz in L. Agassiz & Desor, 1847: 144.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E29FFF6FF551EDFFBC8F9D4.taxon	materials_examined	Material examined. Ceará: 2 spms, Canopos Bank [UFPB. Ech. 1973]. Bahia: 2 spms, 12 ° 57 ′ 28 ″ S 38 ° 21 ′ 22 ″ W, Itapuã Beach, Salvador, 1 m, 19. II. 2007 [UFBA 00493]; 1 spm, Busca Vida Beach, Guarajuba, Camaçari, 21 m, VIII. 2010 [UFBA 01190]; 1 spm, Busca Vida Beach, 23 m, VII. 2010 [UFBA 01191]; 1 spm, Busca Vida Beach, 23 m, VII. 2010 [UFBA 01192].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E29FFF6FF551EDFFBC8F9D4.taxon	description	Description. Test elliptical or oval (TL = 3.3 to 21.9 mm; TH = 2.9 to 9.4 mm), low, with rounded ends (Fig. 9 A, C, D) and densely covered by slender, glassy and short spines. Peristome and periproct on oral surface (Fig. 9 C). Apical system tetrabasal with four genital pores (Fig. 9 A, B). Ambulacra narrow, ambulacra II, III and IV slightly narrower than ambulacra I and V (Fig. 9 A, D). Petals undeveloped, pores double throughout and arranged uniserially. Interambulacra large, 1, 4 and 5 larger than 2 and 3 (Fig. 9 A). Ambulacral plate trigeminate. Tubercles imperforate, smooth and non-crenulate (Fig. 9 A, C, D). Pedicellariae present over whole test. Oral surface flat, except peristome, which is slightly concave (Fig. 9 C). Periproct oval, close to the peristome, about twice the size of peristome (Fig. 9 C). Peristomial plates diminishing in size from margin towards center. Peristome central or slightly anterior, oblique, and depressed (Fig. 9 C). Periproctal plates imbricate, decreasing in size from margin to center. Pedicellariae. According to Westergren (1911), ophicephalous pedicellariae are the most numerous, especially on the aboral side. Tridentate pedicellariae scarce and found only on the oral side of the test, near or a short distance from the peristome and periproct. Globiferous pedicellariae found only near the ambitus, and rarely on the apical system. Triphyllous pedicellariae found only rarely; when present, they occur mostly in the ambulacra. Colour. From pale brownish cream to deep red, generally light red, with red tube feet (H. L. Clark 1921). According to Hendler et al. (1995), the tube feet vary from conspicuously brown to light brown and the spines are whitish to light brown. Naked test white or yellowish (Fig. 9 A – D).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E29FFF6FF551EDFFBC8F9D4.taxon	distribution	Distribution. Circumtropical (Pawson et al. 2009). In the Western Atlantic from Mexico, Florida, Bahamas, Bermudas, Cuba, Jamaica, Puerto Rico, Haiti, Dominican Republic, Belize, Honduras, Bonaire, Costa Rica, Panama, Colombia, Aruba, Venezuela, and Brazil (Engel 1939; H. L. Clark 1925; Bravo-Tzompatzi et al. 1999; Borrero-Pérez et al. 2002; Alvarado 2011; Rodríguez-Barreras et al. 2012; Solís-Marín et al. 2013). In Brazil, from Bahia (Magalhães et al. 2005). In this study, we provide the first record for the State of Ceará. From 1.5 to 570 m (del Valle García et al. 2005), but most commonly occurs at shallow depths (Hendler et al. 1995).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E29FFF6FF551EDFFBC8F9D4.taxon	discussion	Remarks. Echinoneus cyclostomus is the only extant species known in the genus. In the present study, both young and adult specimens were observed. Only small variations were recorded. In young individuals, the peristome is three to four times larger than the periproct. In adults this proportion is inverted. Young specimens have proportionally large tubercles relative to the corona. In adults, these tubercles are short. Westergren (1911) presented a broad morphological study for E. cyclostomus, illustrating in detail external and internal morphological characters and comparing specimens of different sizes from regions in the Pacific and Caribbean. Ecological notes. This small urchin is usually found under rock slabs or fragments of coral rubble, sandy bottoms, seagrass beds, and muddy substrates (Hendler et al. 1995; Solís-Marín et al. 2013), or on coral reefs and isolated rocks over sandy substrates with abundant vegetation, mainly Thalassia testudinum (del Valle García et al. 2005). Echinoneus cyclostomus is a detritus-feeder, selecting food with the tube feet and spines, primarily ingesting Foraminifera and algal fragments (Rose 1976; De Ridder & Lawrence 1982). Echinoneus cyclostomus also ingests organic material that adheres to coarse sand and shell fragments (Hendler et al. 1995).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E28FFF1FF551CBAFD13F985.taxon	description	Figure 5 H – I	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E28FFF1FF551CBAFD13F985.taxon	description	& Sovierzoski, 2013: 680.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E28FFF1FF551CBAFD13F985.taxon	materials_examined	Material examined. Photographic record.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E28FFF1FF551CBAFD13F985.taxon	description	Description (modified from Souto et al. 2011). Test small, wide and low, covered by short primary and secondary spines (Fig. 5 H). Aboral surface inflated, with greatest height posterior to apical system, on small elevation above periproct. Apical system anterior and monobasal. Petaloid region occupying 1 / 3 of aboral region (Fig. 5 H). Petals narrow, short, almost equal in length, and opened distally (Fig. 5 H). Petal I slightly shorter than petal V. Poriferous zones almost equal in length. Interambulacral plates wide at ambitus and narrowing towards the apical disc. Four types of primary spines cover test. Aboral spines short, thick, distally serrated, with enlarged tip, sometimes club-shaped. Spines at bourrelets thick, serrated, slightly curved and with enlarged tip. Oral spines near ambitus are long, straight, smooth proximally, distally serrated and pointing towards centre line of test, partially covering naked zone. Spines above periproct long and straight, covering whole depression, and proximally serrated. Miliary spines throughout test small, serrated, widening to small crown distally. Tubercles slightly crenulate and perforate. Eight large periproctal plates and four to seven small periproctal plates. Oral surface concave, especially along midline. Phyllodes single-pored, parallel and enlarged near peristome, with 6 – 8 pores in each row. Peristome slightly wider than long and located in depression on oral region of the test, displaced towards anterior edge (Fig. 5 I). Buccal spines long, obscuring peristomial opening. Pedicellariae. Ophicephalous pedicellariae sparsely distributed on the aboral surface, but numerous near the ambitus. Tridentate pedicellariae only around the periproct and on periproctal plates (Souto et al. 2011). Colour. Information not available for living specimens. Whitish in ethanol.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E28FFF1FF551CBAFD13F985.taxon	distribution	Distribution. Endemic to Brazil, known from Bahia and Sergipe (R. B. Moura, personal observation). From depths of 3.5 to 11 m (Manso et al. 2008; Souto et al. 2011).