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Kroh, Andreas, Gaudin, Jimmy, Reich, Mike (2022): Oloughlinia, a replacement name for Clarkiella Heding in Heding & Panning 1954 (Echinodermata: Holothuroidea: Sclerodactylidae). Zootaxa 5178 (4): 397-399, DOI: https://doi.org/10.11646/zootaxa.5178.4.6

Genus Oloughlinia Kroh, Gaudin & Reich, 2022 nom. nov. = homonym Clarkiella Heding in Heding & Panning, 1954 non Lambert, 1916 (pro Clarkiella Heding in Heding & Panning, 1954, p. 119, non Lambert, 1916, p. 169 [Echinoidea: Faujasiidae]) Synonymy: non 1916 Clarkiella —Lambert, p. 169. non 1953 Genus Clarkiella Lambert —Cooke, p. 18. 1954 Gattung Clarkiella Heding —Heding & Panning, p. 119. 1956 Clarkiella gen. nov. … Heding & Panning—Clark, p. 13. 1966 Clarkiella Heding & Panning 1954 —Edwards & Hopwood, p. 58. 2017 Clarkiella —Martins, p. 18. 2022 Genus Clarkiella Heding, in Heding & Panning, 1954 —Martins & Tavares, p. 569. Type species: Clarkiella discoveryi Heding in Heding & Panning, 1954, pp. 119–120 is designated as type species of Oloughlinia Kroh, Gaudin & Reich, 2022 nom. nov. here, following recommendation 60A of the Code. Gender: feminine Etymology: named after Peter Mark O’Loughlin (Museum Victoria), in acknowledgement of his extensive research on dendrochirote sea cucumbers. Species Oloughlinia deichmannae (O’Loughlin, 2009) nov. comb. —southern Indian Ocean. Species Oloughlinia discoveryi (Heding in Heding & Panning, 1954) nov. comb. —South Atlantic Ocean., Published as part of Kroh, Andreas, Gaudin, Jimmy & Reich, Mike, 2022, Oloughlinia, a replacement name for Clarkiella Heding in Heding & Panning 1954 (Echinodermata: Holothuroidea: Sclerodactylidae), pp. 397-399 in Zootaxa 5178 (4) on page 398, DOI: 10.11646/zootaxa.5178.4.6, http://zenodo.org/record/7031833, {"references":["Heding, S. G. & Panning, A. (1954) Phyllophoridae. Eine Bearbeitung der polytentaculaten dendrochiroten Holothurien des Zoologischen Museums in Kopenhagen. Spolia Zoologica Musei Hauniensis [Skrifter udgivet af Universitetets Zoologiske Museum KObenhavn], 13, 1 - 209.","Lambert, J. (1916) Echinodermes. Revue Critique de Paleozoologie, 20 (4), 166 - 176. Available from: https: // www. biodiversitylibrary. org / page / 33069430 (accessed 16 July 2022)","O'Loughlin, P. M. (2009) BANZARE holothuroids (Echinodermata: Holothuroidea). Zootaxa, 2196 (1), 1 - 18. https: // doi. org / 10.11646 / zootaxa. 2196.1.1"]}

Genus Oloughlinia Kroh, Gaudin & Reich, 2022 nom. nov. = homonym Clarkiella Heding in Heding & Panning, 1954 non Lambert, 1916 (pro Clarkiella Heding in Heding & Panning, 1954, p. 119, non Lambert, 1916, p. 169 [Echinoidea: Faujasiidae]) Synonymy: non 1916 Clarkiella —Lambert, p. 169. non 1953 Genus Clarkiella Lambert —Cooke, p. 18. 1954 Gattung Clarkiella Heding —Heding & Panning, p. 119. 1956 Clarkiella gen. nov. … Heding & Panning—Clark, p. 13. 1966 Clarkiella Heding & Panning 1954 —Edwards & Hopwood, p. 58. 2017 Clarkiella —Martins, p. 18. 2022 Genus Clarkiella Heding, in Heding & Panning, 1954 —Martins & Tavares, p. 569. Type species: Clarkiella discoveryi Heding in Heding & Panning, 1954, pp. 119–120 is designated as type species of Oloughlinia Kroh, Gaudin & Reich, 2022 nom. nov. here, following recommendation 60A of the Code. Gender: feminine Etymology: named after Peter Mark O’Loughlin (Museum Victoria), in acknowledgement of his extensive research on dendrochirote sea cucumbers. Species Oloughlinia deichmannae (O’Loughlin, 2009) nov. comb. —southern Indian Ocean. Species Oloughlinia discoveryi (Heding in Heding & Panning, 1954) nov. comb. —South Atlantic Ocean.

Clypeaster brigitteae n. sp. Figures 3–8, tables 2–5, 11. Diagnosis. Among all known extant taxa, very few Clypeaster species possess completely unique features (i.e. autapomorphies). Therefore, early workers used unique combinations of relatively conspicuous features to diagnose new species, and this is also the approach here. Nevertheless, among the Clypeaster species recorded from the Philippines, Clypeaster brigitteae n. sp. is unique with respect to certain characters, and these are marked with an asterisk in the following list of relevant characters: Clypeaster with all petals widely open*; fewer than 9 respiratory pore pairs counted within a section 5 mm long along the length of single column in each petal; only 1 or 2 primary tubercles in poriferous zone between adjacent respiratory pore pairs*; interambulacral basicoronals separated from first pair of post-basicoronals by two plates from each adjacent ambulacrum; single occluded distal plate (very rarely two) in petal*; aboral primary tubercles not densely distributed (only about 50/cm 2)*; valves of ophicephalous large (largest of trio usually about 400 µm long). Type and other material studied. A total of 12 specimens (Table 2), consisting of a holotype (CASIZ 234824, Mactan Island, Cebu, Philippines) and 11 individually cataloged paratypes (CASIZ 187409, 187411, 187450, 234819–234823, 234825–234827, Balicasag Island, Bohol and Balut Island, Mindanao, Philippines), were available for study, obtained in a donation to the California Academy of Sciences by the second author in 2008–2009, or purchased with export permits by the first author through online dealers in the Philippines. See Table 2 for a complete list of all material of the new species studied, repository numbers, localities and depth, condition of each specimen as of this writing, and morphometrics for major features. Etymology. The new species is named after the late spouse of the second author, Brigitte. Description. This description relies primarily on the holotype, CASIZ 234824 (Figs. 3, 5C, 7, 8). However, in order to maintain this specimen intact with nearly complete spination, other specimens were used in dissections and other preparations to obtain additional data and imagery (as noted in Table 2 and in the figure captions). Meristics and percentage calculations are from the holotype unless otherwise noted. Measurements from all specimens are given in Table 2. Holotype 93.8 mm TL (size range of type series 62.7–94.6 mm TL, Table 2), 85.1 mm TW, 25.2 mm TH. Ambitus relatively thick, rounded, approximately 8% TL thick through point at which oral surface begins to flatten (Fig. 3). Aboral surface flat near ambitus, but rising close to perimeter to form slight dome (Fig 3). Oral surface flat near ambitus, but deeply indented upwards in centre to form well-developed infundibulum (Fig. 3, and see below). Test broad (TW 90.7% TL), perimeter pentangular with broadly rounded vertices in ambulacra, very shallow indentations in paired interambulacra, slightly convex in posterior interambulacrum (V). Highest point of test at centre of apical system (madreporic plate), approximately 50% TL from anterior end. TH 26.9% TL, but variable among known specimens, see Table 2). Apical system —Monobasal, pentangular (not star-shaped), 49.9% TL from ocular III to anterior edge of test, length ~6% TL, numerous hydropores scattered over slightly domed madreporic plate. Five gonopores, one in each interambulacrum, located at suture between madreporic plate and first adapical plates of interambulacral column (Fig. 4A). Peristome —Circular to slightly pentangular, relatively small, about 6.6% TL. Anterior edge of peristome 48.1% TL from anterior edge of test. Peristome at top of deep infundibulum that extends 12.3 % TL (46.0% TH) upward from oral surface (Figs. 3, 5A). Numerous irregular stereom spicules embedded in peristomial membrane, but none bearing spines (Fig. 7H). Periproct —Small, length about 4.5% TL (92.0% TL from anterior ambitus, that is to say, posterior edge of the periproct situated approximately length of periproct from ambitus on oral surface). Adoral edge of periproct in con- tact with third and fourth post-basicoronals (interambulacral plates 4a and 5b), aboral edge with fourth and fifth (5a and 6b) (Fig. 5B). Periproctal membrane slightly sunken, covered in small plates (slightly larger near peristomial edge) surrounding small, central anal opening, plates bearing dense cover of miliary spines, triphyllous pedicellariae, and relatively small tridentate pedicellariae, but no primary spines (Fig. 5C). Ambulacra —Petaloid adapically, petals broad, with pore pair columns continuously diverging or parallel distally, only slightly converging in some cases (Figs. 3, 4B, C). Ambulacra widen near ambitus in oral view, approximately twice width of interambulacra at ambitus (Figs. 5A, B), most commonly with 8 to 10 plates in any given column (half-ambulacrum). Ambulacra with paired basicoronals, all in agreement with Lovén’s Rule (sensu David et al. 1996), ambulacral plates Ib, IIb, IIIa, IVb, Va about 2/3 length of other plate in pair (Fig. 5B). All ambulacral plates with densely arranged unipores scattered between spine tubercles, each unipore supporting single accessory tube foot. Petals —Posterior paired petals (I and V) slightly longer than anterior paired petals (II and IV). Posterior paired petals longest relative to TL, petal V 60.0% length of corresponding test radius in planar view, but 32.5% TL; anterior paired petals next longest, petal IV 64.8% length of corresponding test radius, but 31.2% TL; anterior unpaired petal shortest, petal III 59.4% length of corresponding test radius, but 29.6% TL. Petal V width at widest point 19.0% TL, interporiferous zone 12.0% TL, distance between inner pores of distalmost respiratory pore pair in petal 14.2% TL (117.7% of maximum interporiferous zone width and a measure of the “openness” of the petal at its distal end, see Materials and Methods and Fig. 2); petal IV width 16.3% TL, interporiferous zone 11.2% TL, distance between inner pores of distalmost respiratory pore pair in petal 11.2% TL (100.0% of maximum interporiferous zone width and a measure of the “openness” of the petal at its distal end, see Materials and Methods and Fig. 2); petal III width 18.1% TL, interporiferous zone 11.8% TL (92.8% of maximum interporiferous zone width and a measure of the “openness” of the petal at its distal end, see Materials and Methods and Fig. 2). Respiratory tube foot pore pairs strongly conjugated, inner pore slightly elongate or almost circular, outer pore only slightly elongated (Figs. 4B, C, D). Column a of petal V with 42 pore pairs; of petal IV with 44; of petal III with 41. One or two (rarely three) primary tubercles situated along ridge between adjacent pore pairs. Plating within petals of typical clypeasterid architecture, with primary plate spanning entire distance from outer edge of petal to perradial suture, alternating with demiplate spanning just less than half that distance from outer edge of petal (Figs. 4B, C, D). One or two occluded plates present at tips of petals (e.g. Fig. 4D). Interambulacra —Narrower than ambulacra and straight on oral surface, slightly widening towards ambitus, but containing paired, zig-zag plates between petals right up to madreporic plate. Typically, five, occasionally six, post-basicoronal plates in each half-interambulacrum in oral interambulacra. Single, narrow basicoronal in each interambulacrum (Fig. 5D), typically but not always reaching ends of adjacent ambulacral basicoronals. Interambulacra disjunct, separated from first pair of interambulacral post-basicoronals by two adjacent oral ambulacral postbasicoronals (Fig. 5B, 11). In some specimens, first interambulacral post-basicoronal in one column can be greatly reduced so that it is not contiguous with second post-basicoronal in corresponding column (see interambulacra 2 and 3 in Fig. 5B), but it is not known if this is typical or unique, as it was decided not to prepare additional specimens to produce plate maps for other members of type series. Tubercles —Of two main types, primary and miliary. Aboral miliary tuberculation homogeneous, primary tubercles relatively sparse, only about 50/cm 2. Oral tubercles slightly larger than those on aboral surface, particularly in interambulacra. Tubercles perforate, slightly crenulate, primaries with distinct, deeply sunken areole, miliaries with shallow areole, filling spaces between primaries and greatly outnumbering them (orally by at least 10 to 1, and approximately 20 to 1 on aboral surface). Abundant miliary tubercles on apical system (Fig. 4A) and periproctal membrane (Fig. 5C), primary tubercles lacking in those areas. No tubercles on peristomial membrane. Food grooves —Unbranched, extending from edge of peristome, following perradial suture, shallowing and narrowing gradually, fading to merge with test surface just short of ambitus (Fig. 5A). Populated by specialized food groove miliary spines (Figs. 7F, G). Internal structure —Strongly developed, with four or five peripheral, circumferentially arranged walls (sensu Mooi 1990; Mihaljević et al. 2011) in ambulacra, spanning space between floor and ceiling of test (Figs. 6A, B). Some reduced walls on inner side of channel that bears gut. Complicated masses of bars in interambulacra, continuing on proximal side of gut towards and beneath lantern (Figs. 6A, B). Minute, extremely thin, attenuated needles on both floor and ceiling of test (Figs. 6D–F), most strongly expressed in ambulacra. Some needles span entire space between floor and ceiling to join together to form thin columns or pillars (Fig. 6F). Lantern —Of typical clypeasterine morphology (Mooi 1989, 1990), not protrusible, with strongly developed outer wings separated from almost rudimentary inner wings by continuous, ring-like arrangement of supra-alveolar processes at summit of lantern (Fig. 6G). Epiphyses and rotules rudimentary, compasses absent, as typical in clypeasteroids (Mooi 1989, 1990). Wings of each pyramid, particularly outer wings, strongly convoluted with subdivided walls connecting ambulacrally situated pyramidal muscle attachment surfaces to interambulacrally positioned symphysis between demi-pyramids underlying tooth slide (Figs. 6G, H). Tooth slide almost vertical, but biting surfaces of keeled tooth strongly angled relative to longitudinal axis of tooth to meet along nearly vertical axis from mouth through top of lantern. When in life position over peristome at top of infundibulum, entire lantern slightly canted forward so that supra-alveolar processes at anterior of lantern noticeably lower than at posterior (Fig. 6C). Lantern supports (auricles) of typical clypeasterid configuration (Kier 1970; Mooi 1989, 1990), consisting of pairs of elongated, relatively thin, upright processes, one support for every pair contributed by adjacent ambulacral basicoronal (Fig. 6B). Spines —Of two main types, primary (Figs. 7A, B) and miliary (Figs. 7C–F), corresponding to tubercle types described above, and as typical for other Clypeaster (Telford et al. 1987). Aboral primaries only about half length of oral primaries, much narrower, finer with about 75% number of longitudinal ribs (Figs. 7A, B). Core of spine sometimes hollow, sometimes with nearly solid meshwork (Fig. 7A, B). So-called “hyaline point” (Serafy 1971) present at tips of undamaged, uneroded primaries on both aboral and oral surfaces (Figs. 7A, B). Miliary spines of variable length (Figs. 7C–F), rounded, blunt, slightly swollen distally but not crown-shaped, longest in interambulacral regions of oral surface, aboral interambulacral miliaries (Fig. 7D) about 80% length of oral interambulacral miliaries (Fig. 7C), aboral ambulacral miliaries shorter still, only 50% length of oral interambulacral miliaries (Fig. 7E). Miliary spines within food grooves short, only about half length of miliaries from adjacent oral ambulacra (Fig. 7F). authors (CASIZ 232217). ...Continued on the next page 1 Known upper size limit, rounded up to the nearest 5 mm, based on examination of CASIZ, MNHN, and USNM specimens, and descriptions in the literature and of type series. 2 Approximate, range based on examination of CASIZ, MNHN, and USNM specimens, and descriptions in the literature and of type series. 3 Percentage based on approximate point at which test begins to inflect upwards towards the peristome – measured from the ambitus along oral interambulacrum 4. 4 In C. reticulatus, the margin is inflated relative to the rest of the distal part of the oral surface so that the distal ends of the petals are below the altitude of the margin. 5 Based on new observations and radiographs in Mihaljević et al. (2011). Structure described is only that distal to the path of the gut. Terminology adopted from Mooi (1989, 1990) and Mihaljević et al. (2011): bars = interambulacral, peripheral radiating buttresses (but not actually “radial” as used in Mihaljević et al. [2011]); walls = concentric, circumferential buttresses made of peripheral ambulacral pillars connected to form nearly solid partitions. 6 Number of ambulacral plates in each column adjacent to interradial suture between basicoronal and first post-basicoronal in posterior interambulacrum (interambulacrum 5). See also Figs. 5B, 11. Pedicellariae — Of three main types: triphyllous (Fig. 8F), tridentate (Figs. 8C–E), and ophicephalous (Figs. 8A, B). Following observations pertain to all specimens with well preserved complement of external appendages. Triphyllous widely distributed over entire test, extremely small and numerous, with long, flexible necconnect- ing three rounded valves to thin but robust stem, flexible neck about same length as stem, valves of usual shape for Clypeaster, with extremely short teeth on distal edges (Fig. 8F). Tridentates principally of two sizes, with three valves, those of smaller form (Fig. 8C) only slightly shorter than those of larger form (Fig. 8E). Smaller version scattered widely over oral surface, densely distributed (as many as 5/mm 2), particularly in infundibulum, much rarer aborally, nearly absent in aboral regions where ophicephalous pedicellariae most densely distributed. Larger tridentates tending to be more common near peristome and periproct. Only small version found on periproctal membrane. Valves of small form meet distally for about one third length of valve when pedicellaria closes, versus about one quarter in large form. Valves of both types with small proximal teeth along edges where valves meet, but with one, or more typically two longer distal teeth (Figs. 8C–E). Entire tridentate jaw connected to robust stem (Fig. 8D) by flexible neck about half as long as stem. Ophicephalous pedicellariae apparently absent on oral surface, but on aboral surface, extremely densely distributed (as many as 4/mm 2) all around peripheral portion of test inward from ambitus to point at which test slopes steeply upward towards apical system. Ophicephalous rare in this inner region, particularly proximal to apical system. Ophicephalous of typical Clypeaster morphology (Figs. 8A, B), with 3-valved jaw directly attached to top of vase-like stem (Fig. 8B), each valve connected to stereom meshwork in inner region of distal cup at top of stem by collagenous straps weaving between handles, inserting proximal to hinge of each valve. Handles always of three different lengths in single jaw, smallest nesting within medium-sized, then medium-sized within longest (Fig. 8A). Distal part of each valve with rounded, helmet-shaped head, inner surface of head with unbroken, oval ring of>30 prominent, sharp teeth, usually 4–6 additional teeth between head and top of hinge (Fig. 8A). Tube feet —Of respiratory and accessory types, as typical for Clypeaster (Mooi 1986). Respiratory tube feet leaf-like, straddling conjugated respiratory tube foot pores in petals as described above, lacking spicules. Accessory tube feet varying in size and shape, distributed as typical for other Clypeaster (Mooi 1986; Telford et al. 1987), tube feet extending from unipores distributed on all ambulacral plates, occasionally on interambulacrals in areas bordering the ambulacra as well, although not on interambulacral basicoronals (Fig. 5D). Smaller accessory tube feet in food grooves, small number of somewhat larger food groove tube feet just distal to slightly enlarged buccal tube feet as described for other species (Mooi 1986). Spicules in tips of accessories (not known for buccals) like those of other Clypeaster, but elaborate, with two to three concentric rings of fenestrae, and peripheral projections pointing both proximally and distally (Fig. 7I). Sphaeridia —Two per ambulacrum, one in each ambulacral basicoronal, located at outer edges of food groove near peristome just distal to buccal tube feet (Fig. 5D), consisting of solid sphere of imperforate stereom inside chamber entirely enclosed within surrounding ambulacral plate. Colour. Unknown in living specimens. Aboral surface of dried specimens brown to purplish or grayish brown, oral surface slightly paler, approaching beige in interambulacra of some specimens, centres of some plates slightly darker, particularly aborally, so that major sutures and tube foot fields stand out in contrast (Fig. 3). Occurrence. Holotype collected by tangle net off Mactan Island, Cebu, Philippines, four paratypes from Mactan Island, four from Balicasag Island, Bohol, and three from Balut Island, Mindanao (Fig. 1, Table 1), at depths from 100 to 200 m. Biology. C. brigitteae n. sp. has not been observed in its habitat, but gut and food groove contents suggest that it lives on soft bottoms with low particle size ranges. Food preferences are only partially known, because the gut contents are significantly broken up or pulverized, presumably by chewing action of the lantern (Telford et al. 1987). Materials remaining in the food grooves suggest that the animal is a detritivore, with some preference for foraminiferans. However, as the specimens were primarily collected by tangle net, material normally gathered by the animals before capture could have been lost, and no quantitative assessments were possible. Low gonopore size suggests that the species possesses planktotrophic larvae (F. Armstrong and R. Mooi unpublished data concerning gonopore size in the genus). Remarks. Clypeaster brigitteae n. sp. appears to fit some of the features listed by Mortensen (1948b) for the section (or subgenus) Stolonoclypus A. Agassiz, 1863, given the well-developed bars and walls of the internal skeleton. However, even Mortensen (1948b: 88) was concerned that the “numerous species here enclosed in this section evidently do not form a natural group”. Mihaljević et al. (2011) were also unable to recover these so-called subgenera as monophyletic taxa, and this nomenclature is therefore abandoned, as indicated in Kroh & Mooi (2020). C. brigitteae n. sp., Published as part of Mooi, Rich & Noordenburg, Henk Van, 2021, A new species and comparative morphology of Philippine sea biscuits (Echinoidea Clypeaster), pp. 1-36 in Zootaxa 4964 (1) on pages 9-18, DOI: 10.11646/zootaxa.4964.1.1, http://zenodo.org/record/4706613, {"references":["Loven, S. (1874) Etudes sur les echinoidees. Kongelige Svenska Vetenskaps-Akademiens Handlingar, 11, 1 - 91.","David, B., Mooi, R. & Telford, M. (1996) The ontogenetic basis of Loven's Rule clarifies homologies of the echinoid peristome. In: Emson, R., Smith, A. B. & Campbell, A. (Eds.), Echinoderm Research 1995. A. A. Balkema, Rotterdam, pp. 155 - 164.","Mooi, R. (1990) Paedomorphosis, Aristotle's lantern, and the origin of the sand dollars (Echinodermata: Clypeasteroida). Pa- leobiology, 16, 25 - 48. https: // doi. org / 10.1017 / S 0094837300009714","Mihaljevic, M., Jerjen, I. & Smith, A. B. (2011) The test architecture of Clypeaster (Echinoidea, Clypeasteroida) and its phylogenetic significance. Zootaxa, 2983 (1), 21 - 38. https: // doi. org / 10.11646 / zootaxa. 2983.1.2","Mooi, R. (1989) Living and fossil genera of the Clypeasteroida (Echinoidea: Echinodermata): An illustrated key and annotated checklist. Smithsonian Contributions to Zoology, 488, 1 - 51. https: // doi. org / 10.5479 / si. 00810282.488","Kier, P. M. (1970) Lantern support structures in the clypeasteroid echinoids. Journal of Paleontology, 44, 98 - 109.","Telford, M., Mooi, R. & Harold, A. S. (1987) Feeding activities of two species of Clypeaster (Echinoides, Clypeasteroida): Further evidence of clypeasteroid resource partitioning. Biological Bulletin, 172, 324 - 336. https: // doi. org / 10.2307 / 1541712","Serafy, D. K. (1971) A new species of Clypeaster (Echinodermata, Echinoidea) from San Felix Island, with a key to the recent species of the Eastern Pacific Ocean. Pacific Science, 25, 165 - 170.","Koehler, R. (1922) Echinides du Musee Indien a Calcutta, II. Clypeastrides et Cassidulides. Echinoderma of the Indian Museum, 9, 5 - 161.","Leske, N. G. (1778) Jacobi Theodori Klein naturalis dispositio echinodermatum …, edita et descriptionibus novisque inventis et synonomis auctorem aucta. Addimenta ad I. T. Klein naturalem dispositionem Echinodermatum. G. E. Beer, Leipzig, xxii + 278 pp.","Doderlein, L. (1885) Seeigel von Japan un den Liu-Kiu-Inselin. Archiv fur Naturgeschichte, 51, 73 - 112. https: // doi. org / 10.5962 / bhl. part. 1569","Lamarck, J. - B. (1816) Histoire Naturelle des Animaux sans Vertebres, presentant les caracteres generaux et particuliers de ces animaux, leur distribution, leur classes, leurs familles, leurs generes, et le citation des principales especes qui s'y rapportent; precedee d'une Introduction offrant la Determination des caracteres essentielles de l'animal, sa distinction du vegetal et des autres corps naturels, enfin, l'Exposition des Principes fondamentaux de la Zoologie. Tome Troisieme. Verdiere, Paris, 586 pp. https: // doi. org / 10.5962 / bhl. title. 40014","Mortensen, T. (1948 a) Contributions to the biology of the Philippine Archipelago and adjacent regions. Report on the Echinoidea collected by the United States Fisheries Steamer \" Albatross \" during the Philippine Expedition, 1907 - 1910. Part 3: The Echinoneidae, Echinolampidae, Clypeastridae, Arachnoididae, Laganidae, Fibulariidae, Urechinidae, Echinocorythidae, Palaeostomatidae, Micrasteridae, Palaeopneustidae, Hemiasteridae, and Spatangidae. Smithsonian Institution, United States National Museum Bulletin, 100, 93 - 140.","Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. 1. Editio Decima, reformata [10 th Revised Edition]. Laurentius Salvius, Holmiae, 824 pp. https: // doi. org / 10.5962 / bhl. title. 542","Mooi, R. (1986) Non-respiratory podia of clypeasteroids (Echinodermata, Echinoides): II. Diversity. Zoomorphology, 106, 75 - 90. https: // doi. org / 10.1007 / BF 00312110","Mortensen, T. (1948 b) A Monograph of the Echinoidea. IV, 2. Clypeastroida. Clypeastridae, Arachnoididae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp.","Agassiz, A. (1863) List of the echinoderms sent to different institutions in exchange for other specimens, with annotations. Bulletin of the Museum of Comparative Zoology at Harvard College, 1, 17 - 28.","Kroh, A. & Mooi, R. (2020) World Echinoidea Database. Available from: http: // www. marinespecies. org / echinoidea (accessed 15 May 2020) https: // doi. org / 10.14284 / 355","Clark, H. L. (1941) Reports on the scientific results of the Atlantis expeditions to the West Indies, under the joint auspices of the University of Havana and Harvard University. The Echinoderms (other than holothurians). Memorias de la Sociedad Cubana de Historia Natural, 15, 1 - 154.","Agassiz, A. (1869) Preliminary report on the Echini and star-fishes dredged in deep water between Cuba and the Florida Reef, by L. F. de Pourtales, Assist. U. S. Coast Survey. Bulletin of the Museum of Comparative Zoology at Harvard College, 1, 253 - 308.","Tenison-Woods, J. E. (1878) The Echini of Australia. Proceedings of the Linnean Society of New South Wales, 2, 145 - 176."]}