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E28FFF1FF551CBAFD13F985.taxon	discussion	Remarks. Four extant Cassidulus species are known, two of which are recorded from Brazil: C. infidus Mortensen, 1948 [from Bahia and Sergipe] and C. mitis Krau, 1954 [endemic to Rio de Janeiro]. Krau (1960) described C. delectus from Baía de Sepetiba (RJ), later synonimzed with C. mitis by Tommasi & Lima-Verde (1970). Mortensen (1948 b) described C. infidus on the basis of a single naked test. Mooi (1990) summarized data, and provided a key on living cassiduloids, including both C. mitis and C. infidus. Twenty years later, Souto et al. (2011), redescribed C. infidus, illustrated several morphological characters, and presented a modified identification key for the species of the genus provided by Mooi (1990). Cassidulus infidus is apparently rare. On the other hand, Souto et al. (2011) believed that the difficulty in finding this species may be related to the fact that they are very small, fragile, and live buried in sand. Ecological notes. This species lives in shallow and warmer waters, buried in medium-grained sand (Manso et al. 2008; Souto et al. 2011). According to Telford & Mooi (1996), Cassidulus species are not selective deposit feeders and their juveniles inhabit exactly the same sediment as the adults, feeding on the same array of particle sizes. Souto et al. (2011) found sediment and Foraminifera in the gut contents of one specimen of C. infidus. Specimens measuring 10 mm in test length already have completely developed genital pores (Tommasi & Lima- Verde 1970). As in C. caribaearum Lamarck, 1801 and C. mitis, C. infidus broods its young among the spines of its aboral surface (Tommasi & Lima-Verde 1970; Gladfelter 1978; Souto et al. 2011). According to Souto et al. (2011), C. mitis may have gregarious habits.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E2FFFF0FF551C69FA13F85B.taxon	description	Figure 10 A – H	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E2FFFF0FF551C69FA13F85B.taxon	materials_examined	Material examined. Bahia: 1 spm, Salvador, 14. IX. 1997 [UFBA 00072]; 1 spm, 12 ° 54 ′ S 28 ° 29 ′ W, Salvador, 22. VIII. 1993 [UFBA 00461].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E2FFFF0FF551C69FA13F85B.taxon	description	Description. Test large (TL = 139 mm; TW = 118 mm), oval, with rounded margin, uniformly covered by short, thin spines, petaloid region slightly inflated (Fig. 10 A – C). Apical system central, monobasal, with five gonopores (Fig. 10 F). Petaloid large, slightly longer than 50 % of test (Fig. 10 A, D). Petals large, leaf-shaped (Fig. 10 A, D). Paired petals nearly closed distally (Fig. 10 A, D). Lunule lacking. Aboral primary spines long, slightly club-shaped. Oral primary spines long, terminating in a small hyaline point. Tubercles perforate. Areoles of aboral primary tubercles uniform in size. Oral surface flat, slightly concave around peristome (Fig. 10 B, E). Peristome circular, central (Fig. 10 B, E, G). Food grooves unbranched, fairly deep, not reaching ambitus (Fig. 10 E). Periproct oral, towards posterior margin, located between third and fifth pairs of post-basicoronal interambulacral plates (Fig. 10 B, E). Periproctal membrane densely covered by spines, with some small bivalved triphyllous pedicellariae. Pedicellariae. Tridentate pedicellariae more abundant in areas close to peristome and to food grooves. Ophicephalous pedicellariae more abundant on margins and triphyllous pedicellariae distributed over entire test. Tridentate pedicellariae with short stalk and neck. Valves long, narrow, with denticulate margin and slightly enlarged tip. Ophicephalous pedicellariae short, without a neck. Valves narrow, with enlarged tips, and foramen (sensu Coppard et al. 2012) denticulate (Fig. 10 H). Bivalved triphyllous pedicellariae small, with long neck and small head. Valves rounded. Some quadridentate pedicellariae were also observed. Colour. Colour in life yellowish, reddish, very deep brown (Mortensen 1948 a), or olive green (Tommasi 1966 a). Spines brownish-green. Preserved specimens are light brownish (Mortensen 1948 a). Naked test white.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E2FFFF0FF551C69FA13F85B.taxon	distribution	Distribution. Florida, North Carolina, Mexico, Cuba, Jamaica, Dominican Republic, Panama, Colombia, Venezuela, and Brazil (Alvarado 2011; Tommasi 1966 a). In Brazil from BA, RJ, SP and SC (Tommasi 1966 b; Alves & Cerqueira 2000; Magalhães et al. 2005; Xavier 2010). From 5 to 210 m, more common between 5 and 50 m (Serafy 1979; Hendler et al. 1995; Laguarda-Figueras et al. 2005 a).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E2FFFF0FF551C69FA13F85B.taxon	discussion	Remarks. Presently 49 extant species of Clypeaster are known, making it the most speciose extant echinoid genus (Mihaljević et al. 2011). In Brazil, there are records of five species [C. rosaceus (Linnaeus, 1758), C. subdepressus, C. lamprus H. L. Clark, 1914, C. aloysioi (Brito, 1959) and C. oliverai Krau, 1952]. Both C. aloysioi and C. oliverai are endemic to southeastern Brazil. Clypeaster aloysioi differs from C. subdepressus in its pentagonal test, and narrower and more widely open petals. Clypeaster oliverai differs by having a large petaloid occupying almost the entire aboral surface, and slightly conical test in lateral view. Clypeaster lamprus differs by having a brown-reddish colour, a slightly pentagonal test with thick borders, and spatulate oral spines. Mortensen (1948 a) subdivided Clypeaster into ten subgenera. In a study on the architecture of the test Mihaljević et al. (2011) suggested that the subgenera proposed by Mortensen (1948 a) do not represent clades. Thus, Mihaljević et al. (2011) agree with Serafy (1970), Mooi (1989) and Kroh & Mooi (2016) in not adopting any subgenera, although recognizing considerable heterogeneity among the species. In the present study, only two adults were examined, in which no morphological variations were observed. Tommasi (1964) gave a good discussion on the possible morphological variations found in C. subdepressus. Ecological notes. This species lives in sandy substrates or sediments formed by shell fragments. It can be found in seagrass and on sandy bottoms near coral reefs (del Valle García et al. 2005). Telford et al. (1978) studied the feeding activities of C. subdepressus and established that individuals select particles above 400 µm in diameter for ingestion. According to Gladfelter (1978), the gastropod Cassis tuberosa is a common predator of this species. Clypeaster subdepressus is common along the littoral region of southeastern Brazil, but appears to be rare in northeastern Brazil.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E11FFCEFF551A15FF7CFCC1.taxon	description	Figure 11 A – I	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E11FFCEFF551A15FF7CFCC1.taxon	materials_examined	Material examined. Ceará: 2 spms, Canopus Bank [UFPB / ECH. 1974].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E11FFCEFF551A15FF7CFCC1.taxon	description	Description. Test oval or round (TL = 6.6 to 9.8 mm; TH = 2.6 to 8 mm), usually low and flat, oral region slightly concave (Fig. 11 A – D). Test covered by short, slender, hyaline spines (Fig. 11 A). Petals small and open distally (Fig. 11 C, E). Madreporite slightly prominent. Genital and ocular pores of same size (Fig. 11 C, E). Four genital and five ocular pores form a conspicuous circle or pentagon around the madreporite (Fig. 11 C, E). Primary spines elongate and lanceolate. Miliary spines shorter than primary spines, with terminal crown. Peristome central, circular, three times size of periproct (Fig. 11 B, D). Periproct small, circular, inframarginal, and usually covered by 5 to 6 regular plates that never bear spines. Phyllodes lacking. Pedicellariae. According to Mortensen (1907), there are three types of pedicellariae (ophicephalous, tridentate and triphyllous) (Fig. 11 F – H). In the present study, we have observed only two types, ophicephalous and triphyllous ones. The absence of tridentate pedicellariae is probably the result of their loss by inadequate fixation or conservation. Triphyllous and ophicephalous pedicellariae over entire test. Ophicephalous pedicellariae with long stalk and head, without a neck. Valves with margin denticulate. Colour. According to Tommasi (1966 b), living specimens are brown-yellow. White, cream-coloured, greenish or gray (naked test) in alcohol (Fig. 11 A, D).