A new species of clypeasterid sea biscuit, Clypeaster brigitteae n. sp., is described from material collected in the Philippines at depths between 100 and 200 m. The new taxon increases the number of Clypeaster species recorded from the Philippines to nine, representing nearly a quarter of the world’s diversity of the genus. Other Philippine species include: C. annandalei Koehler, 1922; C. fervens Koehler, 1922; C. humilis (Leske, 1778); C. japonicus Döderlein, 1885; C. latissimus (Lamarck, 1816); C. pateriformis Mortensen, 1948; C. reticulatus (Linnaeus, 1758); and C. virescens Döderlein, 1885. Using type material where available, each of these species is compared and contrasted with C. brigitteae n. sp. in tables consisting of new data derived from general test shape and size, petal structure, food grooves, plate architecture, internal structure, and morphology of spines, pedicellariae, and tube feet.

Madeira, Patrícia, Kroh, Andreas, Cordeiro, Ricardo, De, António M., Martins, Frias, Ávila, Sérgio P. (2019): The Echinoderm Fauna of the Azores (NE Atlantic Ocean). Zootaxa 4639 (1): 1-231, DOI: https://doi.org/10.11646/zootaxa.4639.1

Echinocyamus scaber macrostomus Mortensen, 1907 Reports for the Azores Echinocyamus macrostomus $ Mortensen, 1907: 36–37, pl. 12, figs. 2, 7, 17, 24; $ Koehler 1909: 235, pl. 4, figs. 9–10; Mortensen 1927a: 315, 1948: 183–184; García-Diez et al. 2005: 51; Madeira et al. 2011: 255; Echinocyamus scaber macrostomus Mortensen, 1907 — $ Mironov & Sagaidachny 1984: 186–187, fig. 2.2; Mironov 2006: 113– 114; Mironov 2014: 124. See: Mortensen (1907; 1927b: 30–31); Mironov & Sagaidachny (1984). Occurrence: North Atlantic, in the west from the Blake Plateau to Cuba (Mironov 2014), in the east from Portugal to Cape Verde (Mortensen 1907, 1927b) including the archipelagos of the Azores and Madeira (Mortensen 1907) and the Josephine, Tropic and Meteor seamounts (Mironov 2006). The subspecies E. scaber scaber de Meijere, 1903 is reported from the Indo-Pacific (Mironov & Sagaidachny 1984). Depth: 1,010 –2,820 m, though bare tests have been reported as deep as 3,140 m (Mironov & Sagaidachny 1984); AZO: 1,560 –2,178 m (Mortensen 1907, Koehler 1909). Habitat: mud to sand (Koehler 1909). Remarks: Mortensen (1907) described two species of Echinocyamus, E. macrostomus and E. grandiporus using material, which included animals collected in Azorean deep waters. Mortensen (1907) considered his two new deep-water species closely related though E. macrostomus tended to live in deeper waters. In contrast, Mironov & Sagaidachny (1984) considered E. macrostomus closely related with E. scaber and downgraded Mortensen species to a variety of the later. Additionally, Mironov (2006) observed that some of the specimens from Meteor and Antialtair seamounts presented intermediate characteristics between this subspecies and E. grandiporus, suggesting that they could represent hybrids., Published as part of Madeira, Patrícia, Kroh, Andreas, Cordeiro, Ricardo, De, António M., Martins, Frias & Ávila, Sérgio P., 2019, The Echinoderm Fauna of the Azores (NE Atlantic Ocean), pp. 1-231 in Zootaxa 4639 (1) on pages 134-135, DOI: 10.11646/zootaxa.4639.1, http://zenodo.org/record/3342161, {"references":["Mortensen, T. (1907) Echinoidea (Part 2). In: The Danish Ingolf-Expedition 1895 - 1896. 4 (2). Bianco Luno, Copenhagen, pp. 1 - 200.","Koehler, R. (1909) Echinodermes provenant des campagnes du yacht Princesse-Alice (Asteries, Ophiures, Echinides et Crinoides). Resultats des campagnes scientifiques accomplies sur son yacht par Albert Ier Prince Souverain de Monaco, 34, 1 - 317.","Mortensen, T. (1927 a) Handbook of the echinoderms of the British Isles. Oxford University Press, viii + 471 pp. https: // doi. org / 10.5962 / bhl. title. 6841","Mortensen, T. (1948) A Monograph of the Echinoidea. IV. 2. Clypeasteroida. Clypeasteridae, Arachnoidae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp.","Garcia-Diez, C., Porteiro, F. M., Meirinho, A., Cardigos, F. & Tempera, F. (2005) Taxonomic review of selected invertebrate groups collected during the Campaigns of the Prince Albert I of Monaco in the Azorean waters. Arquipelago. Life and Marine Sciences, 22 A, 35 - 59.","Madeira, P., Kroh, A., Cordeiro, R., Meireles, R. & Avila, S. P. (2011) The fossil echinoids of Santa Maria Island, Azores (Northern Atlantic Ocean). Acta Geologica Polonica, 61 (3), 243 - 264.","Mironov, A. N. & Sagaidachny, A. Y. (1984) [Morphology and distribution of the recent echinoids of the genus Echinocyamus (Echinoidea: Fibulariidae)]. Trudy Instituta Okeanologii AN, USSR, 119, 179 - 204. [in Russian, English summary]","Mironov, A. N. (2006) Echinoids from seamounts of the north-eastern Atlantic, onshore / offshore gradients in species distribution. In: Mironov, A. N., Gebruk, A. V. & Southward, A. J. (Eds.), Biogeography of the North Atlantic Seamounts. KMK Scientific Press, Russian Academy of Sciences, P. P. Shirshov Institute of Oceanology, Moscow, pp. 96 - 133.","Mironov, A. N. (2014) Deep-sea fauna of European seas: An annotated species check-list of benthic invertebrates living deeper than 2000 m in the seas bordering Europe. Echinoidea. Invertebrate Zoology, 11 (1), 120 - 129. https: // doi. org / 10.15298 / invertzool. 11.1.12","Mortensen, T. (1927 b) Sur les echinides recueillis par l'expedition du \" Travailleur \" et du \" Talisman. \" Archives du Museum d'Histoire Naturelle, 2 (6), 21 - 34.","Meijere, J. C. H. de (1903) Vorlaufige Beschreibung der neuen, durch die Siboga-Expedition gesammelten Echiniden. Tijdschrift der Nederlandsche Dierkundige Vereeniging, Series 2, 8 (1), 1 - 16."]}

Fibularia coffea sp. nov. [New Japanese name: Kohi-mame-uni] Figs. 3���8; Tables 1, 2; Electronic Supplementary Table S1. Non Fibularia japonica Shigei 1982: 11 ���16, figs. 1���48 (part). Non Fibularia plateia Schultz 2005: 321, fig. 603. Fibularia n. sp. ���bean��� Gomes & Mooi 2015: poster presentation. Material examined. Holotype: NSMT E-10371, whole, with spines, Ayamaru Cape, Amami-Oshima Island, Kagoshima Prefecture, Japan (28��28���26���N, 129��43���09���E), depth 1 m, snorkeling, coll. H. Tanaka, 23 Mar. 2015. Paratypes: 1 specimen, NSMT E-10372, whole, denuded, SEM stub, Tsuchi-Hama, Amami-Oshima Island, Kagoshima Prefecture, Japan (28��24���36���N, 129��40���47���E), depth 2 m, snorkeling, coll. H. Tanaka and L. Sakamoto, 20 Mar. 2015; 2 specimens, NSMT E-10373, whole, denuded, SEM stub, Kunigami, Nishinoomote city, Tanegashima Island, Kagoshima Prefecture, Japan (30��48���19���N, 131��01���24���E), depth 4���5 m, scuba diving, coll. H. Yamasaki and R. Yoshida, 1 Mar. 2014; 1 specimen, NSMT E-10374, whole, denuded, SEM stub, Tateyama Bay, Tateyama city, Chiba Prefecture, Japan (34��58���56���N, 139��48���43���E), depth 6.9���8.1 m, dredging, coll. Y. Yoshida, 14 Jan. 2015; 1 specimen, NSMT E-10375, whole, denuded, SEM stub, Tateyama Bay, Tateyama city, Chiba Prefecture, Japan (34��59���05���N, 139��48���57���E), depth 9���12 m, dredging, coll. Y. Yoshida, 12 May. 2014; 1 specimen, NSMT E-10376, whole, denuded, SEM stub, Tateyama Bay, Tateyama city, Chiba Prefecture, Japan (34��58���40���N, 139��46���05���E), depth 5 m, scuba diving, coll. K. Kosoba, 30 Aug. 2015; 29 specimens, NSMT E-10377, dead tests, Tomioka, Amakusa group, Kumamoto Prefecture, Japan, dredging, coll. H. Tanaka, 21 Mar. 2015; 1 specimen, NSMT E-10378, dead test, Ayamaru Cape, Amami-Oshima Island, Kagoshima Prefecture, Japan (28��28���30���N, 129��43���05���E), beachcombing, coll. H. Tanaka, 23 Mar. 2015; 7 specimens, NSMT E-10379, dead tests, Ayamaru Cape, Amami-Oshima Island, Kagoshima Prefecture, Japan (28��28���30���N, 129��43���05���E), beachcombing, coll. H. Arima, 11 Dec. 2013 ��� 27 Mar. 2014; 7 specimens, NSMT E-10380, dead tests, Kosyuku, Amami-Oshima Island, Kagoshima Prefecture, Japan (28��24���10���N, 129��28���04���E), beachcombing, coll. H. Arima, 13 Dec. 2013 ��� 22 Mar. 2015; 2 specimens, NSMT E-10381, dead tests, Kosyuku, Amami-Oshima Island, Kagoshima Prefecture, Japan (28��24���10���N, 129��28���04���E), beachcombing, coll. H. Tanaka, 22 Mar. 2015; 5 specimens, NSMT E-10382, dead tests, Tsuchi-Hama, Amami-Oshima Island, Kagoshima Prefecture, Japan (28��24���39���N, 129��40���37���E), beachcombing, coll. H. Tanaka, 15 Jul. 2014; 3 specimens, NSMT E-10383, dead tests, Tsuyazaki, Fukutsu city, Fukuoka Prefecture, Japan (33��45���59���N, 130��23���03���E), beachcombing, coll. K. Wakabayashi, 17 Jun. 2005; 10 specimens, NSMT E-10384, dead test, Wada Beach, Oi county, Fukui Prefecture, Japan (35��29���41���N, 135��34���32���E), beachcombing, coll. H. Tanaka, 3 Jan. 2014; 4 specimens, NSMT E-10385, dead tests, Yoan Beach, Amami-Oshima Island, Kagoshima Prefecture, Japan (28��24���12���N, 129��38���35���E), beachcombing, coll. H. Tanaka and L. Sakamoto, 31 Jan. 2016; 48 specimens, NSMT E-10386, dead tests, Zushi Beach, Zushi city, Kanagawa Prefecture, Japan (35��17���18���N, 139��34���25���E), beachcombing, coll. H. Tanaka, 21 Jul. 2012 ��� 2 Feb. 2014; 2 specimens, NSMT E-10387, whole, denuded, Tsuchi-Hama, Amami-Oshima Island, Kagoshima Prefecture, Japan (28��24���36���N, 129��40���47���E), intertidal, snorkeling, coll. H. Tanaka and L. Sakamoto, 24 Mar. 2015; 1 specimen, NSMT E-10388, whole, denuded, Tsuchi-Hama, Amami-Oshima Island, Kagoshima Prefecture, Japan (28��24���36���N, 129��40���47���E), intertidal, snorkeling, coll. H. Tanaka and L. Sakamoto, 22 Oct. 2016. One specimen, UMUTZ-Ecn-SG10-17T No. 7 (as a paratype of Fibularia japonica), dead test, off Misaki Marine Biological Station, Sagami Bay, sublittoral zone, coll. K. Aoki, J. Deguchi, T. Sekimoto, H. Suzuki, and M. Shigei, 1926���1978. GenBank accession number. LC388935 (holotype: NSMT E-10371, Amami-Oshima Island, Kagoshima Prefecture, Japan), LC388934 (paratype: NSMT E-10374, Tateyama Bay, Tateyama city, Chiba Prefecture, Japan). Diagnosis. Test outline elliptical when viewed from above; height low; oral surface slightly depressed toward the peristome. Periproct outline round square shaped. Petaloid region large; number of pores of petal III continues to increase with the test growth, reaching over 20 at TL> 5 mm. Diameter of genital pores equal to or smaller than that of petaloid pores in mature individuals. Two hydropores opening in an irregularly-shaped groove. Black pigments on each aboral perradius, forming symmetric pentaradial in living animals. Description. The test is very small (TL = 1.53���9.73 mm) (Fig. 3), flattened (TH/TL = 0.38���0.55) (Fig. 3C), elliptical when viewed from above (TW/TL = 0.67���0.86), and truncated posteriorly (Figs. 3A, B). The test proportion hardly changes with the test growth (slope value is 1.0 between TW and TL, and 1.1 between TH and TL in the allometry regression, Table 1). The oral surface is slightly depressed around the peristome. The aboral surface is slightly arched convex. There are no internal buttresses. Food grooves are absent (Fig. 3B). The ambulacra are almost the same width as the interambulacra (Figs. 3 D���F). The height of both ambulacral and interambulacral plates are lower than the width at the ambitus (Fig. 3F). The petaloid region is large (PL/TL = 0.46���0.75, PW/TL = 0.33���0.61). The ratio of petaloid region size to test size becomes larger with test growth (slope value is 1.3 between PL and TL, as well as between PW and TL in the allometry regression). Each petal is composed of almost parallel series of pore pairs lying oblique, and crossing the ambulacral plates (Fig. 3D). The number of pores of petal III, IV, and V is continuously increasing up to 40, 28 and 36, respectively, with the test growth (Fig. 4). The pores become larger towards the distal tip of the petals. The peristome, situated at the center of the oral side, is small (SL/TL = 0.17���0.34, SW/TW = 0.14���0.30) and slightly elongated antero-posteriorly (Figs. 3B, E). The ratio of peristome size to test size becomes smaller with the test grows (slope value is 0.8 between SL and TL, as well as between SW and TL in the allometry regression). The peristomial membrane lacks spine and pedicellariae. Two buccal pores are situated in each ambulacrum at the edge of the peristome. Single sphaeridium is fully enclosed within the test, and in the sphaeridial chamber in each ambulacrum near the peristome. The rounded square shaped periproct is located halfway between the peristome and posterior margin of the test and is smaller than the peristome (AL/TL = 0.08���0.15, AW/TW = 0.08���0.16) (Figs. 3B, E) and covered by 4���6 (mainly 5) naked radiating periproctal plates. The ratio of periproct size to test size hardly changes with the test grows (slope value is 1.0 between AL and TL, and 0.9 between AW and TL in the allometry regression). The apical system is situated slightly anterior on the aboral surface (Figs. 3A, D). It consists of four genital pores, five ocular pores in small ocular plates, and two hydropores in a deep, irregularly-shaped groove (Fig. 5A). The diameter of the genital pores (Color. The color is yellow to brown in life (Figs. 8A, B) but changes to green when preserved in ethanol. Black pigments are apparent in each ambulacrum along the pore pair columns, forming a pentaradially symmetric pattern on the aboral surface (Figs. 8A, B). The black pigments remain even after preservation in ethanol. The denuded test is whitish. Distribution. This species is recorded in Japanese waters from Sagami Bay to the Amami-Oshima Islands; 1��� 12 m in depth (present study). Schultz (2005) recorded F. plateia from Queensland, Australia but the figured specimen seems to be F. coffea from the flattened test and the large number of pores of petals (the number of pores of petal III, IV and V are 32, 22 and 32, respectively). In addition, Gomez & Mooi (2015) recorded an unknown species Fibularia n. sp. ���bean��� from the Philippines and Micronesia that seems to be the same as F. coffea from the flattened test and the large number of pores of petals (the number of pores of petal III, IV and V are 34, 26 and 33, respectively; counted from the sketch image). Therefore, this species is considered to be widely distributed across the Indo-Pacific. Habitat. The micro-habitat of this new species is presumed to thin sand deposits directly on a hard substrate like rock-reefs. No living individual of this species was found in the sandy or muddy substrate around rock-reef although such substrates are inhabited by many species of clypeasteroids (Mooi 1990). At Amami-Oshima Island, six live sea urchins (NSMT E-10371, E-10372, E-10373, and E-10387) were collected from the thinly deposited sand of 1���2 cm thick on a rock-reef (Figs. 8C, D). In addition, one live specimen (NSMT E-10388) was found in the thin accumulation of sand of ca. 1 cm in thickness trapped by branched calcareous algae on a vertical surface of a rock-reef. In Tateyama, one live F. coffea (NSMT E-10376) was also collected from a 1���2 cm thin sand layer on a rock-reef. Two sea urchins (NSMT E-10374 and NSMT E-10375) were collected by dredging from sediments containing rubble and stones where rock-reef and sand bottom are interfingering. * These data were calculated by the authors based on the descriptions in each reference. Etymology. The species name is derived from the Latin ��� coffea ���, meaning ���coffee���, because the elliptical outline of test and the brownish color of living specimens resemble coffee beans. Remarks. F. coffea can be easily distinguished from all other extant species of Fibularia except F. ovulum by the mode of increase of the number of pores in the petal. The number of pores in the petal III of F. coffea is greater than in F. japonica, F. plateia, F. cribellum, and F. nutriens, reaching 20 in specimens of TL> 5.0 mm and 30 in specimens of TL> 7.5 mm (Fig. 4). On the other hand, the number of pores of petal III is up to 14 for F. japonica even in the specimens of TL> 7.5 mm (Fig. 4), 7 for the holotype of F. plateia (TL = 6.25 mm) (H.L. Clark 1928), 8 or less for F. cribellum (TL = 6.0��� 6.1 mm) (de Meijere 1903; Schultz 2009). Moreover, the number of pores of petal III of F. coffea continues to increase even after TL = 5.0 mm (Fig. 4). In contrast, in F. japonica, F. plateia, F. cribellum and F. nutriens, increase in pore pair number significantly slows down at TL = ca. 3.0 mm and almost stops after TL = ca. 4.0 mm (Fig. 4) (Gomez & Mooi 2015). F. ovulum is the most similar species to F. coffea. The number of pores of petal III of F. ovulum increases like in F. coffea below TL = 5.0 mm. However, in F. ovulum, that increase slows down above 5.0 mm TL, and only a maximum of 30 pores is reached in the largest specimens studied (9.45 mm TL). In F. coffea, that increase continues, up to 40 (Fig. 4). F. coffea can be more clearly distinguished from F. ovulum by its lower (TH/TL = 0.38���0.55 in F. coffea vs. 0.59���0.84 in F. ovulum) and less wide test (TW/TL = 0.67���0.86 in F. coffea vs. 0.77���0.92 in F. ovulum). The difference in these proportions of the test is consistent throughout their growth (slope value is 1.0 and 1.0 between TW and TL, and 1.1 and 1.1 between TH and TL in the allometry regression for F. coffea and F. ovulum), so it is more useful for identification between two species. Moreover, F. coffea can be distinguished by the larger peristome (SL/TL = 0.17���0.34 in F. coffea vs. 0.12���0.24 in F. ovulum; SW/TL = 0.14���0.30 in F. coffea vs. 0.11���0.23 in F. ovulum). In addition, the area around the peristome is depressed in F. coffea but inflated in F. ovulum. In life, F. coffea can be distinguished from F. ovulum by its coloration: the former has black pigment on the aboral surface (Fig. 7A, B), and the latter lacks this pigment and usually has purplish accessory tube feet (Mortensen 1948)., Published as part of Tanaka, Hayate, Wakabayashi, Kaori & Fujita, Toshihiko, 2019, A new species of Fibularia from Japanese waters with a redescription of F. japonica and F. ovulum (Echinodermata: Echinoidea: Clypeasteroida), pp. 241-260 in Zootaxa 4543 (2) on pages 244-251, DOI: 10.11646/zootaxa.4543.2.4, http://zenodo.org/record/2617781, {"references":["Shigei, M. (1982) A new species of the fibulariid sea-urchin, Fibularia japonica, from Japanese waters. Publications of the Seto Marine Biological Laboratory, 27, 11 - 16. https: // doi. org / 10.5134 / 176047","Schultz, H. (2005) Sea-Urchins, a Guide to Worldwide Shallow Water Species. Heinke & Peter Schultz Partner Scientific Publication, Hemdingen, 484 pp.","Gomez, C. & Mooi, R. (2015) New fossil and extant species of Fibularia illuminate evolution of the most highly miniaturized \" sand dollars \". Integrative and Comparative Biology, 55, 264. [E 264]","Mooi, R. (1990) Paedomorphosis, Aristotle's lantern, and the origin of the sand dollars (Echinodermata: Clypeasteroida). Paleobiology, 16 (1), 25 - 48. https: // doi. org / 10.1017 / S 0094837300009714","Clark, H. L. (1928) The sea-lilies, sea-stars, brittle stars and sea-urchins of the South Australian Museum. Records of The South Australian Museum, 3, 361 - 482.","Schultz, H. (2009) Sea-Urchins II, Worldwide Irregular Deep Water Species. Heinke & Peter Schultz Partner Scientific Publication, Hemdingen, 365 pp.","Mortensen, T. (1948) A Monograph of the Echinoidea. Fol. 3. Clypeastroida. Clypeastridae, Arachnoididae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp."]}

Fibularia ovulum Lamarck, 1816 [Japanese name: Maru-mame-uni (Shigei 1986)] Figs. 4, 6, 14; Tables 1, 2; Electronic Supplementary Table S1. Fibularia ovulum Lamarck, 1816: 16; Mortensen 1948: 208 ���210, figs, 102d, 104e, 106, 118, pl. 46, figs. 15���17, 21���24; A.M. Clark & Rowe 1971: 167 ���170, fig. 83���, pl. 25, figs. 6���8; Liao and A.M. Clark 1995: 382 ���383, fig. 224. Fibularia craniolaris ��� H.L. Clark 1914: 57. Material examined. 50 specimens, NSMT E-10393, dead tests, Kosyuku, Amami-Oshima Island, Kagoshima Prefecture, Japan (28��24���10���N, 129��28���04���E), beachcombing, coll. H. Tanaka, 22 Mar. 2015; 28 specimens, NSMT E-10392, dead tests, Vavvaru Island, Easter Beach, Lhaviyani Atoll, Maldives (5��25���4���N, 73��21���17���E), beachcombing, coll. A. Kroh and J. Herler, 7 Sept. 2014 ��� 14 Sept. 2014. Diagnosis. Test outline slightly elliptical when viewed from above; height high; oral surface inflated toward the peristome. Periproct from round to elliptical. Petaloid region large; number of pores in petal III continues to increase with the test growth, reaching over 20 at TL> 5 mm. Diameter of genital pores equal to or smaller than that of petaloid pores in mature individuals. Two hydropores in an irregularly-shaped groove. Description. The test is very small (TL = 2.05���9.45 mm) (Fig. 14), high (TH/TL = 0.59���0.84) (Fig. 14C), and elliptical when viewed from above (TW/TL = 0.77���0.92) (Figs. 14A, B). The test proportion hardly changes with the test growth (slope value is 1.0 between TW and TL, and 1.1 between TH and TL in the allometry regression; Table 1). The oral and aboral surfaces are inflated. There are no internal buttresses. Food grooves are absent (Fig. 14B). The ambulacra are almost the same width as the interambulacra (Figs. 14 D���F). The height of both ambulacral and interambulacral plates are lower than the width at the ambitus (Fig. 14F). The petaloid region is large (PL/TL = 0.49���0.69, PW/TW = 0.43���0.60). The ratio of petaloid region size to test size hardly change with the test growth (slope value is 1.0 between PL and TL, and 1.1 between PW and TL in the allometry regression). Each petal is composed of two almost parallel series of pore pairs lying oblique, and crossing the ambulacral plates (Fig. 14D). The number of pores of petal III, IV, and V is continuously increasing up to 30, 24, and 28, respectively, with the test growth (Fig. 4). The rate of increase slows down once a TL of more than 5.0 mm is reached. The pores become larger towards the distal tip of the petals. The peristome, situated at the center of the oral side, is small (SL/TL = 0.12���0.23, SW/TW = 0.11���0.23) and slightly elongated antero-posteriorly (Figs. 14B, E). The ratio of peristome size to test size becomes smaller as test grows (slope value is 0.7 between SL and TL, as well as between SW and TL in the allometry regression). Two buccal pores are situated in each ambulacrum at the edge of the peristome. Single sphaeridium is fully enclosed within the test, and in the sphaeridial chamber in each ambulacrum near the peristome. The round or roundish diamond shaped periproct is located halfway between the peristome and posterior margin of the test and is smaller than the peristome (AL/TL = 0.08���0.14, AW/TW = 0.09���0.15) (Figs. 14B, E). The ratio of peristome size to test size hardly changes as the test grows (slope value is 1.1 between AL and TL, and 0.9 between AW and TL in the allometry regression). The apical system is situated at the midpoint of the anterior-posterior axis on the aboral surface (Figs. 14A, D). It consists of four genital pores, five ocular pores in small ocular plates, and two hydropores in a deep, irregularlyshaped groove. The diameter of the genital pores (Color. The denuded test is whitish. Distribution. This species is recorded from East Africa and the Red Sea, the Maldives, Bay of Bengal, Xisha Island, the Philippines, the East Indies to the Gilbert Islands, and south of the Tokara Islands, from 0���385 m depth (Shigei 1981; Liao & A.M. Clark 1995; present study)., Published as part of Tanaka, Hayate, Wakabayashi, Kaori & Fujita, Toshihiko, 2019, A new species of Fibularia from Japanese waters with a redescription of F. japonica and F. ovulum (Echinodermata: Echinoidea: Clypeasteroida), pp. 241-260 in Zootaxa 4543 (2) on pages 257-259, DOI: 10.11646/zootaxa.4543.2.4, http://zenodo.org/record/2617781, {"references":["Lamarck, J. - B. (1816) Histoire Naturelle des Animaux sans Fertebres, presentant les caracteres generaux et particuliers de ces animaux, leur distribution, leur classes, leurs familles, leurs generes, et le citation des principales especes qui s'y rapportent; precedee d'une Introduction offrant la Determination des caracteres essentiells de l'animal, sa distinction du vegetal et des autres corps naturels, enfin, l'Exposition des Principes fondamentaux de la Zoologie. Tome Troisieme. Verdiere, Paris, 586 pp.","Shigei, M. (1986) The Sea Urchins of Sagami Bay. Maruzen, Tokyo, 173 pp.","Mortensen, T. (1948) A Monograph of the Echinoidea. Fol. 3. Clypeastroida. Clypeastridae, Arachnoididae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp.","Clark, A. M. & Rowe, F. W. E. (1971) Monograph of shallow-water Indo-west Pacific Echinoderms. Trustees of the British Museum (Natural History), London, 238 pp.","Liao, Y. & Clark, A. M. (1995) The Echinoderms of Southern China. Science Press, New York, 614 pp.","Clark, H. L. (1914) Hawaiian and other Pacific Echini. The Clypeasteridae, Arachnoididae, Laganidae, Fibulariidae, and Scutellidae. Memoirs of the Museum of Comparative Zoology, 46, 1 - 80.","Shigei, M. (1981) A study on the echinoid fauna of the East China Sea and the coastal waters of southern Korea, Kyushu, Ryukyu, and Taiwan. Publications of the Seto Marine Biological Laboratory, 26, 191 - 241. https: // doi. org / 10.5134 / 176013"]}

Fibularia japonica Shigei, 1982 [Japanese name: Nihon mame-uni (named by Shigei 1986)] Figs. 4, 6, 9���13; Tables 1, 2; Electronic Supplementary Table S1. Fibularia sp. nov.��� Shigei 1981: 202 (probably part). Fibularia japonica Shigei, 1982: 11 ���16, figs. 1���48 (part). Fibularia japonica Shigei 1986: 116 ���117, Pl. 92, figs. 4���9; Schultz 2005: 321, fig. 602; Shigei 2006: 318 ���319; Yoshigou 2009: 35 ���36, pl. 1, fig. 4; Gomez & Mooi 2015: poster presentation. Material examined. Holotype: 1 specimen, UMUTZ-Ecn-SG10-16T, denuded test, off Misaki Marine Biological Station, Sagami Bay, sublittoral zone, coll. K. Aoki, J. Deguchi, T. Sekimoto, H. Suzuki, and M. Shigei, 1926��� 1978. Paratypes: 1 specimen, UMUTZ-Ecn-SG10-18T, dead test, 3 km off Futamachiya, Sagami Bay (35��08.5���N, 139��35.0���E), depth 45 m, coll. H. Suzuki, M. Sekimoto, K. Shimazaki, and M. Shigei, 4 Jul. 1979; 6 specimens, UMUTZ-Ecn-SG10-17T No. 2���6, 8, dead tests, off Misaki Marine Biological Station, Sagami Bay, sublittoral zone, coll. K. Aoki, J. Deguchi, T. Sekimoto, H. Suzuki, and M. Shigei, 1926���1978; 1 specimen, UMUTZ-Ecn-SG10-19T, dead test, Tomioka, Amakusa group, Kumamoto Prefecture, Japan, depth 30 m, coll. T. Kikuchi, 14 Feb. 1963; 26 specimens, UMUTZ-Ecn-SG10-20T, dead tests, Sagami Bay, sublittoral zone; 3 specimens, UMUTZ-Ecn-SG10-21T, dead tests, Sagami Bay, sublittoral zone; 1 specimen, UMUTZ-Ecn-SG10-22T, dead test, Suruga Bay, sublittoral zone, coll. Y. Okada. Non-type specimens: 45 specimens, NSMT E-10389, dead tests, Sagami Bay (35��09���18���N, 139��35���12���E), depth 74.1 m, dredging, coll. H. Tanaka, 23 Jan. 2014; 6 specimens, NSMT E-10390, whole, denuded, SEM stub, Sagami Bay (35��09���18���N, 139��35���13���E), depth 74.1 m, dredging, coll. H. Tanaka, 23 Jan. 2014; 15 specimens, NSMT E-10391, dead tests, Sagami Bay (35��08���09���N, 139��34���47���E), depth 87.5���88.6 m, dredging, coll. S. Teruya, 14 Mar. 2012. One paratype specimen (UMUTZ-Ecn-SG10-17T No. 7) was identified as F. coffea in this study. GenBank accession number. LC388936 (non-type specimen: NSMT E-10390, Sagami Bay, Japan) Emended diagnosis. Test outline elliptical when viewed from above; height low; oral surface not depressed. Periproct outline round to oblong. Petaloid region large; number of pores in petal III up to 14 even in the specimens of TL> 7.5 mm. Diameter of genital pores equal to or larger than that of petaloid pores in mature individuals. Two hydropores opening in an irregularly-shaped groove. Black pigments not forming symmetric pentaradial in living animals. Spatula-like primary spines around periproct. Description. The test is very small (TL = 2.19���9.70 mm) (Fig. 9), flattened (TH/TL = 0.49���0.80) (Fig. 9C), and elliptical when viewed from above (TW/TL = 0.67���0.83) (Figs. 9A, B). The test proportion hardly changes with the test growth (slope value is 1.0 between TW and TL, as well as between TH and TL in the allometry regression; Table 1). The oral surface is flattened. The aboral surface is slightly arched convex. There are no internal buttresses. Food grooves are absent (Fig. 9B). The ambulacra are almost the same width as the interambulacra (Fig. 9 G���I). The height of both ambulacral and interambulacral plates are lower than the width at the ambitus (Fig. 9I). The petaloid region is large (PL/TL = 0.33���0.66, PW/TW = 0.28���0.51). The ratio of petaloid size to test size increases with the test grows (slope value is 1.1 between PL and TL, as well as between PW and TL in the allometry regression). Each petal is composed of two almost parallel series of pore pairs lying oblique, and crossing the ambulacral plates (Fig. 9G). The number of pores of petal III, IV, and V increases up to 14, 12, and 14, respectively, before reaching TL = ca. 3 mm, and hardly increases after that size has been reached (Fig. 4). The pores become larger towards the distal tip of the petals. The peristome, situated at the anterior-posterior midpoint of the oral side, is small (SL/TL = 0.13���0.27, SW/ TW = 0.12���0.25) and slightly elongated antero-posteriorly (Figs. 9B, E, H). The ratio of peristome size to test size becomes smaller with the test grows (slope value is 0.7 between SL and TL, as well as between SW and TL in the allometry regression). Two buccal pores are situated in each ambulacrum at the edge of the peristome. Single sphaeridium is fully enclosed within the test, and in the sphaeridial chamber in each ambulacrum near the peristome. The round to oblong shaped periproct is located halfway between the peristome and posterior margin of the test and is smaller than the peristome (AL/TL = 0.09���0.15, AW/TW = 0.09���0.16) (Figs. 9B, E, H), and covered by 4���6 (usually 5) naked radiating periproctal plates. The ratio of periproct size to test size becomes smaller with the test grows (slope value is 1.0 between AL and TL, and 0.8 between AW and TL in the allometry regression). The apical system is situated slightly anteriorly on the aboral surface (Figs. 9A, D, G). It consists of four genital pores, five ocular pores in small ocular plates, and two hydropores in a deep, irregularly-shaped groove (Figs. 10A, B). The diameter of the genital pores (F. coffea. Each valve has 20���25 teeth, and each tooth has 1���2 denticles. The proximal end of the handle on the largest valve is inserted into a depression at the distal end of the pedicellarial stalk (Fig. 11 Dii). The tridentate pedicellariae (Fig. 11E) occur only around the peristome and periproct. These pedicellariae consist of a head with three slender valves (Fig. 11 Ei), short neck, and stem. The valves possess ca. 11���20 teeth on the edge and without denticles (Fig. 11E). Some valves have ca. 1���4 teeth in the inner area (Fig. 11 Eiv). The accessory tube feet lack a calcareous disk or spicules. Color. The color is white to yellow in life (Fig. 13A) but changes to green when preserved in ethanol. Black pigments are distributed in speckles over the entire test (Fig. 13B) and remain even after preservation. The denuded test is whitish. Distribution. F. japonica has so far been recorded in Japanese waters, from Sagami Bay to Kyushu; 30���100 m in depth (Shigei 1986; present study). Schultz (2009) reported this species from also the Philippines. Habitat. Live specimens collected from sandy bottoms by dredging suggest that F. japonica inhabits sandy substrate. Remarks. In the original description of F. japonica, Shigei (1982) noted that ���each pore series of petals consists of only 2���3 pore pairs in adult specimens, while 4���5 in young specimens [sic].��� His statement that the number of pores decreases with growth is erroneous because it always increases with growth in clypeasteroids (Zachos 2015) and all other echinoids. As a result of the re-examination of the type specimens of F. japonica, one of the paratypes (UMUTZ-Ecn-SG10-17T No. 7) showed the morphology of F. coffea. This paratype specimen is 4.11 mm in TL, has five pore pairs in each pore series (total 20 pores) in petal III (x-mark in Fig. 4). We confirmed that the number of pores in petal III does not exceed 15 in any of the paratypes of F. japonica. We assumed that this relatively small specimen of F. coffea, mixed in the type specimens of F. japonica, caused his strange statement in the original description (Shigei 1982). True F. japonica is a species with less than 4 pore pairs in each pore series (a total of max. 16 pores per petal). Shigei (1982) described a valve of ophicephalous pedicellariae that had no trace of intertwined loops (Shigei 1982: fig. 48). However, we observed that the valves of ophicephalous pedicellariae of F. japonica are characterized by well-developed intertwined loops (Fig. 11 Di). Therefore, the valves of an ophicephalous pedicellaria do not separate from each other even after the soft tissues are removed by bleaching. The valve suggested to be from an ophicephalous pedicellaria illustrated by Shigei (1982: fig. 48) is more similar to that of a tridentate pedicellaria (Fig. 11E), suggesting that Shigei mislabeled the valve in his illustration., Published as part of Tanaka, Hayate, Wakabayashi, Kaori & Fujita, Toshihiko, 2019, A new species of Fibularia from Japanese waters with a redescription of F. japonica and F. ovulum (Echinodermata: Echinoidea: Clypeasteroida), pp. 241-260 in Zootaxa 4543 (2) on pages 252-257, DOI: 10.11646/zootaxa.4543.2.4, http://zenodo.org/record/2617781, {"references":["Shigei, M. (1982) A new species of the fibulariid sea-urchin, Fibularia japonica, from Japanese waters. Publications of the Seto Marine Biological Laboratory, 27, 11 - 16. https: // doi. org / 10.5134 / 176047","Shigei, M. (1986) The Sea Urchins of Sagami Bay. Maruzen, Tokyo, 173 pp.","Shigei, M. (1981) A study on the echinoid fauna of the East China Sea and the coastal waters of southern Korea, Kyushu, Ryukyu, and Taiwan. Publications of the Seto Marine Biological Laboratory, 26, 191 - 241. https: // doi. org / 10.5134 / 176013","Schultz, H. (2005) Sea-Urchins, a Guide to Worldwide Shallow Water Species. Heinke & Peter Schultz Partner Scientific Publication, Hemdingen, 484 pp.","Shigei, M. (2006) A systematical study on the echinoids (Echinodermata, Echinoidea) from the Sagami Sea. Memoirs of the National Science Museum, 41, 305 - 327.","Yoshigou, H. (2009) The Fibulariidae (Echinoidea: Clypeasteroida) collected from sea sand of factory materials (In Japanese). Hibakagaku, 231, 33 - 40.","Gomez, C. & Mooi, R. (2015) New fossil and extant species of Fibularia illuminate evolution of the most highly miniaturized \" sand dollars \". Integrative and Comparative Biology, 55, 264. [E 264]","Schultz, H. (2009) Sea-Urchins II, Worldwide Irregular Deep Water Species. Heinke & Peter Schultz Partner Scientific Publication, Hemdingen, 365 pp.","Zachos, L. G. (2015) Holistic morphometric analysis of growth of the sand dollar Echinarachnius parma (Echinodermata: Echinoidea: Clypeasteroida). Zootaxa, 4052 (2), 151 - 179. https: // doi. org / 10.11646 / zootaxa. 4052.2.1"]}

A new species, Fibularia coffea sp. nov., occurs from shallow waters in Japan. This new species is distinguished from the other species of Fibularia by the following characters: test height is low, oral surface is slightly depressed toward the peristome, number of pores of petal III continues to increase with the test growth, reaching over 30 at TL > 7.5 mm, and black pigments form symmetric pentaradial on aboral surface in living animals. Two further Japanese species, Fibularia japonica and F. ovulum, are redescribed based on the type specimens (F. japonica) and additional specimens (F. ovulum), respectively. A tabular key to the extant species of Fibularia is also provided. A partial fragment of the mitochondrial gene cytochrome oxidase subunit I (COI) of the type specimens of F. coffea sp. nov. and the additional specimen of F. japonica was sequenced for barcoding in future works.