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E11FFCEFF551A15FF7CFCC1.taxon	distribution	Distribution. Mexico, Gulf of Mexico, Florida, Bahamas, Antilles, Cuba, Jamaica, Colombia, and Brazil (Borrero-Pérez et al. 2002; Alvarado 2011, Solís-Marín et al. 2013; Martínez-Melo et al. 2015). In Brazil from MA, PB, AL, BA, RJ, and SP (Tommasi 1966 a; Kempf 1972; Oliveira et al. 2010; Smithsonian Database). In this study, we record the species for the first time from the coast of Ceará. From 110 to 2500 m (Pawson et al. 2009).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E11FFCEFF551A15FF7CFCC1.taxon	discussion	Remarks. Presently, 15 – 20 recent species of Echinocyamus are known (Markello 2015), of which only three occur in the Atlantic [E. pusillus (Müller, 1776), E. grandiporus and E. macrostomus Mortensen, 1907]. Echinocyamus grandiporus differs from E. pusillus, which has larger petals, a more rounded test in lateral view, and ophicephalous pedicellariae with a more serrate blade. Echinocyamus macrostomus differs in having ocular and genital pores of different sizes, test more elongated, and large peristoma. Mortensen (1907) provided an excellent discussion regarding morphological differences among E. grandiporus, E. pusillus and E. macrostomus. Echinocyamus grandiporus is a small echinoid which rarely exceeds 20 mm TL (Borrero-Pérez et al. 2002). However, specimens larger than 15 mm TL are sometimes seen (A. Kroh, personal communication). In this study, the largest specimen is 9.17 mm TL. Ecological notes. Found in muddy and sandy bottoms. Tommasi (1966 b) recorded this species between 110 and 130 m on bottoms with deep-sea scleractinian corals of the genera Deltocyathus Milne Edwards & Haime, 1848, Trochocyathus Milne Edwards & Haime, 1848 and Cladocora Ehrenberg, 1834, together with many brachiopods, ophiuroids, and gastropods. Kroh & Mooi (2016) cite the presence of the species on submarine seamounts. Almost nothing is known about its biology and ecology, probably due to its occurrence in deeper waters.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E10FFC8FF551F69FB05FDA6.taxon	description	Figures 12 A – J, 18 F, 19 F, G	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E10FFC8FF551F69FB05FDA6.taxon	materials_examined	Material examined. Ceará: 3 spms, Ponta Grossa, Aracati, 24. VI. 1982 [UFPB / ECH. 1231]; 1 spm, Arpoeira Beach, Aracaraú, 07. VIII. 1982 [UFPB / ECH. 1232]. Rio Grande do Norte: 5 spms, Timbau Beach, 29. II. 1980 [UFPB / ECH. 1205]; 3 spms, Diogo Lopes, Macau, 11. XI. 2007 [UFPB / ECH. 1228]; 1 spm, Diogo Lopes, Macau, 12. XI. 2007 [UFPB / ECH. 1314]; 7 spms, Ponta Santo Cristo, to the east of São Miguel do Gostoso, 21. VI. 1982 [UFPB / ECH. 1894]; 1 spm, Ponta do Tubarão, criffs in front of Chico Martins, Macau, 02. II. 2011 [UFPB / ECH. 1929]; 7 spms, river mouth of Rio Tubarão, Diogo Lopes, Macau, 15. XI. 2009 [UFPB / ECH. 1930]. Paraíba: 12 spms, Baía da Traição, Baía da Traição, 04. VI. 1979 [UFPB / ECH. 350]; 1 spm, Santa Catarina Beach, Cabedelo, 18. V. 2007 [UFPB / ECH. 616]; 13 spms, Baía da Traição, Baía da Traição, 24. XI. 2007 [UFPB / ECH. 1015]; 6 spms, Baía da Traição, 04. V. 2008 [UFPB / ECH. 1167]; 11 spms, Bessa Beach, João Pessoa, 18. II. 1979 [UFPB / ECH. 1206]; 2 spms, Carapibus Beach, Conde, 21. III. 2008 [UFPB / ECH. 1207]; 3 spms, Carapibus Beach, Conde, 08. III. 2008 [UFPB / ECH. 1208]; 1 spm, Cabo Branco Beach, João Pessoa, 05. XII. 1 983 [UFPB / ECH. 1213]; 1 spm, Lucena Beach, Lucena, 07. X. 2007 [UFPB / ECH. 1227]; 21 spms, Cabo Branco Beach, João Pessoa, 18. VIII. 1978 [UFPB / ECH. 1229]; 2 spms, Lucena Beach, Lucena, 07. X. 2007 [UFPB / ECH. 1230]; 1 spm, Coqueirinho Beach, Conde, 03. VI. 2008 [UFPB / ECH. 1285]; 37 spms, Ponta de Campina, Cabedelo, 19. XI. 1983 [UFPB / ECH. 1292]; 1 spm, Bessa Beach, João Pessoa, 18. XII. 2007 [UFPB / ECH. 1293]; 2 spms, Costinha Beach, Lucena, 07. X. 2007 [UFPB / ECH. 1294]; 1 spm, Cabo Branco Beach, João Pessoa, 25. X. 2007 [UFPB / ECH. 1295]; 2 spms, Barra de Mamanguape Reefs, Rio Tinto, 10. I. 2009 [UFPB / ECH. 1458]; 2 spms, Maceiozinho Beach, Jacumã, Conde, 06. V. 2008 [UFPB / ECH. 1474]; 11 spms, Santa Catarina Beach, Cabedelo, 05. V. 2009 [UFPB / ECH. 1931]; 4 spms, Cabo Branco Beach, João Pessoa, 09. II. 2001 [UFPB / ECH. 1942]; 1 spm, Barra de Mamanguape, Rio Tinto, 13. I. 2017 [UFPB / ECH. 2287]. Alagoas: 1 spm, Pontal do Peba, Piaçabuçu, 27. I. 1983 [UFPB / ECH. 1233]; 8 spms, Japaratinga Beach, Japaratinga, 18. IX. 2009 [UFPB / ECH. 1892]. Sergipe: 1 spm, Mosqueiro Beach, Aracajú, 25. IV. 2010 [UFPB / ECH. 1920]; 1 spm, Sarney Beach, Aracajú, 30. VIII. 1995 [UFSITAB- 6]; 1 spm, Sarney Beach, Aracaju, 30. VII. 1995 [UFSITAB- 117]; 1 spm, Aruana Beach, Aracajú, 21. I. 2007 [UFSITAB- 120]. Bahia: 3 spms, Iemanjá Beach, Salvador, 10. I. 2007 [UFSITAB- 116].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E10FFC8FF551F69FB05FDA6.taxon	description	Description. Test wider than long (TL = 71.7 mm; TW = 78.7 mm), with thin ambitus, domed anteriorly, highest point of test located slightly anterior to apical system and curving down abruptly to anterior margin (Fig. 12 A, C). Apical system monobasal, with four gonopores (Fig. 12 I). Petaloid large, corresponding to a little more than 50 % TL (Fig. 12 A, C). Posterior petals slightly longer than anterior ones (Fig. 12 A, C). Five lunules, anal lunule longer than paired ambulacral lunules (Fig. 12 A – D). Aboral primary spines short and club-shaped, covering the test uniformly (Fig. 12 A, H). Each Miliary spines each with sac-like structure on tip, densely scattered among primary spines. Lunules with two types of spines: aboral ones longer, flat and enlarged at tip (similar to swim fins). Inner spines elongate and tapering. Oral surface flat, with long, thin locomotory spines that are slightly curved at the base, occupying the centre of interambulacra (Fig. 12 b, D). Geniculate spines short and slightly curved, in fields from margins of interambulacra to margins of food grooves. Among these are long, slender spines in areas near pressure drainage channels (Fig. 12 B). Peristome pentagonal and slightly anterior (Fig. 12 B, D). Periproct oval to triangular, very close to peristome, deeply indenting basicoronal plate (Fig. 12 B, D). Periproctal membrane covered by large plates of different sizes and shapes. Interambulacral basicoronal plates longer than ambulacral ones. Food grooves bifurcating at distal edges of ambulacral basicoronals and highly branched distally (Fig. 12 D). Pedicellariae. Pedicellariae scattered over entire test, being more abundant in oral interambulacra and in areas close to lunules, peristome, and periproct. Bidentate pedicellariae with a long neck (as long as or shorter than stalk) and short head. Valves short, with narrow and slightly curved, median region enlarged, margin slightly serrate, and a terminal tooth that crosses with the tooth of the opposing valve (this tooth is seen only in the larger pedicellariae) (Fig. 12 E – G). Colour. Greenish or brownish (Fig. 12 A, B, 19 F). Naked test white (Fig. 12 C, D).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E10FFC8FF551F69FB05FDA6.taxon	distribution	Distribution. In Brazil from MA, PI, CE, PB, PE, AL, SE, BA, ES, RJ, SP, SC, and RS (Brito 1962; Tommasi 1964, 1966 a, 1974; Lima-Verde 1969; Harold & Telford 1990; Fernandes et al. 2002; Magalhães et al. 2005; Martins & Martins de Queiroz 2006; Gondim et al. 2008; Lima & Fernandes 2009; Gondim & Giacometti 2010; Oliveira et al. 2010; Xavier 2010). In this study, we provide the first record of the species for the State of Rio Grande do Norte. From depths between 0 and 550 m, being more common between 1 and 3 m (Tommasi 1974).