Fibularia coffea sp. nov. [New Japanese name: Kohi-mame-uni] Figs. 3–8; Tables 1, 2; Electronic Supplementary Table S1. Non Fibularia japonica Shigei 1982: 11 –16, figs. 1–48 (part). Non Fibularia plateia Schultz 2005: 321, fig. 603. Fibularia n. sp. “bean” Gomes & Mooi 2015: poster presentation. Material examined. Holotype: NSMT E-10371, whole, with spines, Ayamaru Cape, Amami-Oshima Island, Kagoshima Prefecture, Japan (28°28′26″N, 129°43′09″E), depth 1 m, snorkeling, coll. H. Tanaka, 23 Mar. 2015. Paratypes: 1 specimen, NSMT E-10372, whole, denuded, SEM stub, Tsuchi-Hama, Amami-Oshima Island, Kagoshima Prefecture, Japan (28°24′36″N, 129°40′47″E), depth 2 m, snorkeling, coll. H. Tanaka and L. Sakamoto, 20 Mar. 2015; 2 specimens, NSMT E-10373, whole, denuded, SEM stub, Kunigami, Nishinoomote city, Tanegashima Island, Kagoshima Prefecture, Japan (30°48′19″N, 131°01′24″E), depth 4–5 m, scuba diving, coll. H. Yamasaki and R. Yoshida, 1 Mar. 2014; 1 specimen, NSMT E-10374, whole, denuded, SEM stub, Tateyama Bay, Tateyama city, Chiba Prefecture, Japan (34°58′56″N, 139°48′43″E), depth 6.9–8.1 m, dredging, coll. Y. Yoshida, 14 Jan. 2015; 1 specimen, NSMT E-10375, whole, denuded, SEM stub, Tateyama Bay, Tateyama city, Chiba Prefecture, Japan (34°59′05″N, 139°48′57″E), depth 9–12 m, dredging, coll. Y. Yoshida, 12 May. 2014; 1 specimen, NSMT E-10376, whole, denuded, SEM stub, Tateyama Bay, Tateyama city, Chiba Prefecture, Japan (34°58′40″N, 139°46′05″E), depth 5 m, scuba diving, coll. K. Kosoba, 30 Aug. 2015; 29 specimens, NSMT E-10377, dead tests, Tomioka, Amakusa group, Kumamoto Prefecture, Japan, dredging, coll. H. Tanaka, 21 Mar. 2015; 1 specimen, NSMT E-10378, dead test, Ayamaru Cape, Amami-Oshima Island, Kagoshima Prefecture, Japan (28°28′30″N, 129°43′05″E), beachcombing, coll. H. Tanaka, 23 Mar. 2015; 7 specimens, NSMT E-10379, dead tests, Ayamaru Cape, Amami-Oshima Island, Kagoshima Prefecture, Japan (28°28′30″N, 129°43′05″E), beachcombing, coll. H. Arima, 11 Dec. 2013 – 27 Mar. 2014; 7 specimens, NSMT E-10380, dead tests, Kosyuku, Amami-Oshima Island, Kagoshima Prefecture, Japan (28°24′10″N, 129°28′04″E), beachcombing, coll. H. Arima, 13 Dec. 2013 – 22 Mar. 2015; 2 specimens, NSMT E-10381, dead tests, Kosyuku, Amami-Oshima Island, Kagoshima Prefecture, Japan (28°24′10″N, 129°28′04″E), beachcombing, coll. H. Tanaka, 22 Mar. 2015; 5 specimens, NSMT E-10382, dead tests, Tsuchi-Hama, Amami-Oshima Island, Kagoshima Prefecture, Japan (28°24′39″N, 129°40′37″E), beachcombing, coll. H. Tanaka, 15 Jul. 2014; 3 specimens, NSMT E-10383, dead tests, Tsuyazaki, Fukutsu city, Fukuoka Prefecture, Japan (33°45′59″N, 130°23′03″E), beachcombing, coll. K. Wakabayashi, 17 Jun. 2005; 10 specimens, NSMT E-10384, dead test, Wada Beach, Oi county, Fukui Prefecture, Japan (35°29′41″N, 135°34′32″E), beachcombing, coll. H. Tanaka, 3 Jan. 2014; 4 specimens, NSMT E-10385, dead tests, Yoan Beach, Amami-Oshima Island, Kagoshima Prefecture, Japan (28°24′12″N, 129°38′35″E), beachcombing, coll. H. Tanaka and L. Sakamoto, 31 Jan. 2016; 48 specimens, NSMT E-10386, dead tests, Zushi Beach, Zushi city, Kanagawa Prefecture, Japan (35°17′18″N, 139°34′25″E), beachcombing, coll. H. Tanaka, 21 Jul. 2012 – 2 Feb. 2014; 2 specimens, NSMT E-10387, whole, denuded, Tsuchi-Hama, Amami-Oshima Island, Kagoshima Prefecture, Japan (28°24′36″N, 129°40′47″E), intertidal, snorkeling, coll. H. Tanaka and L. Sakamoto, 24 Mar. 2015; 1 specimen, NSMT E-10388, whole, denuded, Tsuchi-Hama, Amami-Oshima Island, Kagoshima Prefecture, Japan (28°24′36″N, 129°40′47″E), intertidal, snorkeling, coll. H. Tanaka and L. Sakamoto, 22 Oct. 2016. One specimen, UMUTZ-Ecn-SG10-17T No. 7 (as a paratype of Fibularia japonica), dead test, off Misaki Marine Biological Station, Sagami Bay, sublittoral zone, coll. K. Aoki, J. Deguchi, T. Sekimoto, H. Suzuki, and M. Shigei, 1926–1978. GenBank accession number. LC388935 (holotype: NSMT E-10371, Amami-Oshima Island, Kagoshima Prefecture, Japan), LC388934 (paratype: NSMT E-10374, Tateyama Bay, Tateyama city, Chiba Prefecture, Japan). Diagnosis. Test outline elliptical when viewed from above; height low; oral surface slightly depressed toward the peristome. Periproct outline round square shaped. Petaloid region large; number of pores of petal III continues to increase with the test growth, reaching over 20 at TL> 5 mm. Diameter of genital pores equal to or smaller than that of petaloid pores in mature individuals. Two hydropores opening in an irregularly-shaped groove. Black pigments on each aboral perradius, forming symmetric pentaradial in living animals. Description. The test is very small (TL = 1.53–9.73 mm) (Fig. 3), flattened (TH/TL = 0.38–0.55) (Fig. 3C), elliptical when viewed from above (TW/TL = 0.67–0.86), and truncated posteriorly (Figs. 3A, B). The test proportion hardly changes with the test growth (slope value is 1.0 between TW and TL, and 1.1 between TH and TL in the allometry regression, Table 1). The oral surface is slightly depressed around the peristome. The aboral surface is slightly arched convex. There are no internal buttresses. Food grooves are absent (Fig. 3B). The ambulacra are almost the same width as the interambulacra (Figs. 3 D–F). The height of both ambulacral and interambulacral plates are lower than the width at the ambitus (Fig. 3F). The petaloid region is large (PL/TL = 0.46–0.75, PW/TL = 0.33–0.61). The ratio of petaloid region size to test size becomes larger with test growth (slope value is 1.3 between PL and TL, as well as between PW and TL in the allometry regression). Each petal is composed of almost parallel series of pore pairs lying oblique, and crossing the ambulacral plates (Fig. 3D). The number of pores of petal III, IV, and V is continuously increasing up to 40, 28 and 36, respectively, with the test growth (Fig. 4). The pores become larger towards the distal tip of the petals. The peristome, situated at the center of the oral side, is small (SL/TL = 0.17–0.34, SW/TW = 0.14–0.30) and slightly elongated antero-posteriorly (Figs. 3B, E). The ratio of peristome size to test size becomes smaller with the test grows (slope value is 0.8 between SL and TL, as well as between SW and TL in the allometry regression). The peristomial membrane lacks spine and pedicellariae. Two buccal pores are situated in each ambulacrum at the edge of the peristome. Single sphaeridium is fully enclosed within the test, and in the sphaeridial chamber in each ambulacrum near the peristome. The rounded square shaped periproct is located halfway between the peristome and posterior margin of the test and is smaller than the peristome (AL/TL = 0.08–0.15, AW/TW = 0.08–0.16) (Figs. 3B, E) and covered by 4–6 (mainly 5) naked radiating periproctal plates. The ratio of periproct size to test size hardly changes with the test grows (slope value is 1.0 between AL and TL, and 0.9 between AW and TL in the allometry regression). The apical system is situated slightly anterior on the aboral surface (Figs. 3A, D). It consists of four genital pores, five ocular pores in small ocular plates, and two hydropores in a deep, irregularly-shaped groove (Fig. 5A). The diameter of the genital pores ( The primary tubercles are hemispherical, crenulate, and perforate (Figs. 5C, D). The diameters of oral and aboral primary tubercles are almost equal, ca. 150–200 µm. Their mamelons are constricted at the base (Fig. 5D). The miliary tubercles are hemispherical, poorly to non-crenulate, and indistinctly to non-perforate (Figs. 5C, D). They scattered around the primary tubercles. The diameters of oral and aboral miliary tubercles are almost equal, ca. 50–60 µm. The glassy tubercles occur between primary and miliary tubercles (Fig. 5C, D). The primary spines are ca. 350–450 µm in length (Fig. 7 Ai). The number of wedges in a primary spine is 8–10, and each wedge has many small granules and a series of distinct denticles (Figs. 7 Aii, Aiii). The tip of each primary spine is somewhat truncated (Fig. 7 Aii). The distal end of primary spines around the peristome is slightly curved towards the peristome, and their shafts are somewhat broadened and flattened (Fig. 7B). The miliary spines are ca. 250–350 µm in length (Fig. 7 Ci). Each miliary spine bears a distal crown (Fig. 7 Cii), and the number of wedges in a miliary spine is six for all examined spines. Each wedge has many small granules along its outer surface (Fig. 7 Ciii). Two types of pedicellariae, ophicephalous and tridentate, are present (Figs. 7D, E). These two types pedicellariae occur on small tubercles similar to those of miliary spines. The ophicephalous pedicellariae (Fig. 7D) are numerous and occur over the entire test surface. The head is ca. 80–100 µm in length (Fig. 7 Dii), and consists of three valves that differ from each other in size and shape. The largest valve has a large, bilaterally symmetric handle. The medium-sized valve has a left-right asymmetric handle. The smallest valve has a small, bilaterally symmetric handle. The left and right ends of each valve have a finger-like structure near the hinge (Fig. 7 Dvi), making an “intertwined loop” (Mooi 1990). The valves are connected to each other by these intertwined loops (Figs. 7 Dii, Dv–Dvii). Each valve has 20–25 teeth, and each tooth has 1–2 denticles (Fig. 7 Div). The proximal end of the handle on the largest valve is inserted into a depression at the distal end of the pedicellarial stalk (Figs. 7 Di– Diii). The tridentate pedicellariae (Fig. 7E) occur only around the peristome and periproct. These pedicellariae consist of a head with three slender valves (Fig. 7 Ei), short neck, and stem. The valves possess ca. 11–15 teeth on the edge and without denticles (Figs. 7E). Some valves have ca. 1–4 teeth in the inner area (Figs. 7 Ei, Eii, Eiv). The accessory tube feet lack a calcareous disk or spicules. Color. The color is yellow to brown in life (Figs. 8A, B) but changes to green when preserved in ethanol. Black pigments are apparent in each ambulacrum along the pore pair columns, forming a pentaradially symmetric pattern on the aboral surface (Figs. 8A, B). The black pigments remain even after preservation in ethanol. The denuded test is whitish. Distribution. This species is recorded in Japanese waters from Sagami Bay to the Amami-Oshima Islands; 1– 12 m in depth (present study). Schultz (2005) recorded F. plateia from Queensland, Australia but the figured specimen seems to be F. coffea from the flattened test and the large number of pores of petals (the number of pores of petal III, IV and V are 32, 22 and 32, respectively). In addition, Gomez & Mooi (2015) recorded an unknown species Fibularia n. sp. “bean” from the Philippines and Micronesia that seems to be the same as F. coffea from the flattened test and the large number of pores of petals (the number of pores of petal III, IV and V are 34, 26 and 33, respectively; counted from the sketch image). Therefore, this species is considered to be widely distributed across the Indo-Pacific. Habitat. The micro-habitat of this new species is presumed to thin sand deposits directly on a hard substrate like rock-reefs. No living individual of this species was found in the sandy or muddy substrate around rock-reef although such substrates are inhabited by many species of clypeasteroids (Mooi 1990). At Amami-Oshima Island, six live sea urchins (NSMT E-10371, E-10372, E-10373, and E-10387) were collected from the thinly deposited sand of 1–2 cm thick on a rock-reef (Figs. 8C, D). In addition, one live specimen (NSMT E-10388) was found in the thin accumulation of sand of ca. 1 cm in thickness trapped by branched calcareous algae on a vertical surface of a rock-reef. In Tateyama, one live F. coffea (NSMT E-10376) was also collected from a 1–2 cm thin sand layer on a rock-reef. Two sea urchins (NSMT E-10374 and NSMT E-10375) were collected by dredging from sediments containing rubble and stones where rock-reef and sand bottom are interfingering. * These data were calculated by the authors based on the descriptions in each reference. Etymology. The species name is derived from the Latin “ coffea ”, meaning “coffee”, because the elliptical outline of test and the brownish color of living specimens resemble coffee beans. Remarks. F. coffea can be easily distinguished from all other extant species of Fibularia except F. ovulum by the mode of increase of the number of pores in the petal. The number of pores in the petal III of F. coffea is greater than in F. japonica, F. plateia, F. cribellum, and F. nutriens, reaching 20 in specimens of TL> 5.0 mm and 30 in specimens of TL> 7.5 mm (Fig. 4). On the other hand, the number of pores of petal III is up to 14 for F. japonica even in the specimens of TL> 7.5 mm (Fig. 4), 7 for the holotype of F. plateia (TL = 6.25 mm) (H.L. Clark 1928), 8 or less for F. cribellum (TL = 6.0– 6.1 mm) (de Meijere 1903; Schultz 2009). Moreover, the number of pores of petal III of F. coffea continues to increase even after TL = 5.0 mm (Fig. 4). In contrast, in F. japonica, F. plateia, F. cribellum and F. nutriens, increase in pore pair number significantly slows down at TL = ca. 3.0 mm and almost stops after TL = ca. 4.0 mm (Fig. 4) (Gomez & Mooi 2015). F. ovulum is the most similar species to F. coffea. The number of pores of petal III of F. ovulum increases like in F. coffea below TL = 5.0 mm. However, in F. ovulum, that increase slows down above 5.0 mm TL, and only a maximum of 30 pores is reached in the largest specimens studied (9.45 mm TL). In F. coffea, that increase continues, up to 40 (Fig. 4). F. coffea can be more clearly distinguished from F. ovulum by its lower (TH/TL = 0.38–0.55 in F. coffea vs. 0.59–0.84 in F. ovulum) and less wide test (TW/TL = 0.67–0.86 in F. coffea vs. 0.77–0.92 in F. ovulum). The difference in these proportions of the test is consistent throughout their growth (slope value is 1.0 and 1.0 between TW and TL, and 1.1 and 1.1 between TH and TL in the allometry regression for F. coffea and F. ovulum), so it is more useful for identification between two species. Moreover, F. coffea can be distinguished by the larger peristome (SL/TL = 0.17–0.34 in F. coffea vs. 0.12–0.24 in F. ovulum; SW/TL = 0.14–0.30 in F. coffea vs. 0.11–0.23 in F. ovulum). In addition, the area around the peristome is depressed in F. coffea but inflated in F. ovulum. In life, F. coffea can be distinguished from F. ovulum by its coloration: the former has black pigment on the aboral surface (Fig. 7A, B), and the latter lacks this pigment and usually has purplish accessory tube feet (Mortensen 1948).

Family Astriclypeidae Stefanini, 1912 Genus Echinodiscus Leske, 1778, Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on page 57, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460

The earliest information on Sri Lankan echinoid species belonging to the Irregularia dates back to Alexander Agassiz (1872). However, the current knowledge of diversity and distribution of irregular echinoids from Sri Lanka (formerly Ceylon) remains sparse. In addition, there are no recent taxonomic studies or biodiversity surveys for irregular echinoids, and no illustrated field-guides or reference collections are available specifically for Sri Lanka. To address these gaps, left open for more than 100 years since the work of Clark (1915), this study was conducted as an island-wide systematic sampling survey. Over 200 echinoid specimens were collected from 24 localities in Sri Lankan coastal waters by snorkelling and SCUBA diving down to 33 m depth. The collected specimens were identified using existing keys and authenticated with specimens available at the Natural History Museum in Vienna, Austria. The present study records 22 irregular echinoid species belonging to 15 genera and nine families in four orders. Among the identified irregular echinoids, six species, Echinocyamus megapetalus H.L. Clark, 1914, Fibularia ovulum Lamarck, 1816, Fibulariella angulipora Mortensen, 1948, Echinodiscus cf. truncatus L. Agassiz, 1841, Peronella oblonga Mortensen, 1948 and Brissus cf. agassizii Döderlein, 1885, are new records for Sri Lanka. Four unidentified, possibly new species belonging to the genera Fibularia, Jacksonaster and Metalia are reported, but kept in open nomenclature until more material becomes available. At present, the diversity of irregular echinoids from Sri Lanka now stands at 37 species representing 11 families in four orders. A dichotomous key is presented for all Sri Lankan irregular echinoids.

Family Astriclypeidae Stefanini, 1912 Genus Echinodiscus Leske, 1778

Genus Sculpsitechinus Stara & Sanciu, 2014 1778 Echinodiscus auritus Leske: p. 138 1948b Echinodiscus auritus Leske. —Mortensen: p. 400–403; pl. 56: figs. 2, 3; pl. 57: fig. 7; pl. 71: figs. 1–5, 10–15, 17, 22. 2014 Sculpsitechinus auritus (Leske, 1778) —Stara & Sanciu: p. 348–351; pl. 21: figs. 1–7. Material studied. Five denuded tests: WUSL /EI/57, from Silavathurai, Mannar; WUSL /EI/59, from Kirinda; WUSL /EI/128, EI/129, and EI/130, from Mandathiv, Jaffna. Description. Shape and size —Tests large, 62.6–111.7 mm TL; outline of anterior of test rounded, posterior end truncated with two deep, open lunules; greatest width lies about half-way between apex and posterior edge; TW 98–104% TL; test depressed in lateral view, c. 8% TL in height; margin thin and sharp, anterior margin thicker than posterior; oral side flat, food grooves bifurcating, closely following lunules; oral ambulacra with much more densely packed and smaller tubercles than interambulacra. Apical system —Monobasal, with four small, circular gonopores; subcentral, 44–51% TL from anterior margin. Ambulacra —Petaloid region medium-sized, length 44–47% TL; petals closed; petal III slightly longer (length 24% TL) and broader (10% TL in width) than others; anterior paired petals (mean 20.4% TL) and posterior paired petals (mean 19.8% TL) similar in length, 20% TL and width, c. 9% TL; interporiferous zones of petals small and about equal in width, c. 3% TL; poriferous zones and interporiferous zones about equal in width; pores are elongate anisopores with sharp furrows that deepen towards outer pore; 7–15 primary tubercles on ridges between furrows; food groves bifurcating close to peristome, with well-developed lateral branches encroaching upon interambulacra; 2 to 3 pairs of plates between ambulacral petals and lunules. Lunules —Open, slit-like; of variable length; width c. 5% TL; angle between lunules 57° to 60°; lunules surrounded by 3–4 plates per column on oral and 4–5 plates per column on aboral side. Interambulacra —Disjunction of oral interambulacra 1 to 4 variable, sometimes all in contact, in other specimens all are disjunct; interambulacrum 5 always disjunct. Five post-basicoronal plates in oral interambulacrum 5, two in column a and three in column b. Peristome —Round to subpentagonal, small, diameter 3–4% TL; 44–46% TL from anterior margin. Periproct —Small, diameter c. 2% TL; situated close to posterior margin of test, 14–19% TL from posterior edge; distance between periproct and peristome 29–35% TL. Geographic range. Indo-West Pacific, from Mauritius (De Loriol 1883), East Africa & Madagascar (Brown 1910a), Red Sea (Agassiz 18841), South East Arabia (Mortensen 1948c), Persian Gulf (Mortensen 1940b), India & Pakistan (Clark 1925a), Sri Lanka (Clark 1915; Herdman et al. 1904), Bay of Bengal (Koehler 1922), East Indies (de Meijere 1904), North Australia (Clark 1938) to Philippine Islands (Agassiz 1872), China Sea & South Japan (Mortensen 1948b). Bathymetric range. Tidal zone to 50 m (Mortensen 1948b). Observed occurrence in Sri Lanka. Specimens were collected from sandy and muddy bottoms at three Sri Lankan localities: Mandathiv on the northern coast, Silavathurai on the north-western coast, and Kirinda on the southern coast at 1, 9, and 27 m respectively (Fig. 61). Bathymetric range of this species in Sri Lanka is 1–29 m. This species was first recorded in Sri Lanka by Döderlein (1888). Remarks. In contrast to Stara & Sanciu’s (2014) description, Sri Lankan S. auritus has two pairs of ambulacral plates between the petal tips and lunules. The periproct is close to the posterior margin (mean=16% TL). However, in Stara & Sanciu’s (2014) examples, there are four to five pairs of plates between the petal tips and lunules, and the periproct is farther from the posterior margin (mean=21% TL). Specimen WUSL/EI/58, collected at a depth of 12 m from Palagala reef (Fig. 1) on the western coast of Sri Lanka, differs from the material described above and may represent a different species (or a hybrid). In contrast to typical S. auritus it has 1) a distinct anterior notch, 2) wide interporiferous zones (4–5% TL), and 3) a markedly structured oral side like in fossil Amphiope (Fig. 60B, D; in contrast to the rather flattened oral surface of typical S. auritus). Specimen WUSL/E1/58 is provisionally identified as Sculpsitechinus sp. until more material of Sculpsitechinus from Sri Lanka becomes available and the degree of intraspecific variation of the Sri Lankan population is better understood. S. auritus can be easily distinguished from other astriclypeids occurring in Sri Lanka (E. bisperforatus and E. cf. truncatus) by its open lunules., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on pages 67-70, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Stara, P. & Sanciu, L. (2014) Analysis of some astriclypeids (Echinoidea: Clypeasteroida). Biodiversity Journal, 5, 291 - 358.","De Loriol, P. (1883) Catalogue raisonne des Echinodermes recueillis par MV de Robillard a l'ile Maurice. Memoires de la Societe de physique et d'histoire naturelle de Geneve, 28, 1 - 64.","Brown, R. N. R. (1910 a) Echinoidea from the Kerimba Archipelago, Portuguese East Africa (Mozambique). Edinburgh Proceedings of the Royal Physical Society, 18, 36 - 44.","Mortensen, T. (1948 c) Report on the Echinoidea of the Murray Expedition. II [Irregular Echinoidea]. Scientific Reports on the John Murray Expedition, 9, 1 - 15.","Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Clark, H. L. (1925 a) A Catalogue of the Recent Sea-Urchins (Echinoidea) in the Collection of the British Museum (Natural History). Oxford University Press, London, 250 pp.","Clark, H. L. (1915) The echinoderms of Ceylon other than holothurians. Spolia Zeylanica, 10, 83 - 102.","Herdman, W. A., Herdman, J. B. & Bell, F. J. (1904) Report on the Echinoderma collected by Professor Herdman, at Ceylon, in 1902. In: Herdman, W. A. (Ed.), Report to the government of Ceylon on the pearl oyster fisheries of the Gulf of Mannar. The Royal Society, London, pp. 137 - 150.","Koehler, R. (1922) Echinoderma of the Indian Museum, Part 9, An account of the Echinoidea. II. Clypeastrides et Cassidulides. Trustees of the Indian Museum, Calcutta, 161 pp.","de Meijere, J. C. H. (1904) Die Echinoidea der Siboga-Expedition. Siboga Expeditie, 43, 1 - 252.","Clark, H. L. (1938) Echinoderms from Australia. Memoirs of the Museum of Comparative Zoology of the Harvard College, 55, 1 - 596.","Agassiz, A. (1872 - 1874) Revision of the Echini. Illustrated Catalogue of the Museum of Comparative Zoology at Harvard College, 7 (Pt. 1 - 2, 3 & 4) i-xii + 1 - 378, pls. 1 - 49 (1872), 379 - 628 + 1, pls. 50 - 77 (1873) & 629 - 762, pls. 78 - 94 (1874).","Mortensen, T. (1948 b) A Monograph of the Echinoidea IV. 2. Clypeasteroida. Clypeasteridae, Arachnoidae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp.","Doderlein, L. (1888) Echinodermen von Ceylon. Bericht uber die von den Herren Dres Sarasin gesammelten Asteroidea, Ophiuroidea und Echinoidea. Zoologische Jahrbucher, Abteilung fur Systematik, Geographie und Biologie der Tiere, 3, 821 - 846. https: // doi. org / 10.5962 / bhl. part. 1933"]}

Echinodiscus cf. truncatus L. Agassiz, 1841 Figures 51B, 55–58 1841 Lobophora truncata L. Agassiz: p. 66–67; pl. 11: figs. 11–16. 1948b Echinodiscus bisperforatus truncatus (L. Agassiz).—Mortensen: p. 410–411. Material studied. Five denuded specimens: WUSL /EI/60, EI/61, and EI/63, from Periyapaduwa and WUSL /EI/62 and EI/64, from Negombo 2, Sri Lanka. Description. Shape and size —Test small, 22.5–48.1 mm TL; TW 107–114% TL; height c. 14% TL; margins of ambulacra II, III, IV, and interambulacrum 5 indented; oral side flat to slightly concave, infundibulum depth 1–4% TL; food grooves bifurcating, closely following lunules and pressure drainage channels; well-defined fields of smaller tubercles in ambulacra compared to interambulacra on oral surface; plate sutures of aboral surface marked with light coloured margins, slightly depressed suture area gives impression of slight inflation of plate centres. Apical system —Monobasal, with four circular gonopores; subcentral, 45–51% TL from anterior margin. Ambulacra —Petals small, closed distally, except for petal III which is slightly open, especially so in small specimens; petaloid length approximately 42% TL; petal III slightly longer than others, length 22% TL, anterior paired petals approximately 20% TL, posterior paired petals shortest, 17% TL; relative width of all petals similar, c. 10% TL; pores distinctly conjugated, with sharply delimited furrows deepening towards outer pore; inner pores small and rounded, outer pores large and elongated; 3–7 primary tubercles on ridges between furrows; pairs of miliary tubercles between consecutive primary tubercles; elongated lunules in posterior ambulacra, mean LL 21% TL; lunules of variable width, 5–11% TL; SI of Stara & Fois (2014) 0.2–0.5; WI 3.7–5.7; LL greater than that of shortest petal; angle between lunules 75°–87°; distance between tips of posterior petals and lunules 7–11% TL; two pairs of plates between petal and lunule; four pairs of basicoronal plates in oral ambulacrum; food grooves bifurcating close to peristome, with more or less well-developed lateral branches encroaching upon interambulacra; oral ambulacra with much more densely packed and smaller tubercles than interambulacra; sharply delimited darker-coloured ambulacra relative to interambulacra; ambulacral width at ambitus 25–28% TL. Interambulacra —Oral interambulacra with less dense field of larger tubercles than ambulacra; interambulacrum 5 width at ambitus 37–40% TL; oral interambulacra all disjunct, widest gap observed in interambulacrum 5. Peristome —Round to subpentagonal, small, 4–7% TL in diameter; absolute peristome size not increasing at same rate as test with growth, relative peristome size is smallest in largest specimen; peristome anterior of centre, 44–48% TL from anterior margin. Periproct —Small, 2–4% TL; lies orally, 4–8% TL from posterior edge; bounded by plates 5.a.2/5.a.3/5.b.3. Geographic range. Indian Ocean, from Red Sea, East Coast of Africa (Mortensen 1948b) to Singapore (Stara & Sanciu 2014). Bathymetric range. Littoral to 12 m (Mortensen 1948b). Observed occurrence in Sri Lanka. Specimens were collected from muddy shorelines along the northwestern coast (Periyapaduwa) at 1–2 m, and near a fish landing site in Negombo 2 Sri Lanka (Fig. 58). Remarks. Following Stara and Sanciu (2014) the specimens in this study were identified as Echinodiscus because of the “almost vertical” (i.e. almost parallel to the anterior-posterior axis) suture between plates 5.a.2 and 5.b.2 and the low number of plates between the ends of the petals and the lunules (Fig. 51B). The shape and size of the lunules show marked variation among the examined specimens. The largest specimen has a low SI, but high WI, small specimens show the opposite trend. The second Sri Lankan Echinodiscus species, E. bisperforatus, has long, slit like lunules (length 34% TL) and a wide angle (112°) between them, whereas Echinodiscus cf. truncatus has shorter and wider lunules (length 21% TL), which form an acute angle of about 81°. The Sri Lankan material differs from E. truncatus described by Stara and Sanciu (2014) by its higher angle between lunules (81° vs. 67°), different periproct position (5.a.2/5.a.3/5.b.3 vs. 5.a.2/5.b.2/5.b.3), higher SI (0.37 vs. 0.22), lower WI (4.7 vs. 8.68), and much simpler food groves. Echinodiscus cf. truncatus differs from Echinodiscus sp. 1 by its large peristome (5.6% TL vs. E. andamanensis Stara & Sanciu, 2014 differs from the examined specimens by the position of the periproct, which opens between the plates 5.b.2/5.a.2 or between 5.b.2/5.a.2/5.a.3. Stara and Sanciu (2014: 305) report a fossil species called Echinodiscus sp. 2 from the Pleistocene of Hurgada (Red Sea) that shows many similarities to the present material, notably in periproct position, angle between lunules, and complexity of food groves. According to Clark (1914) Echinodiscus bisperforatus var. truncatus has “lunules shorter than shortest petal” but in the examined specimens the lunules are usually longer than the shortest petal, sometimes matching petal III in length. According to Stara and Sanciu (2014), Clark’s (1914) E. truncatus may have been a misidentified specimen of Sculpsitechinus tenuissimus., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on pages 63-67, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Agassiz, L. (1841) Monographies d'Echinodermes vivans et fossiles. Echinites. Famille des Clypeasteroides. 2 (Seconde Monographie). Des Scutelles, Neuchatel, iv + 151 pp., 27 pls. https: // doi. org / 10.5962 / bhl. title. 126954","Stara, P. & Fois, M. (2014) Analysis on a sample of Echinodiscus cf. auritus Leske, 1778 (Echinoidea: Clypeasteroida), Biodiversity Journal, 5, 269 - 290.","Mortensen, T. (1948 b) A Monograph of the Echinoidea IV. 2. Clypeasteroida. Clypeasteridae, Arachnoidae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp.","Stara, P. & Sanciu, L. (2014) Analysis of some astriclypeids (Echinoidea: Clypeasteroida). Biodiversity Journal, 5, 291 - 358.","Clark, H. L. (1914) Hawaiian and other Pacific Echini-the Clypeasteridae, Arachnoididae, Laganidae, Fibulariidae, and Scutellidae. Memoirs of the Museum of Comparative Zoology, 46, 1 - 78."]}