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E10FFC8FF551F69FB05FDA6.taxon	discussion	Remarks. Seven extant species of Mellita are known, of which only M. quinquiesperforata had been recorded for Brazil. The taxonomic history of this species is quite complex (Harold & Telford 1990). Among several subspecies, M. quinquiesperforata latiambulacra was cited for Brazil on the basis of the diagnostic character, a more rounded test (Tommasi 1964). Harold & Telford (1990) conducted a morphological analysis of the genus and established the following diagnostic characters for M. quinquiesperforata: sharply inclined anterior profile; position of maximum thickness usually distinctly anterior to centre; test very broad; unique spatulate spines bordering the lunules. Recently, Coppard et al. (2013) conducted a phylogeographic study of the genus Mellita along the coasts of the Americas and concluded that the specimens from Brazil (collected along the littoral of the State of Paraíba, northeast Brazil), represented a new species, yet unnamed and not described. According to these authors, the new species of Mellita is distributed from the Lesser Antilles to tropical Brazil. Herein we provisionally considered our specimens from the northeastern Brazil as M. aff. quinquesperforata, based on the results by Coppard et al. (2013), its morphological similarities with M. quinquiesperforata and recommenations of the ICZN (1999). We only examined adults, in which no significant morphological variations were observed. Mellita aff. quinquiesperforata has four gonopores. Tommasi (1966 a) used this feature as a character to distinguish Encope (five gonopores) and Mellita. Tommasi (1957) examined a single specimen of Mellita with five gonopores, which he considered an anomaly. Ecological notes. This species lives in sandy substrates. On beaches, it occurs mainly near the surf zone. Mellita aff. quinquiesperforata is the most common clypeasteroid along the northeastern coast of Brazil and may form dense populations just behind the surf zone (Borzone 1992). In the present study, we observed high densities of this species in hypersaline estuaries, where it occurred together with Leodia sexiesperforata and Encope emarginata. Several specimens harboured one or two small crabs on the oral surface, possibly of the genus Dissodactylus Smith, 1870. Along the semiarid littoral of the State of Rio Grande do Norte, M. aff. quinquiesperforata is one of the main preys of the gastropod Cassis tuberosa (Mota 2012).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E16FFCAFF551D4AFBB7FE81.taxon	description	Figures 13 A – H, 18 G Echinodiscus sexiesperforatus Leske, 1778: 199.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E16FFCAFF551D4AFBB7FE81.taxon	materials_examined	Material examined. Rio Grande do Norte: 3 spms, Ponta do Rio Tubarão, in front of the cliffs of Chico Martins, Diogo Lopes, Macau, 02. II. 2011 [UFPB / ECH. 1893]; 5 spms, Ponta do Rio Tubarão, in front of the cliffs of Chico Martins, Diogo Lopes, 01. II. 2011 [UFPB / ECH. 1915]. Paraíba: 1 spm, Baía da Traição Beach, Baía da Traição, 24. XI. 2007 [UFPB / ECH. 1013]; 1 spm, Baía da Traição Beach, Baía da Traição, 05. V. 2008 [UFPB / ECH. 1165]; 1 spm, 6 ° 56.139 ′ S 34 ° 49.391 ′ W, Porto Project, Cabedelo, 09. II. 2004 [UFPB / ECH. 1297]; 1 spm, Ponta de Campina, Cabedelo, 20. XI. 1983 [UFPB / ECH. 1303]; 1 spm, reefs facing Ponta de Campina, Poço Beach, Cabedelo, 28. II. 2010 [UFPB / ECH. 1921]. Alagoas: 1 spm, Francês Beach, Marechal Deodoro, 29. I. 1983 [UFPB / ECH. 1307]. Bahia: 1 spm, Ponta da Coroa Vermelha, Santa Cruz da Cabrália, 14. X. 1982 [UFPB / ECH. 1302].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E16FFCAFF551D4AFBB7FE81.taxon	description	Description. Test slightly pentagonal in outline (TL = 84 mm; TW = 79 mm), discoidal, fragile, with slight elevation in centre, ambitus extremely thin and sharp (Fig. 13 A – D). Apical system monobasal (Fig. 13 H), four gonopores (Fig. 13 H). Petaloid small, occupying only central region of test (Fig. 13 C, H). Posterior petals slightly longer than anterior ones (Fig. 13 A, C). Six long, slit-like lunules (Fig. 13 A – C), anal lunule shorter than five ambulacral ones (Fig. 13 A, C). Aboral primary spines strongly club-shaped, uniformly covering test (Fig. 13 A). Miliary spines with sac-like structure on tip, densely scattered among primary spines (Fig. 13 G). Lunules with two types of spines, on aboral margin, spines long, flat and slightly curved. Internally, spines are shorter, slenderer than aboral spines, but also slightly curved. Oral surface flat, with locomotory spines long, slender, and slightly curved at base, being largest spines in this region (Fig. 13 B). Short and slender spines occur among these spines. Spines on ambitus similar to those around lunules. Peristome circular and central (Fig. 13 B, D, E). Periproct small, elongate, close to peristome, slightly indenting basicoronal plate (Fig. 13 B, D). Basicoronal plates reduced in size. Ambulacral basicoronal plates almost triangular. Pressure drainage channels and food grooves well-developed (Fig. 13 D). Food grooves bifurcating after ambulacral basicoronal plates, surrounding lunules and reaching margin of test (Fig. 13 D). Pedicellariae. Pedicellariae over entire test, more abundant in oral inter and near the lunules, peristome, and periproct. Bidentate pedicellariae with a long neck (of same size or slightly smaller than the stalk) and short head. Valves short, with narrow and slightly curved base, median region enlarged with denticulate margin and two end teeth that cross with the teeth of opposing valve (Fig. 13 F). Colour. From yellow to brown or greenish (Hendler et al. 1995). Naked test white.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E16FFCAFF551D4AFBB7FE81.taxon	distribution	Distribution. North Carolina, Bahamas, Florida, Gulf of Mexico, Cuba, Belize, Honduras, Puerto Rico, Costa Rica, Jamaica, Panama, Venezuela, Colombia, Brazil, and Uruguay (Hendler et al. 1995; Alvarado et al. 2008, del Valle García et al. 2008; Francisco & Pauls 2008; Rodríguez-Barreras 2014). In Brazil from CE, RN, PB, PE, AL, BA, RJ, SP, including Trindade Islands (Rathbun 1879; Brito 1962, 1968; Tommasi 1966 b; Lima-Verde 1969; Magalhães et al. 2005; Gondim et al. 2008; Manso et al. 2008; Martins et al. 2018). From depths of 0 to 30 m (Francisco & Pauls 2008).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E16FFCAFF551D4AFBB7FE81.taxon	discussion	Remarks. The genus Leodia is represented by a single extant species. The status of this genus has been the subject of some dispute (Mooi & Peterson 2000), having been for many years considered to be a synonym or a subgenus of Mellita. Among the features that were used to separate these genera were the process of development and the number of the lunules (Lambert & Thiéry 1921; H. L. Clark 1940 a; Mortensen 1948 a). However, Durham (1955) pointed out some important differences between Leodia and Mellita that did not rely on lunule number. A series of authors recognized the distinction of these genera (e. g. Durham 1955; Serafy 1979; Mooi 1989; Hendler et al. 1995; Coppard 2016) and phylogenetic studies support this view (Mooi & Peterson 2000; Coppard et al. 2013). Leodia sexiesperforata is distinguished from the species of Mellita by the slightly pentagonal outline of the strongly flattened test, and very narrow lunules. Kier (1963) and Mooi & Peterson (2000) included other characters that are also used for distinguishing L. sexiesperforata from living representatives of Mellita, for example: five ambulacral lunules, posterior lunule not