Peronella lesueuri (L. Agassiz, 1841) Figures 42–44 1841 Laganum Lesueuri Val. L. Agassiz: p. 116; pl. 24: figs. 3–6. 1948 b Peronella Lesueuri (Valenciennes).—Mortensen: p. 263–269; pl. 47: figs. 1–4, 6, 7; pl. 48: figs. 1, 2; pl. 50: figs. 4–11; pl. 72: figs. 2, 3. Material studied. Three denuded tests: WUSL /EI/33, EI/34 and EI/35, from Chaddy Beach, Jaffna. Description. Shape and size —Test decagonal to elongate, medium-sized, 36.8–40.6 mm TL, longer than broad, width 90–96% TL, greatest width coinciding with tips of anterior paired petals; height 14–18% TL; oral side flat; edge slightly thickened, test raised toward centre. Apical system —Subcentral, c. 52% TL away from anterior margin; with four circular gonopores, hydropores scattered throughout madreporic plate. Ambulacra —Petaloid area broad, more than half TL, 59–64% TL, slightly elevated; petals broad, closed distally; petal III slightly longer than others, c. 30% TL (SD=2); anterior paired petals shortest, c. 26% TL (SD=1); posterior paired petals little shorter than petal III, c. 28% TL (SD=1); petal widths and width of interporiferous zones of all petals similar, c. 13% TL and 8% TL, respectively; width of interporiferous zone of petal III c. 9% TL; ratio between petal width and petal length 0.4–0.5; furrow connecting pores deep and sharply limited; food grooves simple and unbranched, short and inconspicuous. Interambulacra —Slightly inflated adapically between petals; densely packed with small primary tubercles; on oral side, interambulacra forming narrow, straight bands separating broad ambulacra. Tuberculation —Primary tubercles on oral side slightly larger than aboral ones; on oral side, glassy tubercles distributed equally even near peristome, at margin less densely; on aboral side glassy tubercles found only along margin, and are small relative to those on oral surface. Peristome —Small, 6–8% TL, rounded to pentagonal, width c. 7% TL; situated slightly anterior of centre, c. 47% TL away from anterior margin. Periproct —Small, 5% TL; rounded to irregular in outline on oral surface, c. 10% TL away from posterior margin; bounded by second and third postbasicoronal interambulacral plates. Internal buttressing —Well-developed and complicated, most abundant and branched close to margin, radiating inwards along ambulacrum (Fig. 43). Geographic range. Indo-West Pacific, from the West India & Pakistan (Koehler 1922), Maldives area (Koehler 1922), Sri Lanka (Koehler 1922), Bay of Bengal (Brown 1910b), East Indies (Clark 1925a; Mortensen 1948d), North Australia (Clark 1938) to Hervey Bay (Miskelly 2002), China & South Japan (Mortensen 1948b) and Philippine Islands (Clark 1914; Mortensen 1948e). Bathymetric range. Littoral to 70 m (Mortensen 1948b). Observed occurrence in Sri Lanka. Denuded tests only were found on the beach of Chaddy Beach, Jaffna northern coast of Sri Lanka (Fig. 44). Remarks. Sri Lankan P. lesueuri is similar to Peronella lesueuri gadiana Mortensen, 1948. According to Mortensen (1948b: 270), P. lesueuri gadiana differs from typical P. lesueuri “mainly in the tuberculation of the pore-zones, the series of miliary tubercles being interrupted by some primary tubercles”. However, this character could not be observed in the examined materials as the petaloid area was poorly preserved. Gonopores were absent in a specimen 39.8 mm TL (WUSL/EI/33)., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on pages 50-51, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Agassiz, L. (1841) Monographies d'Echinodermes vivans et fossiles. Echinites. Famille des Clypeasteroides. 2 (Seconde Monographie). Des Scutelles, Neuchatel, iv + 151 pp., 27 pls. https: // doi. org / 10.5962 / bhl. title. 126954","Koehler, R. (1922) Echinoderma of the Indian Museum, Part 9, An account of the Echinoidea. II. Clypeastrides et Cassidulides. Trustees of the Indian Museum, Calcutta, 161 pp.","Brown, R. N. R. (1910 b) Echinoidea and Asteroidea from, the Mergui. Archipelago and Moskos Islands, Lower Burma. Edinburgh Proceedings of the Royal Physical Society, 18, 21 - 35.","Clark, H. L. (1925 a) A Catalogue of the Recent Sea-Urchins (Echinoidea) in the Collection of the British Museum (Natural History). Oxford University Press, London, 250 pp.","Mortensen, T. (1948 d) New Echinoidea (Cassiduloida, Clypeasteroida): Preliminary notice. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i KObenhavn, 111, 67 - 72.","Clark, H. L. (1938) Echinoderms from Australia. Memoirs of the Museum of Comparative Zoology of the Harvard College, 55, 1 - 596.","Miskelly, A. (2002) Sea urchins of Australia and the Indo-Pacific. Capricornica Publications, Lindfield, 179 pp.","Mortensen, T. (1948 b) A Monograph of the Echinoidea IV. 2. Clypeasteroida. Clypeasteridae, Arachnoidae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp.","Clark, H. L. (1914) Hawaiian and other Pacific Echini-the Clypeasteridae, Arachnoididae, Laganidae, Fibulariidae, and Scutellidae. Memoirs of the Museum of Comparative Zoology, 46, 1 - 78.","Mortensen, T. (1948 e) Contributions to the Biology of the Philippine Archipelago and adjacent regions. Report on the Echinoidea collected by the United States Fisheries Steamer \" Albatross \" during the Philippine Expedition, 1907 - 1910. Part 3: The Echinoneidae, Echinolampidae, Clypeastridae, Arachnidae, Laganidae, Fibulariidae, Urechinidae, Echinocorythidae, Palaeostomatidae, Micrasteridae, Palaepneustidae, Hemiasteridae, and Spatangidae. Smithsonian Institution, United States National Museum Bulletin, 100, 93 - 140."]}

Suborder Scutellina Haeckel, 1896 Infraorder Laganiformes Desor, 1847, Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on page 32, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Agassiz, L. & Desor, P. J. E. (1847) Catalogue raisonne des especes, des genres, et des familles d'echinides. Annales des Sciences Naturelles, Troisieme Serie, Zoologie 7, 129 - 168."]}

Clypeaster reticulatus (Linnaeus, 1758) Figures 22–24 1758 Echinus reticulatus Linnaeus: p. 666. 1948b Clypeaster (Rhaphidoclypus) reticulatus (Linnaeus).—Mortensen: p. 71–76; pl. 18: figs. 1–21; pl. 26: fig. 3; pl. 65: figs. 2, 13, 16. Material studied. Two denuded specimens: WUSL /EI/19 from Hiriketiya and WUSL /EI/20 from Nilwella, Sri Lanka. Description. Shape and size —Small, 35 and 41 mm TL; test elongate ovoid to pentagonal, distinctly longer than broad, TW 75% and 76% TL; test low, 26% and 31% TL in height; test margin greatly thickened, forming a distinct ring-shaped depression between margin and petalodium; aboral side slightly raised toward apex, concave on oral side, infundibulum deep, bowl-shaped, c. 14% TL and 17% TL. Apical system —Monobasal; situated slightly anterior of centre on aboral surface, c. 45% and 48% TL from anterior margin; circular gonopore in each interambulacrum near madreporic plate; ocular pores small and indistinct. Ambulacra —Petaloid area 63% and 65% TL; petals inflated, closed distally; petal III longest, c. 32% TL; anterior paired petals c. 23% TL, considerably shorter than other petals; posterior paired petals c. 30% TL; paired petals broader distally than other petals; interporiferous zone conspicuously elevated, consisting of 5–6 primary tubercles along widest part of petals; furrow connecting pores in each pore pair deep and sharply delimited; 2–5 primary tubercles between furrows; oral ambulacra simple, inconspicuous food groves present along axis of each ambulacrum. Interambulacra —All interambulacra on oral surface disjunct; on aboral surface, small, shallow pits present in corners of sutures (Fig. 23, A); primary tubercles less dense on oral surface than aborally. Tuberculation —Primary tubercles perforate, areoles sunken, dense miliary tuberculation in between primary tubercles; oral side with adoral region of much smaller tubercles; largest tubercles present toward test edge, intermediate sized tubercles present very close to peristome, smallest tubercles present in between zone of largest tubercles and zone of intermediate-sized tubercles (Fig. 23, B). Peristome —Round to subpentagonal, small, width c. 7% TL, located close to centre, c. 46% and 51% TL from anterior margin. Periproct —Round to transverse oval, length 4% and 5% TL, width 4% and 6% TL; significantly smaller than peristome, c. 64% of peristome length; situated close to posterior edge of test, c. 4% TL away from posterior margin. Internal buttressing —Pillars inside test more abundant along midlines of interambulacra; marginal buttressing largely absent (Fig. 23, C). Geographic range. Indo-West Pacific, from Islands of West Indian Ocean (Clark 1925a), Mauritius (de Loriol 1883), East Africa & Madagascar (Brown 1910a), Red Sea (Agassiz 1872), South East Arabia (Mortensen 1948c), Persian Gulf (Mortensen 1940), West Indian & Pakistan (Koehler 1922), Maldives area (Koehler 1922), Sri Lanka (Herdman et al. 1904; Koehler 1922), Bay of Bengal (Koehler 1922), North Australia (Clark 1925a) and East Indies (de Meijere 1904; Mortensen 1948a) to Philippine Islands (Mortensen 1948e; Mooi & Munguia 2014), China & South Japan (Clark 1925a), South Pacific Islands (Clark 1954) and Hawaiian Islands (Clark 1925b). Bathymetric range. Littoral zone to 125 m (Mooi & Munguia 2014). Observed occurrence in Sri Lanka. On the beach of Hiriketiya Bay and Nilwella along the southern coast of Sri Lanka (Fig. 24), in biogenic sand; first recorded in Sri Lanka by Herdman et al. (1904). Bathymetric range in Sri Lanka 0–100 m. Remarks. C. reticulatus can be distinguished easily from C. humilis by having a greatly thickened test margin that forms a distinct ring-shaped depression between the margin and petaloid area. The oral surface of C. reticulatus is strongly concave, making a bowl-shaped infundibulum. Primary spine tubercles decrease in size towards the peristome, then increase again adjacent to the peristome., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on pages 30-31, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Clark, H. L. (1925 a) A Catalogue of the Recent Sea-Urchins (Echinoidea) in the Collection of the British Museum (Natural History). Oxford University Press, London, 250 pp.","De Loriol, P. (1883) Catalogue raisonne des Echinodermes recueillis par MV de Robillard a l'ile Maurice. Memoires de la Societe de physique et d'histoire naturelle de Geneve, 28, 1 - 64.","Brown, R. N. R. (1910 a) Echinoidea from the Kerimba Archipelago, Portuguese East Africa (Mozambique). Edinburgh Proceedings of the Royal Physical Society, 18, 36 - 44.","Agassiz, A. (1872 - 1874) Revision of the Echini. Illustrated Catalogue of the Museum of Comparative Zoology at Harvard College, 7 (Pt. 1 - 2, 3 & 4) i-xii + 1 - 378, pls. 1 - 49 (1872), 379 - 628 + 1, pls. 50 - 77 (1873) & 629 - 762, pls. 78 - 94 (1874).","Mortensen, T. (1948 c) Report on the Echinoidea of the Murray Expedition. II [Irregular Echinoidea]. Scientific Reports on the John Murray Expedition, 9, 1 - 15.","Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Koehler, R. (1922) Echinoderma of the Indian Museum, Part 9, An account of the Echinoidea. II. Clypeastrides et Cassidulides. Trustees of the Indian Museum, Calcutta, 161 pp.","Herdman, W. A., Herdman, J. B. & Bell, F. J. (1904) Report on the Echinoderma collected by Professor Herdman, at Ceylon, in 1902. In: Herdman, W. A. (Ed.), Report to the government of Ceylon on the pearl oyster fisheries of the Gulf of Mannar. The Royal Society, London, pp. 137 - 150.","de Meijere, J. C. H. (1904) Die Echinoidea der Siboga-Expedition. Siboga Expeditie, 43, 1 - 252.","Mortensen, T. (1948 a) A Monograph of the Echinoidea IV. 1. Holectypoida, Cassiduloida. C. A. Reitzel, Copenhagen, 371 pp.","Mortensen, T. (1948 e) Contributions to the Biology of the Philippine Archipelago and adjacent regions. Report on the Echinoidea collected by the United States Fisheries Steamer \" Albatross \" during the Philippine Expedition, 1907 - 1910. Part 3: The Echinoneidae, Echinolampidae, Clypeastridae, Arachnidae, Laganidae, Fibulariidae, Urechinidae, Echinocorythidae, Palaeostomatidae, Micrasteridae, Palaepneustidae, Hemiasteridae, and Spatangidae. Smithsonian Institution, United States National Museum Bulletin, 100, 93 - 140.","Mooi, R. & Munguia, A. (2014) Sea Urchins of the Philippines. In: Williams, G. C. & Terrence, M. G. (Eds.), The Coral Triangle: The 2011 Hearst Philippine Biodiversity Expedition. California Academy of Sciences, San Francisco, California, pp. 213 - 235.","Clark, A. H. (1954) Records of Indo-Pacific echinoderms. Pacific Science, 8, 243 - 263.","Clark, H. L. (1925 b) Marine zoology of tropical central Pacific: Echinoderms other than sea-stars. Bulletin of the Bernice Pauahi Bishop Museum, 27, 89 - 111."]}

Echinocyamus megapetalus H.L. Clark, 1914 Figures 25���27 1914 Echinocyamus megapetalus H.L. Clark: p. 60; pl. 126. 1948b Echinocyamus megapetalus H. L. Clark. ���Mortensen: p. 190���191; pl. 46: figs. 48���50. Material studied. Nine denuded tests: WUSL /EI/152, EI/153, EI/154, EI/155, EI/156, EI/157, EI/195, EI/196, and EI/197 from Hiriketiya Bay, Sri Lanka. Description. Shape and size ���Test elongate, 3.72���7.35 mm TL; longer than broad, width, 73���80% TL; height, 41���45% TL; oral side deeply sunken, aboral side flattened. Apical system ���Situated close to centre, 50���56% TL from anterior margin; with four circular gonopores, single hydropore, sometimes distorted to resemble groove with two or three hydropores (Fig. 26). Ambulacra ���Petaloid region length more than half TL (75���90% TL); petal III with 7���14 pore pairs, anterior paired petals with 5���12 pore pairs, posterior paired petals with 7���13 pore pairs; petals broad, open distally; pore series diverging; interporiferous zones wider than poriferous zones. Peristome ���Elongate pentagonal; moderate size, length 20���28% TL, width 16���22% TL, slightly anterior of centre, 38���42% TL from anterior margin, infundibulum deep. Periproct ���Small, 9���12% TL; transversely oval; wider than long, width 11���14% TL; inframarginal, 14���23% TL from posterior margin. Geographic range. Indo-West Pacific, from Mauritius (Clark 1914) to Coral Sea (Miskelly 2002), South Pacific Islands (Clark 1954; Mortensen 1948b) and Hawaiian Islands (Mortensen 1948b). Bathymetric range. 20���75 m (Markello 2015). Observed occurrence in Sri Lanka. Only denuded tests were found on the beach of Hiriketiya Bay, southern coast of Sri Lanka (Fig. 27). Remarks. E. megapetalus is recorded for the first time in Sri Lanka. E. megapetalus is similar to E. crispus in having a deep infundibulum but E. megapetalus is easily distinguished from E. crispus and other Indo-Pacific species by having broad, diverging petals that cover the majority of the aboral surface and by its deep infundibulum. However, studied specimens show great variation in test size and petaloid area length., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on pages 32-35, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Clark, H. L. (1914) Hawaiian and other Pacific Echini-the Clypeasteridae, Arachnoididae, Laganidae, Fibulariidae, and Scutellidae. Memoirs of the Museum of Comparative Zoology, 46, 1 - 78.","Miskelly, A. (2002) Sea urchins of Australia and the Indo-Pacific. Capricornica Publications, Lindfield, 179 pp.","Clark, A. H. (1954) Records of Indo-Pacific echinoderms. Pacific Science, 8, 243 - 263.","Mortensen, T. (1948 b) A Monograph of the Echinoidea IV. 2. Clypeasteroida. Clypeasteridae, Arachnoidae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp.","Markello, K. M. (2015) Phylogenetic revision of the micro-echinoid genus, Echinocyamus. San Francisco State University, San Francisco, California, 308 pp."]}

Clypeaster humilis (Leske, 1778) Figures 19–21 1778 Echinanthus humilis Leske: p. 185–189; pl. 17: fig. A; pl. 18: fig. B; pl. 19: figs. A–D. 1948b Clypeaster (Stolonoclypus) humilis (Leske).—Mortensen: p. 88–94; pl. 17: fig. 1; pl. 28: figs. 1–4; pl. 29: figs. 1–3, 5–8; pl. 30: fig. 1; pl. 40: figs. 2, 3; pl. 47: figs. 3, 8, 11, 15–18. Material studied. Six denuded specimens: WUSL /EI/10, from Marawila, WUSL /EI/13, from Mount Lavinia, and WUSL /EI/11, EI/12, EI/14 and EI/15 from Negombo, Sri Lanka. Description. Shape and size —Test pentagonal, large, 70.5–96.9 mm TL, distinctly longer than broad, width 87–93% TL, greatest width at anterior paired petals, more prominently demonstrated in larger specimens; TH 16–21% TL; in larger specimens, all interambulacra (except interambulacrum 5) usually conspicuously indented at test margin; in smaller specimens, posterior end more rounded and all other interambulacra only slightly indented or with regular ovoid outline; oral side more or less flat to slightly concave, particularly near peristome; infundibulum shallow, 6–8% TL in depth. Apical system —Monobasal, situated closer to centre of aboral surface, 50–53% TL (mean=52% TL, SD=1.2) away from anterior margin; a circular gonopore in each interambulacrum adjacent to madreporic plate; ocular pores small and indistinct. Ambulacra —Petaloid small, 53–56% TL (mean=55% TL, SD=1.0), considerably elevated; petals more or less broad, elongate ovoid in shape, closed distally; except in petal III, which is slightly open distally in some specimens; opening 1–4% TL; petal III length 27–30% TL (mean 29% TL) and is distinctly longer than other petals; anterior paired petals length range from 23 to 26% TL (mean 25% TL) and are shorter than other petals; posterior paired petals length 27–28% TL (mean 27% TL). Shape of inner pore of each pore pair circular, outer pore elongate, length about twice diameter of inner pore; interporiferous zones conspicuously elevated, bearing 9–16 primary tubercles across widest part of petals; furrow connecting pores of each pore pair deep and sharply delimited; 3–7 primary tubercles on ridges between furrows; ambulacral furrows of oral side simple unbranched and prominent. Interambulacra —Slightly inflated adapically between petals; densely packed with primary tubercles; all interambulacra on oral surface disjunct and more or less flat. Tuberculation —Primary tubercles perforate, areoles sunken; miliary tubercles more or less densely and evenly distributed among primary tubercles; density and size (mean size=0.23% TL, N=5) of primary tubercles on aboral surface almost homogenous, primary tubercles on oral surface slightly larger (mean=0.30% TL, N=5), density increasing towards edge of test. Peristome —Small, 5–8% TL (mean length=6% TL, mean width=7% TL); slightly wider than long, located slightly anterior of centre, c. 48% TL away from anterior margin. Periproct —Small, 3–6% TL; slightly oval, transversely elongated (mean width=5% TL), situated close to posterior edge, c. 3% TL from posterior margin. Internal buttressing —Marginal buttressing well-developed and reinforced by dense internal pillar system, best developed close to margin. Geographic range. Indo-West Pacific, from East Africa & Madagascar (Brown 1910a; Clark 1923), Red Sea (Koehler 1922), South East Arabia (Clark 1925a; Mortensen 1948c), Persian Gulf (Mortensen 1940), Sri Lanka (Clark 1915; Herdman et al. 1904), North Australia (Clark 1946) and East Indies (de Meijere 1904) to Philippine Islands (Mortensen 1948e; Mooi & Munguia 2014) and South Pacific Islands (Agassiz 1872). Bathymetric range. 0–216 m (Mortensen 1948b). Observed occurrence in Sri Lanka. Sandy bottoms on the western, and north-western coasts of Sri Lanka at depths of 25–30 m (Fig. 21); first recorded in Sri Lanka by Agassiz (1872). Bathymetric range in Sri Lanka is 15–30 m. Remarks. An open petal III was observed in each examined specimen with the degree of opening being extremely variable. In addition, specimens with a highly elevated petaloid area have more open anterior petals. This is consistent with similar observations by Mortensen (1948a: 91)., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on pages 26-30, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Brown, R. N. R. (1910 a) Echinoidea from the Kerimba Archipelago, Portuguese East Africa (Mozambique). Edinburgh Proceedings of the Royal Physical Society, 18, 36 - 44.","Clark, H. L. (1923) The Echinoderm fauna of South Africa. Annals of the South African Museum, 13, 221 - 435.","Koehler, R. (1922) Echinoderma of the Indian Museum, Part 9, An account of the Echinoidea. II. Clypeastrides et Cassidulides. Trustees of the Indian Museum, Calcutta, 161 pp.","Clark, H. L. (1925 a) A Catalogue of the Recent Sea-Urchins (Echinoidea) in the Collection of the British Museum (Natural History). Oxford University Press, London, 250 pp.","Mortensen, T. (1948 c) Report on the Echinoidea of the Murray Expedition. II [Irregular Echinoidea]. Scientific Reports on the John Murray Expedition, 9, 1 - 15.","Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","Clark, H. L. (1915) The echinoderms of Ceylon other than holothurians. Spolia Zeylanica, 10, 83 - 102.","Herdman, W. A., Herdman, J. B. & Bell, F. J. (1904) Report on the Echinoderma collected by Professor Herdman, at Ceylon, in 1902. In: Herdman, W. A. (Ed.), Report to the government of Ceylon on the pearl oyster fisheries of the Gulf of Mannar. The Royal Society, London, pp. 137 - 150.","Clark, H. L. (1946) The echinoderm fauna of Australia. Its composition and its origin. Carnegie Institution of Washington Publication, 566, 1 - 567.","de Meijere, J. C. H. (1904) Die Echinoidea der Siboga-Expedition. Siboga Expeditie, 43, 1 - 252.","Mortensen, T. (1948 e) Contributions to the Biology of the Philippine Archipelago and adjacent regions. Report on the Echinoidea collected by the United States Fisheries Steamer \" Albatross \" during the Philippine Expedition, 1907 - 1910. Part 3: The Echinoneidae, Echinolampidae, Clypeastridae, Arachnidae, Laganidae, Fibulariidae, Urechinidae, Echinocorythidae, Palaeostomatidae, Micrasteridae, Palaepneustidae, Hemiasteridae, and Spatangidae. Smithsonian Institution, United States National Museum Bulletin, 100, 93 - 140.","Mooi, R. & Munguia, A. (2014) Sea Urchins of the Philippines. In: Williams, G. C. & Terrence, M. G. (Eds.), The Coral Triangle: The 2011 Hearst Philippine Biodiversity Expedition. California Academy of Sciences, San Francisco, California, pp. 213 - 235.","Agassiz, A. (1872 - 1874) Revision of the Echini. Illustrated Catalogue of the Museum of Comparative Zoology at Harvard College, 7 (Pt. 1 - 2, 3 & 4) i-xii + 1 - 378, pls. 1 - 49 (1872), 379 - 628 + 1, pls. 50 - 77 (1873) & 629 - 762, pls. 78 - 94 (1874).","Mortensen, T. (1948 b) A Monograph of the Echinoidea IV. 2. Clypeasteroida. Clypeasteridae, Arachnoidae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp.","Mortensen, T. (1948 a) A Monograph of the Echinoidea IV. 1. Holectypoida, Cassiduloida. C. A. Reitzel, Copenhagen, 371 pp."]}