extending far anteriorly between posterior petals, and interambulacral basicoronal plates separated from first interambulacral post-basicoronals by two plates of each adjoining ambulacrum. According to Mooi & Peterson (2000), extant Leodia have aboral primary spines with smooth, very strongly club-shaped tips that are more markedly bent and swollen than those found in any other living mellitid. Ecological notes. This species lives in sandy areas with little or no vegetation (Hendler et al. 1995). It is usually uncommon along the northeastern littoral, yet some sandy beaches may contain significant population densities of this species, as observed in Macau (State of Rio Grande do Norte) and Lucena (State of Paraíba). In the present study, the species was abundant in hypersaline mangroves (State of Rio Grande do Norte), and co-occurred with M. aff. quinquiesperforata and E. emarginata. In many examined specimens we encountered one or two small crustaceans, possibly of the genus Dissodactylus Smith, 1870, associated with the oral surface of the sand-dollar. We further observed small bivalves living inside one of the lunules. McClintock & Marion (1993) recorded the mollusk gastropod Cassis tuberosa as one of the main predators of L. sexiesperforata in the Bahamas. Goodbody (1960) and Telford & Mooi (1986) summarized data on feeding habits of Leodia.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E14FFC4FF551ECAFCADFC86.taxon	description	Figures 14 A – I, 18 H, 19 E Echinodiscus emarginatus Leske, 1778: 136, figs 5 – 6.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E14FFC4FF551ECAFCADFC86.taxon	materials_examined	Material examined. Rio Grande do Norte: 1 spm, Diogo Lopes, Macau, 11. XI. 2007 [UFPB / ECH. 1299]; 2 spms, Ponta do Tubarão, Macau, 02. II. 2011 [UFPB / ECH. 1908]. Paraíba: 3 spms, Ponta de Campina, Cabedelo, 20. XI. 1983 [UFPB / ECH. 615]; 2 spms, 7 ° 8 ′ 12,182 ″ S 34 ° 48 ′ 11,817 ″ W, Project SEMAN, 04. X. 2007 [UFPB / ECH. 865]; 1 spm, Areia Vermelha Reef, Cabedelo, 06. IV. 2008 [UFPB / ECH. 1214]; 2 smps, 6 ° 56,139 ′ S 34 ° 49,391 ′ W, Project Porto, Cabedelo, 09. II. 2004 [UFPB / ECH. 1298]; 1 spm, Ponta de Campina, Cabedelo, 20. XI. 1983 [UFPB / ECH. 1303]; 3 spms, Lucena Beach, Lucena, 07. X. 2007 [UFPB / ECH. 1305]; 18 spms, Ponta de Campina, Cabedelo, 20. XI. 1983 [UFPB / ECH. 1308]; 1 spm, Cabo Branco Beach, João Pessoa, 23. XII. 2007 [UFPB / ECH. 1309]; 5 spms, Costinha Beach, Lucena, 07. X. 2007 [UFPB / ECH. 1312]; 3 spms, receife em frente a Ponta de Campina, Cabedelo, 28. II. 2010 [UFPB / ECH. 1907]; 15 spms, Barra de Mamanguape, Rio Tinto, 13. I. 2017 [UFPB / ECH. 2286]. Pernambuco: 3 spms, Barra de Catuama, Goiana, 0 1.03.2008 [UFPB / ECH. 1304]; 7 spms, Barra de Catuama, Goiana, 01. III. 2008 [UFPB / ECH. 1311]; 2 spms, Cayana, Goiana, 31. X. 1982 [UFPB / ECH. 1313]. Alagoas: 3 spms, Pontal do Peba, Piaçabuçu, 27. I. 1983 [UFPB / ECH. 1291]; 1 spm, Paripoeira Beach, Maceió, 01. II. 1983 [UFPB / ECH. 1300]; 1 spm, Pontal do Peba, Piaçabuçu, 27. I. 1983 [UFPB / ECH. 1306]; 1 spm, Ipioca Beach, Maceió, 30. VI. 2007 [UFSITAB- 121]; 6 spms, Ipioca Beach, Maceió, 30. VI. 2007 [UFSITAB- 127]. Bahia: 5 spms, Ponta da Coroa Vermelha, Santa Cruz da Cabrália, 14. X. 1982 [UFPB / ECH. 1301]; 3 spms, Coroa Vermelha, Santa Cruz da Cabrália, 29. XII. 1984 [UFPB / ECH. 1310]; 1 spm, Coroa Vermelha, Santa Cruz da Cabrália, 29. XII. 1984 [UFPB / ECH. 1360].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E14FFC4FF551ECAFCADFC86.taxon	description	Description. Test oval, domed centrally, with moderately thickened margin (TL = 119.2 mm; TW = 120.1 mm) (Fig. 14 A – D). Apical system monobasal, with five gonopores (Fig. 14 I). Petaloid large (Fig. 14 A, C). Petals bowed and distally open (Fig. 14 C). Posterior petals slightly longer than anterior ones (Fig. 14 C). Six short, broad and drop-shaped lunules, larger in the anterior portion of the body (Fig. 14 A – D). Anal lunule extends between posterior petals (Fig. 14 A – D). Aboral primary spines short and club-shaped, uniformly covering test. Miliary spines with sac-like structure on tip, densely scattered among primary spines. Lunules with two types of spines; long spines, narrow at base and with broad tip (like swim fins) on aboral margins, and long, uniformly thin spines internally. Oral surface flat (Fig. 14 B, D), with long and thin locomotory spines that are slightly curved at base, and short and slightly curved secondary spines. Peristome pentagonal, slightly anterior (Fig. 14 B, D). Periproct oval and anterior to anal lunule (Fig. 14 B, D), not in contact with basicoronal. Periproctal membrane covered by plates of variable shapes and sizes, largest occurring on distal margin of periproct. Basicoronal plate small. Five food grooves bifurcating at edge of basicoronal plates, each branch surrounding the lunules (Fig. 14 D). Pedicellariae. Pedicellariae over entire test, but more abundant on oral surface and near the lunules, peristome, and periproct. Bidentate pedicellariae with long neck and short head. Valves short, with narrow and slightly curved bases, median region enlarged with serrated margins and one end tooth, which crosses terminal tooth of opposing valve (Fig. 14 E – H). Colour. Brown or greenish (Fig. 14 A, B, 18 H). Naked test white (Fig. 14 C, D).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E14FFC4FF551ECAFCADFC86.taxon	distribution	Distribution. Florida, Belize, Guatemala, Panama, Caribbean Sea, Colombia, Venezuela, Brazil, and Argentina (mouth of La Plata River) (H. L. Clark 1925; Tommasi 1966 a; Francisco & Pauls 2008; Alvarado 2011; Kroh & Mooi 2016). In Brazil from AP, MA, PI, CE, PB, PE, AL, BA, RJ, SP, PR, SC and RS (Bernasconi 1955; Brito 1962; Tommasi 1964, 1972; Lima-Verde 1969; Reichholf 1981; Manso 1989; Fernandes et al. 2002; Magalhães et al. 2005; Manso et al. 2008; Lima & Fernandes 2009; Gondim & Giacometti 2010; Oliveira et al. 2010; Gondim et al. 2011, 2013 b; Lopes 2011). In this study, we provide the first record of this species for Rio Grande do Norte. From depths of 0 to 50 m (Tommasi 1966 a).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E14FFC4FF551ECAFCADFC86.taxon	discussion	Remarks. The neotropical genus Encope contains seven extant species, of which only E. emarginata occurs along the Brazilian littoral. Encope emarginata differs from E. michelini L. Agassiz, 1841, from the Gulf of Mexico, as the latter species has open lunules (notches). Young specimens of E. emarginata may be mistaken for E. michelini due to the presence of open lunules in the former when it is young (Francisco & Pauls 2008). In the present study, only adult individuals were examined, and no morphological variations were observed. Ventura et al. (2010) analyzed the presence of morphological variations among seven populations from northeastern and southeastern Brazil and found that the thickness and shape of the test and the position of the lunule relative to the madreporite vary among five of the populations. According to these authors, these variants may be related to different environmental conditions to which the populations are subject to. Tommasi (1964) discussed differences between young and adult individuals. Recently, Coppard & Lessios (2017) conducted a phylogeographic study of Encope, finding that the phylogeny of the genus is characterized by four notable features, among which we highlight extreme morphological plasticity in the widespread species that does not show geographic structure, and an unusually slow rate of molecular evolution. These authors also corroborate that E. oblonga L. Agassiz, 1841, E. subclausa L. Agassiz, 1841, and E. valenciennesii L. Agassiz, 1841 are synonyms of E. emarginata. Ecological notes. This species lives in sandy areas, mainly in the surf zone (Fernandes et al. 2002). According to Brito (1962), E. emarginata is very common in Porto Seguro (BA). It is relatively common along the entire northeastern coast. In the present study, the species was observed to be more numerous in areas of hypersaline mangroves (Rio Grande do Norte), where it occurred together with M. aff. quinquiesperforata and L. sexiesperforata. As for these accompanying species, one or two small crabs, possibly belonging to the genus Dissodactylus, were found associated with the oral surface of E. emarginata (Fig. 19 E). According to Tommasi (1964), young and adult specimens of E. emarginata live segregated into different depths, and the young tend to develop in shallow waters. As they grow, they migrate to deeper waters. Telford (1981) studied the morphology of sand dollars and its relation to hydrodynamic factors. He concluded that E. emarginata is a typically intertidal species with a body shape particularly adapted to its habitat.