Fibulariella angulipora Mortensen, 1948 Figures 34, 35 1948d Fibularia (Fibulariella) angulipora Mortensen: p. 6. 1948b Fibularia (Fibulariella) angulipora Mortensen. ���Mortensen: p. 224���225; pl. 46: figs. 1���4, 12���14. Material studied. Ten denuded tests: WUSL/EI/132, EI/133, EI/144, EI/145, EI/146, EI/147, EI/148, EI/149, EI/ 150, and EI/151, from Etalai, Kalpitiya, Sri Lanka. Description. Shape and size ���Test oval, rounded or pointed at both ends, small size, 3.39���7.14 mm TL, longer than broad, width 65���71% TL, height 43���53% TL; greatest height usually anterior to apical system; oral side convex; raised, rounded margin; aboral surface sloping slightly downward towards posterior end. Apical system ���Subcentral, 41���51% TL from anterior margin; four small, circular gonopores, multiple small hydropores, not in groove, scattered over anterior portion of madreporic plate. Ambulacra ���Petaloid area longer than half of TL, 66���71% TL; petaloid area width 40���46% TL; petals short, with 3���5 pore pairs in posterior paired petals, 3���4 pore pairs in anterior paired petals, 4���5 pore pairs in petal III; pores in posterior petals markedly triangular. Peristome ���Medium-sized, 15���19% TL; usually rounded, width 13���17% TL; lies slightly anterior of centre, 40���47% TL from anterior margin. Periproct ���Small, 8���9% TL; oval in outline, slightly elongate; situated closer to peristome than posterior edge, 17���23% TL from posterior margin. Geographic range. Indo-West Pacific, from Andaman Sea (Putchakarn & Sonchaeng 2004) to Gulf of Siam and the Molucca Sea (Mortensen 1948b). Bathymetric range. 15 m (Mortensen 1948b). Observed occurrence in Sri Lanka. The current study found only denuded tests of F. angulipora on the shore of Etalai, Kalpitiya, in north-western coasts of Sri Lanka (Fig. 35). Remarks. According to (Mortensen 1948b: 225), this species differs from the two other Fibulariella species, F. acuta and F. oblonga, by the large, triangular pores in the petals. F. angulipora is recorded here for the first time from Sri Lanka. This supports Mortensen's (1948b: 225) suggestion that the specimen mentioned as Fibularia volva by H. L Clark (1915) was in reality Fibulariella angulipora, not Fibularia volva., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on pages 41-44, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Putchakarn, S. & Sonchaeng, P. (2004) Echinoderm fauna of Thailand: History and inventory reviews. Science Asia, 30, 417 - 428. https: // doi. org / 10.2306 / scienceasia 1513 - 1874.2004.30.417","Mortensen, T. (1948 b) A Monograph of the Echinoidea IV. 2. Clypeasteroida. Clypeasteridae, Arachnoidae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp.","Clark, H. L. (1915) The echinoderms of Ceylon other than holothurians. Spolia Zeylanica, 10, 83 - 102."]}

Fibularia sp. Figures 27, 32, 33 Material studied. Eleven denuded tests: WUSL /EI/158, EI/159, EI/161, EI/164, EI/165, EI/179, EI/180, EI/181, EI/185, EI/186, and EI/187, from Hiriketiya Bay, Sri Lanka. Description. Shape and size ���Test oval, small, 5.3���8.72 mm TL, longer than broad, width 75���84% TL, height 44���50% TL, posterior part of test wider than anterior, test slightly pointed anteriorly and truncated posteriorly; oral side slightly concave. Apical system ���Situated posterior of centre, 53���59% TL from anterior margin; with four circular small gonopores and small hydropores placed in short groove (Fig. 33, A). Ambulacra ���Petaloid area 69���76% TL, petaloid width 47���49% TL; petals short, with 5���8 pore pairs in each series of petal III and posterior petals, 3���5 pore pairs in each series of anterior-paired petals; pores small, rounded; petals broad, open distally; pore series slightly diverging; interporiferous zones similar in width to poriferous zones. Peristome ���Moderate in size, length 16���22% TL; rounded pentagonal, width 12���17% TL, situated slightly anterior of centre, 39���43% TL from anterior margin. Periproct ���Small, length 11���12% TL; squarish in outline; slightly elongate, width 8���11% TL; situated closer to peristome than posterior edge, 16���22% TL from posterior margin, four plates on periproctal membrane, no spines on plates. Observed occurrence in Sri Lanka. Denuded tests only were found on the beach of Hiriketiya Bay, southern coast of Sri Lanka (Fig. 27). Remarks. This taxon likely represents a new species currently under study by one of us (Rich Mooi). Fibularia sp. can be distinguished from F. cribellum, F. japonica, F. plateia, and F. nutriens in having higher numbers of pore pairs relative to TL. This species is very similar to F. ovulum as both have a relatively high number of pore pairs. However, Fibularia sp. can be consistently distinguished from F. ovulum by its lower test., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on page 39, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460

Jacksonaster sp. 1 Figures 36–38, 40A Material studied. Seven denuded tests: WUSL /EI/23, EI/26, EI/27, EI/140, EI/141, and EI/142 from Thennadi Bay, and WUSL /EI/27 from Chaddy Beach, Jaffna, Sri Lanka. Description. Shape and size —Test pentagonal, posterior end gently indented; medium-sized, 30.0– 47.2 mm TL, slightly longer than broad, width 88–93% TL, larger specimens more elongate than smaller specimens; greatest width at end of anterior paired petals; height 15–17% of TL; oral side flat; edge thin, 7–10% TL, posterior edge slightly thinner than anterior edge. Apical system —Subcentral, approximately 52% TL from anterior margin of test; with five circular gonopores; hydropores in sinuous groove. Ambulacra —Petaloid area large, 67–71% TL, slightly elevated; petals narrow, slightly open distally; pore series straight; pores elongate anisopores; outer pores of pore pairs greatly elongated transversely relative to inner pores; pores conjugate, but connecting furrows shallow; petal III slightly longer than others, approximately 35% TL (SD=2); anterior paired petals shortest, approximately 31% TL (SD=1); posterior paired petals slightly shorter than petal III, approximately 33% TL (SD=2); all petal widths similar, approximately 12% TL; width of poriferous zone IIIa 75–89% of interporiferous zone width at 2/3 of the petal length; food grooves simple, unbranched, lying along each oral perradius, up to 62–70% of corresponding test radius in ambulacrum III. Interambulacra —Oral interambulacra narrow, forming straight bands separating broad ambulacra. Tuberculation —Primary tubercles small and densely distributed among numerous miliary tubercles on aboral side; on oral surface, primary tubercles larger and sparse relative to those on aboral surface; primary tubercles of oral interambulacra close to peristome slightly larger than those of oral ambulacra; smallest primary tubercles near food grooves. Peristome —Small, length 6–8% TL, rounded to pentagonal, width 5–8% TL; situated slightly anterior of centre, 45–48% TL from anterior margin of test. Periproct —Small, length 5% TL, transversely elongate to rounded, width 6–7% TL; lying 11–12% TL from posterior margin of test. Observed occurrence in Sri Lanka. Specimens were collected on the beach at Chaddy Beach, Jaffna, northern coast and Thennadi Bay, eastern coast of Sri Lanka. This species was also observed in Mount Lavinia (at 30 m) and Negombo 1 (at 27 m), western coast and at Mulathiv (at 10 m), northern coast of Sri Lanka (Fig. 38). Remarks. Considering the characters pointed out by Mortensen (1948b), these specimens best match Jacksonaster depressum tenue (Mortensen, 1948). The latter subspecies has a thinner test and margin, and narrower petals with straighter pore series than usually found in J. depressum from other parts of its range. Jacksonaster sp. 1 can be distinguished from Jacksonaster sp. 2 by having a much thinner test, thinner test margin, and slightly open petals with straight pore series. In addition, Jacksonaster sp. 1 has narrower poriferous zones than Jacksonaster sp. 2, being approximately 81% of the width of the interporiferous zone in the former and 116% in the latter (Fig. 40A)., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on pages 44-45, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Mortensen, T. (1948 b) A Monograph of the Echinoidea IV. 2. Clypeasteroida. Clypeasteridae, Arachnoidae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp."]}

Peronella oblonga Mortensen, 1948 Figures 45���49 1948a Peronella oblonga Mortensen: p. 71. 1948b Peronella oblonga Mortensen. ���Mortensen: p. 299���300; pl. 51: figs. 33, 34. Material studied. Seven denuded tests: WUSL /EI/38, EI/39, EI/41, EI/48, EI/49, EI/52, and EI/143 from Casuarina Beach, Jaffna. Description. Shape and size ���Test regularly oval in outline, with a pointed to truncated posterior end; small to medium-sized, 18.03���34.57 mm; longer than broad, width 87���90% TL; height 15���20% TL; oral side slightly concave; margin distinctly thickened, with a distinct depression between margin and slightly elevated central part; anterior edge thicker (approximately 14% TL; SD=1), than posterior edge of test (approximately 12% TL; SD=1). Apical system ���Subcentral, 49���50% TL away from anterior margin; with four circular gonopores. Ambulacra ���Petaloid area large, length 55���59% TL, slightly elevated; petals fairly broad (50���55%TL in width), closed distally; petal III slightly longer than others, c. 26% TL (SD=1); anterior paired petals shorter than others, c. 23% TL (SD=1); posterior paired c. 25% TL (SD=1); petal widths and width of interporiferous zones of anterior paired and posterior paired petals similar, c. 13% TL and 8% TL, respectively; outer pore larger than inner pore; pore pairs distinctly conjugate; ridges between consecutive pore pairs carry single, regular series of 6���8 small tubercles; food grooves simple, unbranched, short and inconspicuous, extending 29���39% of corresponding test radius in ambulacrum III. Interambulacra ���Slightly inflated adapically between petals; on oral side, interambulacra forming narrow, straight bands separating broad ambulacra. Tuberculation ���Primary tubercles relatively large, with well-marked, sunken areole; on aboral surface, primary tubercles small and densely distributed among numerous miliary tubercles; on oral surface, primary tubercles larger and sparse relative to those on aboral surface; glassy tubercles larger than primary tubercles; densely distributed near edge of test on both oral and aboral sides; a few small glassy tubercles scattered over oral surface (Fig. 46, 47). Peristome ���Small, length 8���10% TL; rounded to pentagonal; situated slightly anterior of centre, 43���45% TL away from anterior margin. Periproct ���Small, length 5���6% TL; transverse oval to nearly round in outline; width from 6 to 8% TL; inframarginal, 10���13% TL away from posterior margin; bounded by second and third postbasicoronal interambulacral plates. Geographic range. Indian Ocean, from Tuticorin Beach, India (Mortensen 1948b). Bathymetric range. To date, found only on beach (Mortensen 1948b). Observed occurrence in Sri Lanka. No live specimens were found, dead tests only were collected on Casuarina Beach, Jaffna, northern coasts of Sri Lanka (Fig. 49). Remarks. Based on test characters, the studied material appears closest to P. oblonga as described by Mortensen (1948b). The materials examined show marked variation in glassy tubercle size and distribution on the oral and aboral surfaces. In the largest specimen examined (WUSL/EI/41, TL= 34.57 mm), glassy tubercles are smaller than those in smaller specimens relative to test size (Fig. 46). On the aboral surface close to the edge of the test, glassy tubercles are rare and much less conspicuous compared to the oral side. On the oral surface, small glassy tubercles are dispersed more or less equally even near the peristome but are especially conspicuous near the edge of the test. Glassy tubercle size and distribution are similar to Peronella lesueuri gadiana but P. oblonga is quite distinct from P. l. gadiana by the thicker test margin, and a distinct depression between the margin and slightly elevated central region, small petaloid area, and short petals. In small specimens, glassy tubercles were much more conspicuous than the primary tubercles near the edge of the test, and raised above the primary tubercles, giving the latter a sunken appearance. There is variation in petal length among the studied specimens. Some specimens (WUSL/EI/38 and EI/48) have equally sized anterior and posterior petals while WUSL/EI/143 has longer posterior petals compared to petal III. WUSL/EI/41 and EI/52 have greater test edge thickness relative to test height. A few specimens (WUSL/EI/50, EI/51, and EI/53) collected from the same locality have an unusually pointed anterior end. In addition, these specimens have a higher test edge thickness and apical system situated 51���52% TL from anterior margin (Fig. 48)., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on pages 53-57, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Mortensen, T. (1948 b) A Monograph of the Echinoidea IV. 2. Clypeasteroida. Clypeasteridae, Arachnoidae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp."]}

Fibularia ovulum Lamarck, 1816 Figures 27, 28���31 1816 Fibularia ovulum Lamarck: p. 17. 1948b Fibularia ovulum Lamarck. ���Mortensen: p. 208���212; pl. 46: figs. 48���50, 21���24. Ambulacra ���Petaloid area 61���74% TL, petaloid width 42���51% TL; petals short, with 4���7 pore pairs in each series of petal III and posterior paired petals, 3���5 pore pairs in each series of anterior paired petals; pores small rounded, petals broad, open distally; pore series slightly diverging; interporiferous zones smaller than poriferous zones. Peristome ���Moderate in size, diameter 17���21% TL; rounded pentagonal, width 12���17% TL, situated slightly anterior of centre, 37���42% TL from anterior margin of test. Periproct ���Small, 9���13% TL; squarish in outline; slightly elongate, width, 8���11% TL; situated closer to peristome than posterior edge, 17���26% TL from posterior margin of test. Geographic range. Indo-West Pacific, from the Red Sea (Clark 1925a; Mortensen 1948c) to Coral Sea (Miskelly 2002) and the Philippines (Mortensen 1948b). Bathymetric range. Intertidal zone to 385 m (Mortensen 1948b). Observed occurrence in Sri Lanka. Denuded tests only were found on the beach of Hiriketiya Bay, southern coast of Sri Lanka (Fig. 27). Remarks. Fibularia ovulum differs from Fibularia sp. in having a higher test (Fig. 30). Test height of Sri Lankan Fibularia ovulum is relatively low compared to specimens from the Maldives. However, the total petaloid pore pair number overlaps with that of with Fibularia sp. (Fig. 31)., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on pages 37-38, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Clark, H. L. (1925 a) A Catalogue of the Recent Sea-Urchins (Echinoidea) in the Collection of the British Museum (Natural History). Oxford University Press, London, 250 pp.","Mortensen, T. (1948 c) Report on the Echinoidea of the Murray Expedition. II [Irregular Echinoidea]. Scientific Reports on the John Murray Expedition, 9, 1 - 15.","Miskelly, A. (2002) Sea urchins of Australia and the Indo-Pacific. Capricornica Publications, Lindfield, 179 pp.","Mortensen, T. (1948 b) A Monograph of the Echinoidea IV. 2. Clypeasteroida. Clypeasteridae, Arachnoidae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp."]}

Echinodiscus bisperforatus Leske, 1778 Figures 50, 51A, 52���54 1778 Echinodiscus bisperforatus Leske: p.132���133; pl. 21: figs. A���B. 1948b Echinodiscus bisperforatus Leske. ���Mortensen: p. 406���410; pl. 68: figs. 2, 6���8; pl. 71: figs. 6���9, 18. Apical system ���Subcentral, 48���52% TL away from anterior margin, monobasal with four small circular gonopores. Ambulacra ���Well-developed short petals, more or less closed distally; petaloid area broad, mean petaloid length approximately 44% TL (SD=2); mean width approximately 46% TL (SD=2); petal III slightly longer than other petals, c. 24% TL (SD=1); anterior paired petals c. 21% TL (SD=1); posterior paired petals distinctly shorter than others, c. 17% TL (SD=1); petal III and anterior paired petal slightly wider (mean 13% TL) than posterior paired petal (mean 12% TL); pores in pore pairs distinctly conjugated, with sharply delimited furrows that deepen towards outer pore; 12 to 16 primary tubercles on ridges between furrows. Two slit-like lunules in posterior ambulacra; lunule length c. 34% TL and lunule width c. 3% TL; angles between two lunules range from 108�� to 117��; adapical end of lunule pointed anteriorly and distal end pointed posteriorly; single pair of plates between lunules and tips of posterior petals (Fig. 53A); food grooves bifurcating close to peristome, with more or less welldeveloped lateral branches encroaching upon interambulacra. Interambulacra ���Anal groove distinct; two plates per column in oral interambulacrum 5; oral interambulacra usually disjunct, but in some cases interambulacrum 2 or 3 (rarely both) are contiguous (Fig. 52). Tuberculation ���Oral ambulacra with much more densely packed, small tubercles than in interambulacra. Peristome ���Round to subpentagonal, small, 3���4% TL in diameter; slightly anterior of test centre, 31���36% TL from anterior margin. Periproct ���Small, 1���3% TL in diameter; close to posterior margin of test, 6���8% TL from posterior edge; opens along suture between 5.a.2/5.b.2 or at junction of 5.a.2/5.b.2/5.a.3 or 5.a.2/5.b.2/5.b.3 of post-basicoronal plates (Fig. 53). Geographic range. Indian Ocean, from Mauritius (Clark 1923), East Africa & Madagascar (Agassiz 1872), Red Sea (Clark 1925a), Sri Lanka (Bell 1887a), Bay of Bengal (Koehler 1922) to East Indies (Mortensen 1948b). Bathymetric range. Intertidal zone to 13 m (Koehler 1922). Observed occurrence in Sri Lanka. Specimens were collected from a muddy bottom at 1���2 m along the north-western coast (Periyapaduwa) and at 1 m along the eastern coast (Kaththankudi) of Sri Lanka (Fig. 54). First recorded in Sri Lanka by Bell (1882). Remarks. There is high variability in the position of the periproct (Fig. 53). Therefore, this is not a reliable character to distinguish E. bisperforatus from other species in the genus. The suture between plate 5.a.2 and 5.b.2 is either nearly parallel to the anterior-posterior axis, or distinctly oblique to this axis (close to 45��, see Fig. 55). In the former case, it joins the adradial suture of plate V.b.2 and in the latter case, V.b.3. Otherwise, the studied E. bisperforatus specimens are very similar. This undermines the validity of the genus Paraamphiope recently established solely on this feature by Stara and Sanciu (2014)., Published as part of Arachchige, Gayashan M., Jayakody, Sevvandi, Mooi, Rich & Kroh, Andreas, 2019, Taxonomy and distribution of irregular echinoids (Echinoidea: Irregularia) from Sri Lanka, pp. 1-100 in Zootaxa 4541 (1) on pages 57-60, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Clark, H. L. (1923) The Echinoderm fauna of South Africa. Annals of the South African Museum, 13, 221 - 435.","Agassiz, A. (1872 - 1874) Revision of the Echini. Illustrated Catalogue of the Museum of Comparative Zoology at Harvard College, 7 (Pt. 1 - 2, 3 & 4) i-xii + 1 - 378, pls. 1 - 49 (1872), 379 - 628 + 1, pls. 50 - 77 (1873) & 629 - 762, pls. 78 - 94 (1874).","Clark, H. L. (1925 a) A Catalogue of the Recent Sea-Urchins (Echinoidea) in the Collection of the British Museum (Natural History). Oxford University Press, London, 250 pp.","Bell, F. J. (1887 a) The echinoderm fauna of the island of Ceylon. Scientific Transactions of the Royal Dublin Society, 3, 643 - 658.","Koehler, R. (1922) Echinoderma of the Indian Museum, Part 9, An account of the Echinoidea. II. Clypeastrides et Cassidulides. Trustees of the Indian Museum, Calcutta, 161 pp.","Mortensen, T. (1948 b) A Monograph of the Echinoidea IV. 2. Clypeasteroida. Clypeasteridae, Arachnoidae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagen, 471 pp.","Bell, F. J. (1882) Note on the echinoderm-fauna of the island of Ceylon, together with some observations on heteractinism, XIX. Annals and Magazine of Natural History, Series 5, 10, 218 - 225. https: // doi. org / 10.1080 / 00222938209459697","Stara, P. & Sanciu, L. (2014) Analysis of some astriclypeids (Echinoidea: Clypeasteroida). Biodiversity Journal, 5, 291 - 358."]}

Mellita aff. quinquiesperforata (Leske, 1778) Figures 12 A–J, 18 F, 19 F, G Echinodiscus quinquiesperforata Leske, 1778: 133. Scutella quinquefora Lamarck, 1816: 9. Mellita testudinata Klein, 1734: 422 –425.– Rathbun, 1879: 144. Clypeaster pentaporus Gmelin, 1791: 3189. Mellita latiambulacra H.L. Clark, 1940a: 62. Mellita lata H.L. Clark, 1940a: 437 –439. Mellita quinquiesperforata H.L. Clark, 1911: 599.– Tommasi, 1957: 21, 31–32, fig. 3, pl. 2, figs 1–2, pl. 3, figs 3–4; 1974: 187.– Matos et al. 2000: 741 –745.– Fernandes et al., 2002: 422.– Ventura et al., 2007b: 279, tab.11.2, 281, fig. 11.2f, 290, anexo 11.1.– Gondim et al., 2008: 155; 2013b: 515.– Oliveira et al., 2010: 10, fig. 4g.– Xavier, 2010: 75.– Miranda et al., 2012: 142. Mellita quinquiesperforata latiambulacra Mortensen, 1948a: 426.– Bernasconi, 1955: 63 –64, pl. IV, figs 5, 6.– Tommasi, 1959: 602, 603; 1964: 87–90.– Brito, 1962: 6; 1968: 28, pl. 14, fig. 2–3. Mellita quinquiesperforata tenuis H.L. Clark, 1940a: 442 –444, pl. 60, fig. 2, pl. 61, fig. 2. Mellita (Mellita) quinquiesperforata Brito, 1960a: 6 –7, fig. 2a–c.– Tommasi, 1966a: 25. Mellita (Mellita) quinquiesperforata var. latiambulacra Lima-Verde, 1969: 10. 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]. 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. 12A, C). Apical system monobasal, with four gonopores (Fig. 12I). Petaloid large, corresponding to a little more than 50% TL (Fig. 12A, C). Posterior petals slightly longer than anterior ones (Fig. 12A, 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. 12A, 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. 12b, 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. 12B). Peristome pentagonal and slightly anterior (Fig. 12B, D). Periproct oval to triangular, very close to peristome, deeply indenting basicoronal plate (Fig. 12B, 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. 12D). 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. 12A, B, 19F). Naked test white (Fig. 12C, D). Distribution. In Brazil from MA, PI, CE, PB, PE, AL, SE, BA, ES, RJ, SP, SC, and RS (Brito 1962; Tommasi 1964, 1966a, 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). 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 (1966a) 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)., Published as part of Gondim, Anne Isabelley, Moura, Rafael Bendayan De, Christoffersen, Martin Lindsey & Dias, Thelma Lúcia Pereira, 2018, Taxonomic guide and historical review of echinoids (Echinodermata: Echinoidea) from northeastern Brazil, pp. 1-72 in Zootaxa 4529 (1) on pages 36-38, DOI: 10.11646/zootaxa.4529.1.1, http://zenodo.org/record/2612564, {"references":["Leske, N. G. (1778) Iacobi Theodori Klein naturalis dispositio echinodermatum. Accesserunt lucubratiuncula de aculeis echinorum marinorum et spicilegium de belemnitis. Edita et descriptionibus novisque inventis et synonymis auctorem aucta. Officina Gleditschiana, Lipsiae, 278 pp.","Lamarck, J. B. (1816) Histoire Naturelle des Animaux sans Vertebres, presentant les caracteres generaux et particuliers de ces animaux, leur distribution, leur classes, leurs familles, leurs generes, et le citation des principales especes qui s'y rapportent; precedee d'une Introduction offrant la Determination des caracteres essentiells de l'animal, sa distinction du vegetal et des autres corps naturels, enfin, l'Exposition des Principes fondamentaux de la Zoologie. Tome Troisieme. Verdiere, Paris, 586 pp.","Klein, J. T. (1734) Naturalis dispositio echinodermatum. Accesseit lucubratiuncula de aculeis echinorum marinorum, cum spicilegio de belemnitis. Gedani, Schreiber, 78 pp.","Rathbun, R. (1879) A list of the Brazilian echinoderms, with notes on their distribution, etc. Transactions of the Connecticut Academy of Arts and Sciences, 5, 139 - 151.","Gmelin, J. F. (1791) Vermes. In: Gmelin, J. F. (Ed.) Caroli a Linnaei Systema Naturae per Regna Tria Naturae, Editio Decima Tertia, Aucta Reformata. Tome 1, Pars 6 (Vermes). G. E. Beer, Lipsiae, pp. 3021 - 3910. Available from: http: // www. biodiversitylibrary. org / item / 83098 5 (accessed 13 September 2016)","Clark, H. L. (1940 a) A revision of the keyhole urchins (Mellita). Proceedings of the United States National Museum, 89 (3099), 435 - 444. https: // doi. org / 10.5479 / si. 00963801.89 - 3099.435","Clark, H. L. (1911) The genera of recent clypeasteroids. Annals and Magazine of Natural History, Series 8, 7 (42), 593 - 605. https: // doi. org / 10.1080 / 00222931108692981","Tommasi, L. R. (1957) Os Equinodermos do litoral de Sao Paulo. I. Echinoidea, Crinoidea e Holothuroidea do bentos costeiro. Contribuicoes Avulsas do Instituto Oceanografico, Oceanografia Biologica, 13 (2), 19 - 44.","Matos, E., Matos, P., Corral, L. & Azevedo, C. (2000) Estrutura fina do espermatozoide de Mellita quinquiesperforata Leske (Echinodermata) do litoral norte do Brasil. Revista Brasileira de Zoologia, 17 (3), 741 - 745. https: // doi. org / 10.1590 / S 0101 - 81752000000300018","Fernandes, M. L. B., Tommasi, L. R. & Lima, E. J. B. (2002) Filo Echinodermata de Pernambuco. In: Tabarelli, M. & Silva, J. M. C. (Eds.), Diagnostico da Biodiversidade de Pernambuco. Vol. 2. Massangana, Recife, pp. 405 - 427.","Ventura, C. R. R., Verissimo, I., Lima, R. N. P., Barcellos, C. F. & Oigman-Pszczol, S. S. (2007 b) Capitulo 11. Echinodermata. In: Creed, J., Pires, D. O. & Figueiredo, M. A. O. (Orgs.), Biodiversidade Marinha da Baia da Ilha Grande. Serie Biodiversidade 23. MMA / SBF, Brasilia, pp. 273 - 290.","Gondim, A. I., Lacouth, P., Alonso, C. & Manso, C. L. C. (2008) Echinodermata da praia do Cabo Branco, Joao Pessoa, Paraiba, Brasil. Biota Neotropica, 8 (2), 151 - 159. https: // doi. org / 10.1590 / S 1676 - 06032008000200016","Oliveira, J. P., Oliveira, J. & Manso, C. L. C. (2010) Inventario da colecao de equinodermos do LABIMAR, Campus Prof °. Alberto Carvalho, Universidade Federal de Sergipe. Scientia Plena, 6 (12), 1 - 14.","Xavier, L. A. R. (2010) Inventario dos Equinodermos do Estado de Santa Catarina, Brasil. Brazilian Journal of Aquatic Science and Technology, 14 (2), 73 - 78. https: // doi. org / 10.14210 / bjast. v 14 n 2. p 73 - 78","Miranda, A. L. S., Lima, M. L. F., Sovierzoski, H. H. & Correia, M. D. (2012) Inventory of the Echinodermata collection from the Universidade Federal de Alagoas. Biota Neotropica, 12 (2), 135 - 146. Available from: http: // www. biotaneotropica. org. br / v 12 n 2 / en / abstract? inventory + bn 00812022012 (accessed 1 Agust 2012)","Mortensen, T. (1948 a) A Monograph of the Echinoidea. Vol. IV (2). Clypeasteroida. Clypeasteroidae, Arachnoididae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagem, 471 pp.","Bernasconi, I. (1955) Equinoideos y Asteroideos de la Coleccion del Instituto Oceanografico de la Univerdidad de San Pablo. Primera contribucion. Boletim do Instituto Oceanografico, 6 (1 - 2), 51 - 57. https: // doi. org / 10.1590 / S 0373 - 55241955000100002","Tommasi, L. R. (1959) Equinodermas do Estado do Rio de Janeiro. I-Crinoidea, Asteroidea, Echinoidea e Holothuroidea da regiao compreendida entre o Cabo dos Buzios e Cabo Frio. Anais da Academia Brasileira de Ciencias, 31 (4), 601 - 604.","Brito, I. M. (1962) Ensaio de Catalogo dos Equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 13, 1 - 10.","Brito, I. M. (1960 a) Clypeasteroides do Rio de Janeiro. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 3, 1 - 10.","Tommasi, L. R. (1966 a) Lista dos Equinoides recentes do Brasil. Contribuicoes do Instituto Oceanografico, Serie Oceanografia Biologica, 11, 1 - 50.","Lima-Verde, J. S. (1969) Primeira contribuicao ao inventario dos Echinodermas do nordeste Brasileiro. Arquivos de Ciencias do Mar, 9 (1), 9 - 13.","Tommasi, L. R. (1964) Observacoes sobre equinoides do Brasil. Revista Brasileira de Biologia, 24 (1), 83 - 93.","Tommasi, L. R. (1974) Observacoes sobre a distribuicao batimetrica de seis especies de equinodermes na regiao da desembocadura do Rio Doce (ES). Revista Brasileira de Biologia, 34 (2), 187 - 190.","Harold, A. S. & Telford, M. (1990) Systematics, phylogeny and biogeography of the genus Mellita (Echinoidea: Clypeasteroida). Journal of Natural History, 24, 987 - 1026. https: // doi. org / 10.1080 / 00222939000770621","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do Estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 40, 375 - 381. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Martins, I. X. & Martins de Queiroz, A. C. (2006) Echinodermos do litoral do Estado do Ceara. In: Matthews-Cascon, H. & Lotufo, T. M. C. (Eds.), Biota Marinha da Costa Oeste do Ceara. Ministerio do Meio Ambiente, Brasilia, pp. 199 - 220.","Lima, E. J. B & Fernandes, M. L. B. (2009) Diversidade de equinodermos (Echinodermata) no Estado de Pernambuco (Brasil). Revista Brasileira de Zoociencias, 11 (1), 55 - 63. Available from: https: // zoociencias. ufjf. emnuvens. com. br / zoociencias / article / view / 448 (accessed 20 November 2016)","Gondim, A. I. & Giacometti, A. C. M. (2010) Equinodermos. In: Giacometti; A. C. M. & Loebmann, D. (Org.), Biodiversidade do Litoral do Piaui. Grafica e Editora Paratodos Sorocaba Ltda, Sorocaba, pp. 129 - 133.","Coppard, S. E., Zigler, K. S. & Lessios, H. A. (2013) Phylogeography of the sand dollar genus Mellita: Cryptic speciation along the coasts of the Americas. Molecular Phylogenetics and Evolution, 69, 1033 - 1042. https: // doi. org / 10.1016 / j. ympev. 2013.05.028","International Commission of Zoological Nomenclature (ICZN) (1999) International Code of Zoological Nomenclature. 4 th Edition. The International Trust for Zoological Nomenclature, The Natural History Museum, London, xxx + 306 pp. Available from: http: // iczn. org / (accessed 12 September 2018)","Borzone, C. A. (1992) Spatial distribution and growth of Mellita quinquiesperforata Leske, 1778 on a sandy beach of southern Brazil. Neritica, Curitiba, 7 (1 - 2), 87 - 100.","Smith, S. I. (1870) Notes on American Crustacea. N o I. Ocypodoidea. Transactions of the Connecticut Academy of Arts and Sciences, 2, 113 - 176. https: // doi. org / 10.5962 / bhl. part. 20635","Mota, E. L. S. (2012) Ecologia populacional de Cassis tuberosa (Mollusca: Cassidae) em habitats costeiros no Nordeste brasileiro. M. Sc. Dissertation, Postgraduate Program in Ecology and Conservation, Universidade Estadual da Paraiba, Campina Grande, 89 pp. Available from: http: // tede. bc. uepb. edu. br / jspui / handle / tede / 2037 (Accessed 10 Dec. 2018)"]}

Leodia sexiesperforata (Leske, 1778) Figures 13 A–H, 18 G Echinodiscus sexiesperforatus Leske, 1778: 199. Mellita sexforis Rathbun, 1879: 144. Mellita sexiesperforata Jackson, 1912: 18.― Tommasi, 1959: 602, 603; 1966b: 241–242.― Magalhães et al., 2005: 63. Mellita platensis Bernasconi, 1947: 104; 1956: 124–125, pl. 2, figs 5–6. Mellita (Leodia) sexiesperforata Brito, 1960a: 8, fig. 2–d, fig. 3–b.― Tommasi, 1966a: 25.― Lima-Verde, 1969: 10.― Tommasi et al., 1988a: 5. Leodia sexiesperforata Brito, 1962: 6; 1968: 28–29, pl. 14, figs 4–5.― Alves & Cerqueira, 2000: 547.― Ventura et al., 2007a: 249.― Gondim et al., 2008: 155 (table).― Manso et al., 2008: 183, figs 5a–c.― Queiroz et al., 2011: 65 –67, figs 1af.― Miranda et al., 2012: 142.– Martins et al., 2018: 534, figs 12–13. 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]. 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. 13H), four gonopores (Fig. 13H). Petaloid small, occupying only central region of test (Fig. 13C, H). Posterior petals slightly longer than anterior ones (Fig. 13A, C). Six long, slit-like lunules (Fig. 13 A–C), anal lunule shorter than five ambulacral ones (Fig. 13A, C). Aboral primary spines strongly club-shaped, uniformly covering test (Fig. 13A). Miliary spines with sac-like structure on tip, densely scattered among primary spines (Fig. 13G). 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. 13B). Short and slender spines occur among these spines. Spines on ambitus similar to those around lunules. Peristome circular and central (Fig. 13B, D, E). Periproct small, elongate, close to peristome, slightly indenting basicoronal plate (Fig. 13B, D). Basicoronal plates reduced in size. Ambulacral basicoronal plates almost triangular. Pressure drainage channels and food grooves well-developed (Fig. 13D). Food grooves bifurcating after ambulacral basicoronal plates, surrounding lunules and reaching margin of test (Fig. 13D). 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. 13F). Colour. From yellow to brown or greenish (Hendler et al. 1995). Naked test white. 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 1966b; 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). 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 1940a; Mortensen 1948a). 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., Published as part of Gondim, Anne Isabelley, Moura, Rafael Bendayan De, Christoffersen, Martin Lindsey & Dias, Thelma Lúcia Pereira, 2018, Taxonomic guide and historical review of echinoids (Echinodermata: Echinoidea) from northeastern Brazil, pp. 1-72 in Zootaxa 4529 (1) on pages 38-39, DOI: 10.11646/zootaxa.4529.1.1, http://zenodo.org/record/2612564, {"references":["Leske, N. G. (1778) Iacobi Theodori Klein naturalis dispositio echinodermatum. Accesserunt lucubratiuncula de aculeis echinorum marinorum et spicilegium de belemnitis. Edita et descriptionibus novisque inventis et synonymis auctorem aucta. Officina Gleditschiana, Lipsiae, 278 pp.","Rathbun, R. (1879) A list of the Brazilian echinoderms, with notes on their distribution, etc. Transactions of the Connecticut Academy of Arts and Sciences, 5, 139 - 151.","Jackson, R. T. (1912) Phylogeny of the Echini with a revision of Palaeozoic species. Memoirs of the Boston Society of Natural History, 7, 1 - 491. Available from: https: // archive. org / details / phylogenyofechini 1912 jack (accessed 4 September 2016).","Tommasi, L. R. (1959) Equinodermas do Estado do Rio de Janeiro. I-Crinoidea, Asteroidea, Echinoidea e Holothuroidea da regiao compreendida entre o Cabo dos Buzios e Cabo Frio. Anais da Academia Brasileira de Ciencias, 31 (4), 601 - 604.","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do Estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 40, 375 - 381. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Bernasconi, I. (1947) Distribucion geografica de los equinoideos argentinos. GAEA, 8, 97 - 114.","Brito, I. M. (1960 a) Clypeasteroides do Rio de Janeiro. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 3, 1 - 10.","Tommasi, L. R. (1966 a) Lista dos Equinoides recentes do Brasil. Contribuicoes do Instituto Oceanografico, Serie Oceanografia Biologica, 11, 1 - 50.","Lima-Verde, J. S. (1969) Primeira contribuicao ao inventario dos Echinodermas do nordeste Brasileiro. Arquivos de Ciencias do Mar, 9 (1), 9 - 13.","Tommasi, L. R., Castro, S. M. & Sousa, E. C. P. M. (1988 a) Echinodermata coletados durante as campanhas oceanograficas do N / Oc. \" Almirante Saldanha \" no Atlantico Sul Ocidental. Relatorios Internos do Instituto Oceanografico, Universidade de Sao Paulo, 21, 1 - 11.","Brito, I. M. (1962) Ensaio de Catalogo dos Equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 13, 1 - 10.","Alves, O. F. S. & Cerqueira, W. R. P. (2000) Echinodermata das praias de Salvador (Bahia, Brasil). Revista Brasileira de Zoologia, 17 (2), 543 - 553. https: // doi. org / 10.1590 / S 0101 - 81752000000200024","Ventura, C. R. R., Verissimo, I., Nobre, C. C. & Zama, P. C. (2007 a). Filo Echinodermata. In: Lavrado, H. P. & Viana, M. S. (Eds.), Atlas de Invertebrados Marinhos da regiao central da Zona Economica Exclusiva brasileira. Parte 1. Serie Livros 25. Museu Nacional, Rio de Janeiro, pp. 217 - 258.","Gondim, A. I., Lacouth, P., Alonso, C. & Manso, C. L. C. (2008) Echinodermata da praia do Cabo Branco, Joao Pessoa, Paraiba, Brasil. Biota Neotropica, 8 (2), 151 - 159. https: // doi. org / 10.1590 / S 1676 - 06032008000200016","Manso, C. L. C., Alves, O. F. S. & Martins, L. R. (2008) Echinodermata da Baia de Todos os Santos e da Baia de Aratu (Bahia, Brasil). Biota Neotropica, 8 (3), 179 - 196. https: // doi. org / 10.1590 / S 1676 - 06032008000300017","Queiroz, V., Sales, L., Neves, E. & Johnsson, R. 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Available from: http: // www. redalyc. org / articulo. oa? id = 44920273011 (accessed 29 August 2017)","Rodriguez-Barreras, R. (2014) The shallow-water echinoids (Echinodermata: Echinoidea) of Cuba. Marine Biodiversity Records, 7, e 86, 1 - 8. https: // doi. org / 10.1017 / S 175526721400092 X","Brito, I. M. (1968) Asteroides e Equinoides do Estado da Guanabara e Adjacencias. Boletim do Museu Nacional, 260, 1 - 51.","Tommasi, L. R. (1966 b) Distribuicao geografica de alguns Equinodermas do Brasil. Revista Brasileira de Biologia, 26 (3), 239 - 246.","Mooi, R. & Peterson, D. (2000) A new species of Leodia (Clypeasteroida: Echinoidea) from the Neogene of Venezuela and its importance in the phylogeny of mellitid sand dollars. Journal of Paleontology, 74 (6), 1083 - 1092. Available from: http: // www. jstor. org / stable / 1306965 (accessed 5 Agust 2016)","Lambert, J. M. & Thiery, P. (1921) Essai de nomenclature raisonnee des Echinides. Vol. 5. Chaumont, Paris, 64 pp. [pp. 321 - 384]","Clark, H. L. (1940 a) A revision of the keyhole urchins (Mellita). Proceedings of the United States National Museum, 89 (3099), 435 - 444. https: // doi. org / 10.5479 / si. 00963801.89 - 3099.435","Mortensen, T. (1948 a) A Monograph of the Echinoidea. Vol. IV (2). Clypeasteroida. Clypeasteroidae, Arachnoididae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagem, 471 pp.","Durham, J. W. (1955) Classification of Clypeasteroid echinoids. Publications of the University of California at Los Angeles in Biological Sciences, 31, 73 - 198. Available from: https: // archive. org / stream / classificati 00 durh page / n 5 / mode / 2 up (accessed 29 August 2017)","Serafy, D. K. (1979) Memoirs of the Hourglass Cruises. V (III): Echinoids (Echinodermata: Echinoidea). Florida Department of Natural Resources, St. Petersburg, Florida, 120 pp.","Mooi, R. (1989) Living and fossil genera of the Clypeasteroida (Echinoidea: Echinodermata): An illustrated key and annotated checklist. Smithsonian Contributions to Zoology, 488, 1 - 50. https: // doi. org / 10.5479 / si. 00810282.488","Coppard, S. E. (2016) A new genus of mellitid sand dollars (Echinoidea: Mellitidae) from the eastern Pacific coast of the Americas. Zootaxa, 4111 (2), 158 - 166. http: // doi. org / 10.11646 / zootaxa. 4111.2.4","Coppard, S. E., Zigler, K. S. & Lessios, H. A. (2013) Phylogeography of the sand dollar genus Mellita: Cryptic speciation along the coasts of the Americas. Molecular Phylogenetics and Evolution, 69, 1033 - 1042. https: // doi. org / 10.1016 / j. ympev. 2013.05.028","Kier, P. M. (1963) Tertiary echinoids from the Caloosahatchee and Tamiami Formations of Florida. Smithsonian Miscellaneous Collections, 145 (5), 1 - 63. Available from: https: // repository. si. edu / handle / 10088 / 23786 (accessed 29 September 2009)","Smith, S. I. (1870) Notes on American Crustacea. N o I. Ocypodoidea. Transactions of the Connecticut Academy of Arts and Sciences, 2, 113 - 176. https: // doi. org / 10.5962 / bhl. part. 20635","McClintock, J. B. & Marion, K. R. (1993) Predation by the king helmet (Cassis tuberosa) on six-holed sand dollars (Leodia sexiesperforata) at San Salvador, Bahamas. Bulletin of Marine Science, 52 (3), 1013 - 1017.","Goodbody, I. (1960) The feeding mechanism in the sand dollar, Mellita sexiesperforata (Leske). The Biological Bulletin, 119 (1), 80 - 86. https: // doi. org / 10.2307 / 1538935","Telford, M. & Mooi, R. (1986) Resource partitioning by sand dollars in carbonate and siliceous sediments: evidence from podial and particle dimensions. The Biological Bulletin, 171 (1), 197 - 207. https: // doi. org / 10.2307 / 1541917"]}