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1AFFC6FF551F61FBEFFE81.taxon	description	Figure 15 A – J	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1AFFC6FF551F61FBEFFE81.taxon	materials_examined	Material examined. Sergipe: 2 spms, Atalaia Beach, Aracaju [UFSITAB- 118]. Complementary material: 1 spm, São Sebastião Beach, São Paulo, X. 1955 [MZUSP, without voucher]; 1 spm, 22 ° 30 ′ S 41 ° 23 ′ W, Rio de Janeiro, 25. X. 1963 [EqMN 333].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1AFFC6FF551F61FBEFFE81.taxon	description	Description. Test subspherical or elliptical, covered with short, glassy and often curved spines (TL = 25 to 36.4 mm; TH = 19.3 to 31 mm; TW = 21 to 25.3 mm) (Fig. 15 A – C). Apical system ethmolytic. Two gonopores. Anterior ambulacra (II, III and IV) narrower and longer than posterior ambulacra (I and V) (Fig. 15 A, D). Ambulacrum III deeply sunken and broader than the others (Fig. 15 A, D). Other ambulacra also deeply sunken, forming slit-like depressions on aboral surface (Fig. 15 A, D). Peripetalous fasciole well-developed, bordering margin of petals (Fig. 15 A, D). Lateroanal fasciole well-developed, often appearing as black line (Fig. 15 F). Posterior region of test truncate (Fig. 15 A – F). Periproct oval and longitudinal (Fig. 15 G). Peristome kidney-shaped (Fig. 15 B, E), covered by six or seven large plates. Labrum expanded, almost covering peristome (Fig. 15 E). Amphisternous plastron covered by long and spatula-shaped spines (Fig. 15 B). Crenulate and perforate tubercles (Fig. 15 D – G). Pedicellariae. Pedicellariae not observed, as only naked tests were available. The following descriptions were obtained from the literature. Globiferous pedicellariae with long and slender valves, curving abruptly at the tip. Foramen surrounded by six small teeth. Tridentate pedicellariae with curved and slender valves (Tommasi 1958 a). Chesher (1963) provided excellent illustrations of this character (Fig. 15 H – J). Colour. Light brown or yellowish to white (Hendler et al. 1995). The spines, where more thickly clustered, are brownish (A. Agassiz 1869). Fasciole often dark red-brown, horseshoe-shaped, and partially encircling apical system (Hendler et al. 1995). Naked test white (Fig. 15 D – G).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1AFFC6FF551F61FBEFFE81.taxon	distribution	Distribution. North Carolina, South Carolina, Gulf of Mexico, Florida, Belize, Bermuda, Cuba, Jamaica, Puerto Rico, Dominican Republic, Guadalupe, Haiti, Honduras, Panama, Colombia, Venezuela, and Brazil (H. L. Clark 1925; Borrero-Pérez et al. 2002, 2012; Alvarado et al. 2008; Smithsonian Database). In Brazil from SE, RJ and SP (Tommasi 1958 a; Oliveira et al. 2010). From depths of 0 to 445 m, more common in less than 50 m depth (Serafy 1979; Laguarda-Figueras et al. 2005 b).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1AFFC6FF551F61FBEFFE81.taxon	discussion	Remarks. Five extant species of Moira are known, of which only M. atropos occurs in the Atlantic Ocean. According to Chesher (1963), adults may reach a maximum TL of 60 mm. In the present study, only adults were observed, lacking variations. Moira atropos is a very characteristic and easily identifiable species due to the strongly sunken petals. Ecological notes. This species lives buried up to 15 cm in mud or mud-sand bottoms (del Valle García et al. 2005; Chesher 1963). It is a detritus feeder, extending the tubefeet of ambulacrum III in the direction of the surface through a tunnel constructed with sediment (Chesher 1963). Moira atropos is frequently recorded together with Schizaster spp. and Brissopsis elongata Mortensen, 1907. Among its main predators are the asteroid Luidia clathrata (Say, 1825) and the margate fish Haemulon album Cuvier (Hendler et al. 1995). The species is rarely collected, probably due to its burrowing habit. According to Brito (1962), this is a relative common species in São Sebastião, Santos, and Cananéia (São Paulo). However, in northeastern Brazil it is apparently rare. According to Chesher (1963), M. atropos lacks a mechanism to carry genital products from the neighbourhood of the genital pores to the water overlying the burrow. Therefore, this heart urchin likely ascends to the surface at fairly frequent intervals and there releases any available ripe sperm or eggs (Moore & Lopez 1966). Moore & Lopez (1966) reported a significant correlation between spawning of M. atropos and lunar phases, with peak spawning occurring immediately after a full moon. According to Tommasi (1958 a) the genital pores only appear in specimens larger than 57 mm TL. However, we observed specimens of 24.76 mm with open genital pores.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E18FFC6FF551AEEFBA2F853.taxon	description	Figure 9 E – K	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E18FFC6FF551AEEFBA2F853.taxon	materials_examined	Material examined. Photographic record.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E18FFC6FF551AEEFBA2F853.taxon	description	Description (modified from Chesher 1966, Manso 2004 and Smith & Stockley 2005). Test small, fragile, subglobular, almost as broad as long, slightly larger than high, and vertically truncate posteriorly (Fig. 9 F – G). Test covered by slender, hyaline, and and often curved spines (Fig. 9 F – G), with slightly spatulate tips. Apical system ethmolytic, with four large gonopores. Anterior genital pores about half the size of posterior pores. Anterior paired petals (ambulacra II and IV) slightly curved and two to three times longer than posterior petals (ambulacra I and V) (Fig. 9 E). Ambulacrum III forms a strongly sunken furrow extending toward peristome (Fig. 9 E) that lacks primary spines. Peripetalous and lateroanal fascioles well-developed (Fig. 9 G). Peripetalous fasciole crosses ambulacrum III (Fig. 8 E), and anteriorly over plates 2. a. 4 and 2. b. 4 [4, 4 :: 5 + 6 / 7, 6 (7) :: 10]. Lateroanal fasciole begins close to ambulacra II and IV and extends towards of periproct [–, – :: 5, 4 (5) :: 5 + 4] (Fig. 9 E, G). Plates of ambulacra I and V without spines (Fig. 9 F). Peristome narrow, three times as broad as long (Fig. 9 F). Labrum extends to second plate of interambulacrum 5. Plastron amphisternous, covered by long, narrow spines, spatulate (Fig. 9 F). Periproct large, oval, located almost vertically. Pedicellariae. Globiferous and tridentate pedicellariae distributed over entire test. Rostrate pedicellariae found near periproct (Fig. 9 H). Globiferous pedicellariae with stalk, without a neck, valves with two end teeth (Fig. 9 J, K). Tridentate pedicellariae with valve having an enlarged and serrate tip (Fig. 9 I). Valve of rostrate pedicellariae with a narrow, serrate and spatulate blade (Chesher 1966). Chesher (1966) provides illustrations of all the pedicellariae of this species (Fig. 9 H – K). Colour. Test light pink to yellowish (Hendler et al. 1995). The fascioles and tube feet of petals red, tube feet of anterior groove yellow-orange (Hendler et al. 1995; Manso 2004).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E18FFC6FF551AEEFBA2F853.taxon	distribution	Distribution. Gulf of Mexico, Florida, Bahamas, Bermudas, Belize, Puerto Rico, Honduras, Dominican Republic, Panama, Venezuela, and Brazil (Hendler et al. 1995; Lessios 1998; Manso 2004; Alvarado 2011; Rodríguez-Barreras et al. 2013; Solís-Marín et al. 2013). In Brazil, from BA and RJ (Manso 2004; Oliveira et al. 2010). From depths of 2 to 45 m (Manso 2004).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E18FFC6FF551AEEFBA2F853.taxon	discussion	Remarks. The genus Schizaster contains nine extant species, of which only three are recorded from the tropical Atlantic [S. doederleini (Chesher, 1972), S. orbignyanus A. Agassiz, 1880 and S. floridiensis]. The first two range from the Gulf of Mexico to Venezuela and the latter has Rio de Janeiro as its southern limit. Schizaster floridiensis differs from S. doederleini by its rounded test, presence of less numerous pore-pairs in petal III, a shorter distance from the periproct to the lateroanal fasciole, and a larger petaloid. Schizaster floridiensis also differs from its congener S. orbignyanus, which has two genital pores and ambulacrum III more depressed and larger. According to Chesher (1966), details are lacking for several important taxonomic characters in the original description given by Kier & Grant (1965). Thus, Chesher (1966) redescribed the species and furnished an excellent discussion on variations found in S. floridiensis. Ecological notes. This species lives burrowed in sandy and muddy sediments, at depths of about 25 cm into the sediment (Chesher 1966). Schizaster floridiensis frequently occurs in association with S. doederleini and other fragile, burrowing spatangoids (Hendler et al. 1995). Manso (2004) recorded S. floridiensis together with the ophiuroid Amphioplus camamuensis Manso, 2004, from Baía de Camamu (Bahia).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1EFFC0FF551A2DFF74F84B.taxon	description	Figure 16 A – G	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1EFFC0FF551A2DFF74F84B.taxon	materials_examined	Material examined. Bahia: 1 spm, 12 ° 56 ′ 13 ″ S 38 ° 33 ′ 55 ″ W, 56 m [UFBA 00369]; 2 spms, Busca Vida Beach, Guarajuba, Camaçari, I. 2010, 37 m [UFBA 01195]; 1 spm, Busca Vida Beach, Guarajuba, Camaçari, II. 2008, 45 m [UFBA 01196]; 2 spms, Busca Vida Beach, Guarajuba, Camaçari, II. 2011, 37 m [UFBA 01304]; 1 spm, Baía de Camamu [UFSITAB- 124]; 1 spm, without locality data, 07. XII. 1978 [MZUSP, without voucher]; 1 spm, without locality data [MZUSP, without voucher].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1EFFC0FF551A2DFF74F84B.taxon	description	Description. Test small, oval or rounded, covered by long, slender, vitreous spines (TL = 7.4 to 15.3 mm; TH = 5.3 to 11 mm; TW = 6 to 13 mm) (Fig. 16 A – C). Apical system ethmolytic, displaced towards posterior region of test (Fig. 16 A, D, G). Four gonopores, posterior pores subtly larger than anterior pores (Fig. 16 G). Ambulacrum III weakly depressed, with very small podial pores (Fig. 16 A, D). Ambulacra II and IV weakly sunken, with two columns (one is rudimentary) of pore-pairs and three times longer and slightly slenderer than ambulacra I and V (Fig. 16 A, D). Ambulacra I and V slightly more sunken than anterior paired ambulacra (II and IV), with equally developed columns of pore-pairs (Fig. 16 A, D). Peripetalous fasciole well-developed and adjacent to paired petals (I, II, IV and V) (Fig. 16 A, C, D). Lateroanal fasciole present (Fig. 16 C, F). Cluster of long spines close to periproct. Periproct circular and longitudinal (Fig. 16 D), covered by plates of variable sizes and shapes, larger plates close to edge and smaller plates towards center. Peristome large, D-shaped, covered by many plates (Fig. 16 B, E). Labrum short and wide (Fig. 16 E), in broad contact with sternal plates, and not projecting strongly towards peristome. Amphisternous plastron, with many primary tubercles. Plastron spines large and spatulate (Fig. 16 B, E). Crenulate and perforate tubercles (Fig. 16 D – F). Aboral tuberculation fine, uniform, and dense (Fig. 16 D). Oral tubercles also dense and uniform (Fig. 16 E). Tube feet with spinules of variable shapes. Pedicellariae. Ophicephalous pedicellariae distributed over entire test. Ophicephalous pedicellariae short and without a neck. Valves short, proximal region large, blade narrow with denticulate margin, which enlarges towards the tip, forming a foramen with denticulate margin. Colour. Spines pinkish gray, darkest at the bases, in living specimens (A. Agassiz 1883). Yellowish in alcohol. Naked test white.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1EFFC0FF551A2DFF74F84B.taxon	distribution	Distribution. Florida, Gulf of Mexico, Mexico, Belize, Bahamas, Barbados, Cuba, Dominican Republic, Trinidad and Tobago, Honduras, Puerto Rico, Nicaragua, Panama, Colombia, Venezuela, Guyana, and Brazil (A. H. Clark et al. 1921; Tommasi 1972; Borrero-Pérez et al. 2002, 2012; Alvarado et al. 2008; Solís-Marín et al. 2013). In Brazil from BA (Manso 2004). From depths of 43 to 900 m (Pawson et al. 2009).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1EFFC0FF551A2DFF74F84B.taxon	discussion	Remarks. The genus Agassizia presently consists of only two extant species [A. excentrica and A. scrobiculata Valenciennes, 1846] (Kroh & Mooi 2016). Agassizia excentrica is the only species in the genus recorded from the Atlantic Ocean. A. Agassiz (1869), in the original description, says that he was not sure that his specimen did not belong to A. porifera (Ravenel, 1848) (fossil). Nevertheless, A. Agassiz (1869) presented a brief description of his new species and the features he used to distinguish it from other known species at that time, but did not provide illustrations. Although A. excentrica is well established taxonomically, a reanalysis of its type series and a redescription of this species is still needed. Few individuals were analyzed in the present paper, all of which were similar in size and without observed variations. A. Agassiz (1883, 1904) provided an extended discussion on the ontogenetic development of morphological characters of A. excentrica. Ecological notes. Agassizia excentrica lives buried in sandy or sand-mud substrates. According to Lewis (1963), this species appears to have typical bottom feeding habits and ingests loose bottom material at random. This is a very rare species for which data on ecology and evolutionary biology are scarce. Along the Brazilian littoral, it has been found in fine sediments together with the ophiuroids Microphiopholis atra (Stimpson, 1854), Amphioplus camamuensis Manso, 2004 and Amphiodia pulchella (Lyman, 1869) (Manso 2004; Manso et al. 2008).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1DFFDDFF551D22FE93FAB3.taxon	description	Figure 17 A – K	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1DFFDDFF551D22FE93FAB3.taxon	description	14 – 15.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1DFFDDFF551D22FE93FAB3.taxon	materials_examined	Material examined. Alagoas: Photographic record (R. B. Moura, personal observation). Bahia: 1 spm, Busca Vida Beach, Guarajuba, Camaçari, 01. V. 1993 [UFBA 00522].	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1DFFDDFF551D22FE93FAB3.taxon	description	Description. Test oval, rounded anteriorly and truncate posteriorly (TL = 103.6 mm; TH = 30.8 mm; TW = 12.3 mm), covered by spines (Fig. 17 A – C). Long and slender spines borne on large primary tubercles occur between petals (Fig. 17 A). Apical system ethmolytic. Petaloid large, with long petals. Ambulacrum III slightly depressed. Peripetalous fasciole well-developed. Peristome kidney-shaped, covered by large plates, with small spines (Fig. 17 B) and triphyllous pedicellariae. Subanal fasciole well-developed, with anal fascioles stretching aborally on either side of periproct. Spines from oral surface of test expanded at base. Spines of plastron numerous and slenderer than on rest of oral surface. Flat oral surface (Fig. 17 B). Periproct small. Narrow amphisternous plastron. Pedicellariae. Small three-valved pedicellariae without stalk or neck on ambulacra at margins of the plastron. Ophicephalous pedicellariae with long stalk, short neck, and short bottle-shaped valves (Fig. 17 F, I). Triphyllous pedicellariae with short valves and large blade (Fig. 17 G, H). Tridentate pedicellariae with short stalks and without a neck. Valves narrow, broadened at the base and narrowing toward the tip (Fig. 17 J). Colour. Living specimens are brown. Naked test white.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1DFFDDFF551D22FE93FAB3.taxon	distribution	Distribution. Florida, Gulf of Mexico, Bahamas, Belize, Cuba, Jamaica, Antilles, Panama, Colombia, and Brazil (H. L. Clark 1925; Hendler et al. 1995; del Valle García et al. 2005). In Brazil from AL, BA, SP, and SC, including Trindade Island (ES) (H. L. Clark 1925; Tommasi 1958 a; Martins et al. 2018). This is the first record of this species in Alagoas State. From depths of 1 to 210 m, but most common in less than 50 m (Serafy 1979).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E1DFFDDFF551D22FE93FAB3.taxon	discussion	Remarks. The genus Plagiobrissus contains three extant species [P. africanus (Verrill, 1871), P. pacificus H. L. Clark, 1940 b and P. grandis] (Kroh & Mooi 2016). Plagiobrissus grandis differs from P. africanus, an endemic species from the African coast, by the well-developed labrum, which is composed of two sternal plates of equal size. According to Kier (1975), young individuals of P. grandis can be easily confused with young specimens of Meoma ventricosa (Lamarck, 1816). However, the presence of long, sharp, white spines on the aboral surface of P. grandis readily distinguishes them, as does the more flattened test of the latter. Kier (1975) described detailed morphological variations in the ontogenetic development of P. grandis. In the present study, a single adult was examined that is insufficient to indicate variations. Ecological notes. Plagiobrissus grandis lives buried in sandy areas where seagrass and algae are sparse or absent (Hendler et al. 1995; del Valle García et al. 2005). Serafy (1979) recorded this species from bottoms formed by shell fragments and quartz sand. According to Kier & Grant (1965), P. grandis is typically found buried at a depth of 50 mm. When buried, P. grandis raises and exposes its long aboral spines, presumably as a defence against predators (Hendler et al. 1995). According to Moore (1956) the gastropods Cassis tuberosa and C. madagascariensis spinella Clench (presently accepted as Cassis madagascariensis Lamarck) are its main predators.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E03FFDEFF551EDCFABAFAC6.taxon	materials_examined	Material examined. No specimens from Brazil were obtained for analysis. Complementary material: 1 spm, Gulf of Naples, Italy [EqMN 315]	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E03FFDEFF551EDCFABAFAC6.taxon	description	Description (modified from Chesher 1966 and Manso 2004). Test cordiform, rather thin, truncated, swollen at the posterior region, covered by short, slender, and vitreous spines. Apical system ethmolytic, with four gonopores. Ambulacrum III strongly sunken, with ambulacral pores arranged irregularly, becoming bigger from apical system towards oral surface. Spines from the margin of ambulacrum III larger than remaining aboral spines. Anterior petals longer than posterior petals. Subanal fasciole and inner fasciole well-developed. Three pairs of pore-bearing plates inside subanal fasciole. Peristome ventral, kidney-shaped, wider than long. Labrum short, bordered by two pairs of ambulacral plates. Amphisternous plastron swollen, covered by long, curved spines, with tips spatulate. Periproct oval on vertical truncate face, covered by plates that diminish in size from margin to center. Pedicellariae. Ophicephalous pedicellariae more abundant around the periproct and tridentate pedicellariae distributed over entire test. Fistulate pedicellariae short, lacking a neck, with a crown of spines of variable sizes at the base of the stalk. Valves (shape similar to an Erlenmeyer flask) with proximal base long (corresponding to about a little over half the length of the valve) and broad, blade short and narrow, ending in a small oval foramen with a denticulate margin, and a large area of muscle insertion. Tridentate pedicellariae with stalk and neck short (both equal in length) and head long. Valves with proximal region short and blade long, with margin serrate. Colour. Brown, yellowish or pinkish. Naked test white.	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E03FFDEFF551EDCFABAFAC6.taxon	distribution	Distribution. Cosmopolitan (Baker & Rowe 1990). In Tropical West Atlantic E. cordatum is only found in Brazil (Solís-Marín et al. 2013). In Brazil from BA (Tommasi 1966 a; Magalhães et al. 2005). From depths of 0 to 230 m (Solís-Marín et al. 2013), with densest populations occurring at depths of 40 m (Duineveld & Jenness 1984).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
C4255B094E03FFDEFF551EDCFABAFAC6.taxon	discussion	Remarks. Presently 12 extant species of Echinocardium are known, only three of which occur in the West Atlantic Ocean (E. cordatum, E. laevigaster A. Agassiz, 1869 and E. mortenseni Thiéry, 1909). Echinocardium cordatum differs from E. mortenseni (Gulf of Mexico and Mediterranean), in the latter having an oval to circular, short inner fasciole and subanal fasciole absent. It differs from E. laevigaster (Florida and Gulf of Mexico) mainly by having a narrow, shallow ambulacrum III. The taxonomic status of E. cordatum has been the focus of heated discussion. Some authors detect strong resemblances with E. fenauxi Péquignat, 1963 (Chenuil & Féral 2003) from the Mediterranean Sea. Furthermore, recent studies indicate that E. cordatum is a cryptic species complex (Egea et al. 2016). Despite several attempts, both morphological and genetic, to resolve the taxonomy of this species (e. g. Higgins 1974; Chenuil & Féral 2003; Féral et al. 1995; Laurin et al. 1994), its status remains confused. Féral et al. (1998) analysed specimens of E. cordatum from the Atlantic and the Mediterranean and did not detect genetic differentiation. Gordon (1927) and Duineveld & Jenness (1984) provide an excellent account of the ontogenetic development of E. cordatum. Ecological notes. This species lives buried in the sediment, mostly in fine to muddy sand and mud in depths from 10 to 20 cm (Tommasi 1966 a; Chenuil & Féral 2003). It feeds on decomposing material (Nichols 1959). According to Rolet et al. (2012), E. cordatum is the best-studied spatangoid regarding ecology and physiology, especially in New Zealand, Japan, and the Mediterranean Sea. In Brazil, the species is apparently rare, having been recorded only in estuarine habitats (Baía de Todos os Santos and Baía de Aratú) (Magalhães et al. 2005).	en	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. Zootaxa 4529 (1): 1-72, DOI: 10.11646/zootaxa.4529.1.1