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

Clypeaster subdepressus (Gray, 1825) Figure 10 A–H Echinanthus subdepressus Gray, 1825: 427. Clypeaster subdepressus L. Agassiz, 1836: 20.― Rathbun, 1879: 144.― Krau, 1956: 415 –416, figs 5–10, 13, 15, 17 and 19.― Tommasi, 1957: 21; 1959: 602.― Alves & Cerqueira, 2000: 547.― Ventura et al., 2007b: 279, tab.11.2; 2014: 65.– Xavier, 2010: 75.–Ventura, 2013: 106, fig. 122a, b. Stolonoclypus subdepressus Lambert & Thiéry, 1914: 301. Clypeaster (Stolonoclypus) subdepressus Mortensen, 1948a: 112 –116, pl. 23, figs 1–3, pl. 24, fig. 3, pl. 25, fig. 6, pl. 26, figs 1, 6, pl. 27, fig. 4, pl. 45, figs 4, 11, 14, 15.― Tommasi, 1957: 30 –31, figs 22―24, pl. 2, figs 3―4; 1964: 84–87; 1966a: 24.― Cherbonnier, 1959: 368 –370.― Brito, 1962: 6; 1968: 24–25, pl. 12, fig. 3.― Magalhães et al., 2005: 63. Clypeaster (Stolonoclypus) subdepressus lobatus Bernasconi, 1956: 35.― Brito, 1960a: 3 ―4; 1968: 25–26.― Tommasi, 1964: 84 –87; 1966b: 240–241. Clypeaster subdepressus subdepressus Tommasi, 1959: 602, 603.– Ventura et al., 2007b: 290, app. 11.1. Clypeaster latissimus Krau, 1956: 413 –427. Clypeaster (Stolonoclypus) subdepressus subdepressus Brito, 1960a: 3, pl. 1d.― Tommasi, 1972: 31. Material examined. Bahia: 1 spm, Salvador, 14.IX.1997 [UFBA00072]; 1 spm, 12°54′S 28°29′W, Salvador, 22.VIII.1993 [UFBA00461]. 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. 10F). Petaloid large, slightly longer than 50% of test (Fig. 10A, D). Petals large, leaf-shaped (Fig. 10A, D). Paired petals nearly closed distally (Fig. 10A, 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. 10B, E). Peristome circular, central (Fig. 10B, E, G). Food grooves unbranched, fairly deep, not reaching ambitus (Fig. 10E). Periproct oral, towards posterior margin, located between third and fifth pairs of post-basicoronal interambulacral plates (Fig. 10B, 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. 10H). 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 1948a), or olive green (Tommasi 1966a). Spines brownish-green. Preserved specimens are light brownish (Mortensen 1948a). Naked test white. Distribution. Florida, North Carolina, Mexico, Cuba, Jamaica, Dominican Republic, Panama, Colombia, Venezuela, and Brazil (Alvarado 2011; Tommasi 1966a). In Brazil from BA, RJ, SP and SC (Tommasi 1966b; 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. 2005a). 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 (1948a) 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 (1948a) 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., Published as part of Gondim, Anne Isabelley, Moura, Rafael Bendayan De, Christoffersen, Martin Lindsey & Dias, Thelma Lúcia Pereira, 2018, Taxonomic guide and historical review of echinoids (Echinodermata: Echinoidea) from northeastern Brazil, pp. 1-72 in Zootaxa 4529 (1) on pages 29-30, DOI: 10.11646/zootaxa.4529.1.1, http://zenodo.org/record/2612564, {"references":["Gray, J. E. (1825) An attempt to divide the Echinida or sea-eggs into natural families. Annales de Philosophie, 26, 423 - 431. Available from: https: // archive. org / stream / annalsofphilosop 10 lond page / 430 / mode / 2 up (accessed 26 April 2017)","Agassiz, L. (1836) Prodrome d'une Monographie des Radiaires ou Echinodermes. Memoires de la Societe des Sciences naturelles de Neuchatel, 1, 168 - 199. Available from: http: // www. biodiversitylibrary. org / item / 40297 page / 180 / mode / 1 up (accessed 2 October 2016)","Rathbun, R. (1879) A list of the Brazilian echinoderms, with notes on their distribution, etc. Transactions of the Connecticut Academy of Arts and Sciences, 5, 139 - 151.","Krau, L. (1956) A existencia de Clypeaster latissimus (Lamarck) no Brasil e consideracoes sobre Clypeaster subdepressus (Gray) (Clypeasteroida, Echinoidea). Memorias do Instituto Oswaldo Cruz, 54 (2), 413 - 416. https: // doi. org / 10.1590 / S 0074 - 02761956000200004","Tommasi, L. R. (1957) Os Equinodermos do litoral de Sao Paulo. I. Echinoidea, Crinoidea e Holothuroidea do bentos costeiro. Contribuicoes Avulsas do Instituto Oceanografico, Oceanografia Biologica, 13 (2), 19 - 44.","Alves, O. F. S. & Cerqueira, W. R. P. (2000) Echinodermata das praias de Salvador (Bahia, Brasil). Revista Brasileira de Zoologia, 17 (2), 543 - 553. https: // doi. org / 10.1590 / S 0101 - 81752000000200024","Ventura, C. R. R., Verissimo, I., Lima, R. N. P., Barcellos, C. F. & Oigman-Pszczol, S. S. (2007 b) Capitulo 11. Echinodermata. In: Creed, J., Pires, D. O. & Figueiredo, M. A. O. (Orgs.), Biodiversidade Marinha da Baia da Ilha Grande. Serie Biodiversidade 23. MMA / SBF, Brasilia, pp. 273 - 290.","Xavier, L. A. R. (2010) Inventario dos Equinodermos do Estado de Santa Catarina, Brasil. Brazilian Journal of Aquatic Science and Technology, 14 (2), 73 - 78. https: // doi. org / 10.14210 / bjast. v 14 n 2. p 73 - 78","Lambert, J. & Thiery, P. (1914) Essai de Nomenclature Raisonnee des Echinides. L. Ferriere, Chaumont, 5, 321 - 384.","Mortensen, T. (1948 a) A Monograph of the Echinoidea. Vol. IV (2). Clypeasteroida. Clypeasteroidae, Arachnoididae, Fibulariidae, Laganidae and Scutellidae. C. A. Reitzel, Copenhagem, 471 pp.","Cherbonnier, G. (1959) Echinodermes de la Guyane Francaise (Crinoides, Asterides, Ophiurides, Holothurides). (Note 4). Bulletin du Museum National d'Histoire Naturelle, 2, 367 - 372.","Brito, I. M. (1962) Ensaio de Catalogo dos Equinodermas do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 13, 1 - 10.","Magalhaes, W. F., Martins, L. R. & Alves, O. F. S. (2005) Inventario dos Echinodermata do Estado da Bahia. Brazilian Journal of Aquatic Science and Technology, 40, 375 - 381. https: // doi. org / 10.14210 / bjast. v 9 n 1. p 61 - 65","Bernasconi, I. (1956) Equinoideos y Asteroideos de la Coleccion del Instituto Oceanografico de la Univerdidad de San Pablo. Segunda contribucion. Boletim do Instituto Oceanografico, 7 (1 - 2), 119 - 148. https: // doi. org / 10.1590 / S 0373 - 55241956000100008","Brito, I. M. (1960 a) Clypeasteroides do Rio de Janeiro. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 3, 1 - 10.","Tommasi, L. R. (1964) Observacoes sobre equinoides do Brasil. Revista Brasileira de Biologia, 24 (1), 83 - 93.","Tommasi, L. R. (1959) Equinodermas do Estado do Rio de Janeiro. I-Crinoidea, Asteroidea, Echinoidea e Holothuroidea da regiao compreendida entre o Cabo dos Buzios e Cabo Frio. Anais da Academia Brasileira de Ciencias, 31 (4), 601 - 604.","Tommasi, L. R. (1972) Equinodermes da regiao entre o Amapa (Brasil) e a Florida (E. U. A.). II. Echinozoa. Boletim do Instituto Oceanografico, Sao Paulo, 21, 15 - 67. https: // doi. org / 10.1590 / S 0373 - 55241972000100002","Coppard, S. E., Kroh, A. & Smith, A. B. (2012) The evolution of pedicellariae in echinoids: an arms race against pests and parasites. Acta Zoologica, 93 (2), 125 - 148. https: // doi. org / 10.1111 / j. 1463 - 6395.2010.00487. x","Tommasi, L. R. (1966 a) Lista dos Equinoides recentes do Brasil. Contribuicoes do Instituto Oceanografico, Serie Oceanografia Biologica, 11, 1 - 50.","Alvarado, J. J. (2011) Echinoderm diversity in the Caribbean Sea. Marine Biodiversity, 41, 261 - 285. https: // doi. org / 10.1007 / s 12526 - 010 - 0053 - 0","Tommasi, L. R. (1966 b) Distribuicao geografica de alguns Equinodermas do Brasil. Revista Brasileira de Biologia, 26 (3), 239 - 246.","Serafy, D. K. (1979) Memoirs of the Hourglass Cruises. V (III): Echinoids (Echinodermata: Echinoidea). Florida Department of Natural Resources, St. Petersburg, Florida, 120 pp.","Hendler, G., Miller, J., Pawson, D. & Porter, K. (1995) Sea Stars, Sea Urchins, and Allies. Echinoderms of Florida and the Caribbean. Smithsonian Institution Press, Washington, 390 pp.","Laguarda-Figueras, A., Solis-Marin, F. A., Duran-Gonzalez, A., Gust-Ahearn, C., Buitron-Sanchez, B. E. & Torres-Veja, J. (2005 a) Equinodermos (Echinodermata) del Caribe Mexicano. Revista de Biologia Tropical, 53 (Supplement 3), 109 - 122.","Mihaljevic, M., Jerjen, I. & Smith, A. B. (2011) The test architecture of Clypeaster (Echinoidea, Clypeasteroida) and its phylogenetic significance. Zootaxa, 2983, 21 - 38.","Linnaeus, C. (1758) Systema Natura; per Regna tria Naturae, secundum Classes, Ordines. Genera. Species, cum characteribus, differentiis, synonymis, locis. Edito Decima, Reformata. Tomus 1. Impensis Direct. Laurentii Salvii, Holmire, 824 pp.","Clark, H. L. (1914) Hawaiian and other Pacific Echini. The Clypeasteridae, Arachnoididae, Laganidae, Fibulariidae, and Scutellidae. Memoirs of the Museum of Comparative Zoology at Harvard College, 46 / 1, 1 - 80. Available from: http: // archive. org / stream / memoirs 46 harv page / n 9 / mode / 2 up (accessed 19 September 2017)","Brito, I. M. (1959) Sobre uma nova Clypeaster do Brasil. Universidade do Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, 1, 1 - 4.","Krau, L. (1952) Sobre uma nova especie de Echinoidea Clypeaster oliveirai (Ordem Clypeasteroida). Memorias do Instituto Oswaldo Cruz, 50, 703 - 705. https: // doi. org / 10.1590 / S 0074 - 02761952000100023","Serafy, D. K. (1970) A new species of Clypeaster from the Gulf and Caribbean and a key to the species in the tropical Northwestern Atlantic (Echinodermata: Echinoidea). Biological Results of the University of Miami Deep-Sea Expedition, 58, 662 - 677.","Mooi, R. (1989) Living and fossil genera of the Clypeasteroida (Echinoidea: Echinodermata): An illustrated key and annotated checklist. Smithsonian Contributions to Zoology, 488, 1 - 50. https: // doi. org / 10.5479 / si. 00810282.488","Kroh, A. & Mooi, R. (2016) World Echinoidea Database. Available from: http: // www. marinespecies. org / echinoidea (accessed 3 September 2016)","del Valle Garcia, R., Solis-Marin, F. A., Abreu Perez, M., Laguarda-Figueras, F. & Duran Gonzalez, A. de la L. (2005) Catalogo de los equinodermos (Echinodermata: Crinoidea, Echinoidea, Holothuroidea) neritico-betonico del Arquipielago Cubano. Revista de Biologia Tropical, 53 (Supplement 3), 9 - 28.","Telford, M., Mooi, R. & Harold, A. S. (1978) Feeding activities of two species of Clypeaster (Echinoides, Clypeasteroida): Further evidence of clypeasteroid resource partitioning. The Biological Bulletin, 172, 324 - 336. https: // doi. org / 10.2307 / 1541712","Gladfelter, W. B. (1978) General ecology of the cassiduloid urchin Cassidulus caribbearum. Marine Biology, 47, 149 - 160. https: // doi. org / 10.1007 / BF 00395636"]}

Echinocyamus grandiporus Mortensen, 1907 Figure 11 A–I Echinocyamus grandiporus Mortensen, 1907: 33 –36, pl. 12, figs 1–5.― H.L. Clark, 1925: 165.― Tommasi, 1966a: 26, pl. 8b; 1966b: 239–240.― Tommasi et al., 1988a: 5.― Oliveira et al., 2010: 11. Material examined. Ceará: 2 spms, Canopus Bank [UFPB/ECH.1974]. 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. 11A). Petals small and open distally (Fig. 11C, E). Madreporite slightly prominent. Genital and ocular pores of same size (Fig. 11C, E). Four genital and five ocular pores form a conspicuous circle or pentagon around the madreporite (Fig. 11C, E). Primary spines elongate and lanceolate. Miliary spines shorter than primary spines, with terminal crown. Peristome central, circular, three times size of periproct (Fig. 11B, 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 (1966b), living specimens are brown-yellow. White, cream-coloured, greenish or gray (naked test) in alcohol (Fig. 11A, D). 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 1966a; 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). 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 (1966b) 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., Published as part of Gondim, Anne Isabelley, Moura, Rafael Bendayan De, Christoffersen, Martin Lindsey & Dias, Thelma Lúcia Pereira, 2018, Taxonomic guide and historical review of echinoids (Echinodermata: Echinoidea) from northeastern Brazil, pp. 1-72 in Zootaxa 4529 (1) on pages 35-36, DOI: 10.11646/zootaxa.4529.1.1, http://zenodo.org/record/2612564, {"references":["Mortensen, T. (1907) The Danish Ingolf-Expedition. Vol. IV. Part. 2. Echinoidea (II). H. Hagerup, Copenhagen, 200 pp. Available from: http: // www. biodiversitylibrary. org / page / 19694216 page / 45 / mode / 1 up (accessed 7 November 2016)","Clark, H. L. (1925) A catalogue of the recent sea-urchins (Echinoidea) in the Collection of the British Museum. Trustees of the British Museum, London. The Oxford University Press, London, 250 pp.","Tommasi, L. R. (1966 a) Lista dos Equinoides recentes do Brasil. Contribuicoes do Instituto Oceanografico, Serie Oceanografia Biologica, 11, 1 - 50.","Tommasi, L. R., Castro, S. M. & Sousa, E. C. P. M. (1988 a) Echinodermata coletados durante as campanhas oceanograficas do N / Oc. \" Almirante Saldanha \" no Atlantico Sul Ocidental. Relatorios Internos do Instituto Oceanografico, Universidade de Sao Paulo, 21, 1 - 11.","Oliveira, J. P., Oliveira, J. & Manso, C. L. C. (2010) Inventario da colecao de equinodermos do LABIMAR, Campus Prof °. Alberto Carvalho, Universidade Federal de Sergipe. Scientia Plena, 6 (12), 1 - 14.","Tommasi, L. R. (1966 b) Distribuicao geografica de alguns Equinodermas do Brasil. Revista Brasileira de Biologia, 26 (3), 239 - 246.","Borrero-Perez, G. H., Solano, D. O. & Benavides-Serrato, M. (2002) Lista revisada de los erizos (Echinodermata: Echinoidea) del mar Caribe Colombiano. Biota Colombiana, 3 (1), 141 - 148. Available from: http: // icn. unal. edu. co / publicaciones / art / 206 / 3 - N 1 / Erizos. pdf (accessed 31 March 2017)","Alvarado, J. J. (2011) Echinoderm diversity in the Caribbean Sea. Marine Biodiversity, 41, 261 - 285. https: // doi. org / 10.1007 / s 12526 - 010 - 0053 - 0","Solis-Marin, F. A., Alvarado, J. J., Abreu-Perez, M., Aguilera, O., Alio, J., Bacallado-Aranega, J. J., Barraza, E., Benavides- Serrato, M., Benitez-Villalobos, F., Betancourt-Fernandez, L., Borges, M., Brandt, M., Brogger, M. I., Borrero-Perez, G. H., Buitron-Sanchez, B. E., Campos, L. S., Cantera, J., Clemente, S., Cohen-Renjifo, M., Coppard, S., Costa-Lotufo, L. V., del Valle-Garcia, R., Diaz, Y., Diaz de Vivar, M. E., Diaz-Martinez, J. P., Duran-Gonzalez, A., Epherra, L., Escolar, M., Francisco, V., Freire, C. A., Garcia-Arraras, E., Gil, D. G., Guarderas, P., Hadel, V. F., Hearn, A., Hernandez, J. C., Hernandez-Delgado, E. A., Herrera-Moreno, A., Herrero-Perezrul, M. D., Hooker, Y., Honey-Escandon, M. 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Encope emarginata (Leske, 1778) Figures 14 A–I, 18 H, 19 E Echinodiscus emarginatus Leske, 1778: 136, figs 5–6. Scutella quadrifora Lamarck, 1816: 9. Scutella quinqueloba Eschscholtz, 1831: 18 –19, pl. 20: fig. 1. Scutella cassidulina Des Moulins, 1837: 78. Encope emarginata L. Agassiz, 1841: 47, pls. 7–8 (as E. valensii), pl. 10.― Rathbun, 1879: 145.― Krau, 1950: 358.― Bernasconi, 1955: 64 –65, pl. 3, figs 1–2.― Tommasi, 1957: 21, 34, pl. 3, figs 1–2; 1959: 602, 603; 1964: 91–92; 1966a: 26, figs 54–55, pl. 7a; 1972: 32.― Brito, 1960a: 6; 1962: 6; 1968: 26–27, pl. 13, fig. 1–2.― Lima-Verde, 1969: 10.― Tommasi et al., 1988b: 2.― Manso, 1989: 357.― Fernandes et al., 2002: 422. – Ventura et al., 2007b: 279, tab. 11.2, 290, anexo 11.1; 2014: 65.― Manso et al., 2008: 183, figs 5d–f.― Oliveira et al., 2010: 10, fig. 4f. ― Miranda et al., 2012: 142, fig. 4f. ― Gondim et al., 2013b: 515.– Guilherme et al., 2015: 209 –220.– Alitto et al., 2016: 10. Encope oblonga L. Agassiz, 1841: 53 –54, pl. 9, figs 1–7. Encope subclausa L. Agassiz, 1841: 56 –57, pl. 5. Encope valenciennesii L. Agassiz, 1841: 54 –56, pl. 7–8. Mellita lobata L. Agassiz, 1841: 44, pl. 4a, fig. 13, pl. 16, figs 4–7. Moulinia cassidulina L. Agassiz, 1841:139, pl. 22, figs 1–6. Echinoglycus frondosus Gray, 1855b: 24 –25. Encope ghiesbrechtii Belval, 1863: 419 –424. Encope (Echinodesma) emarginata Phelan, 1972: 127, 128.― Tommasi et al., 1988a: 5.― Magalhães et al., 2005: 63.― Lima & Fernandes, 2009: 57. 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]. 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. 14I). Petaloid large (Fig. 14A, C). Petals bowed and distally open (Fig. 14C). Posterior petals slightly longer than anterior ones (Fig. 14C). 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. 14B, 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. 14B, D). Periproct oval and anterior to anal lunule (Fig. 14B, 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. 14D). 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. 14A, B, 18H). Naked test white (Fig. 14C, D). Distribution. Florida, Belize, Guatemala, Panama, Caribbean Sea, Colombia, Venezuela, Brazil, and Argentina (mouth of La Plata River) (H.L. Clark 1925; Tommasi 1966a; 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, 2013b; 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 1966a). 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. 19E). 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. 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(2010) Morphological variation among seven populations of the sand dollar Encope emarginata (Leske) from the southern to northeastern coast of Brazil. In: Harris, L. H, Bottger, S. A., Walker, C. W. & Lesser, M. P. (Eds.), Proceedings of the 12 th International Echinoderm Conference, 2010. EUA, Durham, New Hampshire, pp. 287 - 291.","Coppard, S. E. & Lessios, H. A. (2017) Phylogeography of the sand dollar genus Encope: implications regarding the Central American Isthmus and rates of molecular evolution. Nature Scientific Reports, 7, 11520. https: // doi. org / 10.1038 / s 41598 - 017 - 11875 - w","Telford, M. (1981) A hydrodynamic interpretation of sand dollar morphology. Bulletin of Marine Science, 31 (3), 605 - 622. Available from: http: // www. ingentaconnect. com / content / umrsmas / bullmar / 1981 / 00000031 / 00000003 / art 00012 (Accessed 10 Dec. 2018)"]}

Placatenella complanata (Brito, 1981) Figures 1–4. 1981 Abertella complanata Brito: 3–4, figs 1–2. 2000 Abertella pirabensis Marchesini Santos—Mooi et al.: 266, in part. Diagnosis. As for the genus, which is emended from Brito's (1981: 3) original diagnosis or description (it is not stated which). Brito's information does not diagnose the species because it does not distinguish it from other members of the genus Abertella to which it was originally assigned (translated from the Portuguese): "Test flattened, flat on the oral and convex on the aboral surface, with a semicircular outline, and a marked anal indentation, a small but sharp indentation in the anterior ambulacrum, and only a sinuosity in the paired ambulacra. Petals closed, measuring approximately two-thirds of the test radius with the poriferous zones slightly narrower than the interporiferous. Central peristome, from which extend five food grooves that soon fork. Periproct ventral in the posterior indentation." Type and other material studied. The holotype, MNRJ 5460 -I, is the only originally described specimen, and is housed at the Museu Nacional, Rio de Janeiro, Brazil. We also examined other specimens assigned to this species by Brito (1986), including MNRJ 5536-I, MPEG-886-I, MPEG 1753-I, MPEG 2405-I, and DNPM 6217. These are generally of higher quality than the holotype, revealing new information concerning the species. Description. Brito (1986: 2) expanded upon the original description/diagnosis of the species based on new material, adding that there were five distinct genital pores. This is evidently an error, as Brito's (1986) own photos and original description indicate four gonopores. Our emended description is as follows. Holotype (Fig. 2A, B) approximately 25 mm TL (measured as described in Table 1, from junction of perradial suture of ambulacrum III with anterior edge of test to junction of interradial suture of interambulacrum 5 with posterior edge of test [i.e. inside notch]). Largest known specimen (Brito 1986: Fig. 1) approximately 45 mm TL, almost 55 mm in test width at widest point. Best preserved specimen (MNRJ 5536-I, Fig. 2C, D) approximately 38 mm TL (Fig. 2C, D), 57 mm test width. Ratio of width to length including lobes on either side of notch 1.17. All ensuing percentage calculations for species are from MNRJ 5536-I (Fig, 2C, D). Aboral surface slightly domed, oral surface flat. Highest point of test approximately 17% TL, located at apical system. Very sharply defined, parallel-sided, narrow posterior notch in largest specimens, depth just over 25% TL, width 10% TL, notch slightly widening near ambitus in holotype (Fig. 2A, B). Broad but shallow marginal indentations present where perradial suture meets ambitus in each ambulacrum, particularly in posterior paired ambulacra. Apical system monobasal, pentagonal (not star-shaped), 49% TL from ocular III to anterior edge of test, length 11% TL, numerous hydropores scattered over madreporic plate. Four gonopores, one in each of paired interambulacra and located at suture between madreporic plate and first adapical plates of interambulacral column. Ambulacra petaloid adapically. Posterior paired petals (I and V) noticeably longest, each extending 67% of corresponding test radius, but 39% TL; anterior paired petals (II and IV) 59% of corresponding test radius, but 33% TL; anterior unpaired (III) shortest, 68% of corresponding test radius, but 31% TL. Petal V width at widest point 16% TL, interporiferous zone 7% TL; petal IV width 18% TL, interporiferous zone 10% TL; petal III width 16% TL, interporiferous zone 9% TL. Petals lyrate, almost closed distally, with four or five trailing tube feet (sensu Mooi 1989) at distal end of each column of respiratory tube feet (Fig. 3A). Respiratory tube foot pore pairs strongly conjugated, inner pore slightly elongate or almost circular, outer pore extremely elongated, comprising about half length of pore pair, apparently subdivided by stereom septae. Four or five occluded plates present at tips of petals. At ambitus, ambulacra strongly widened, forming strip-like ambital plates that follow contour of shallow indentation at each perradius, and curving strongly adapically to form test wall along each side of posterior notch (Fig. 4). Ambulacra all in agreement with Lovén’s Rule (sensu David et al. 1996). Ambulacral basicoronal plates all similar, narrow and straight with almost parallel radial sutures on each side (Fig. 4). Interambulacra narrow and straight on oral surface, narrowing towards ambitus, but containing paired, zig-zag plates right up to madreporic plate. On oral surface, two postbasicoronal plates in each half-interambulacrum in interambulacrum 5, about four in interambulacra 1 and 4, and about five in interambulacra 2 and 3. Widest point of each interambulacrum about two thirds of way from basicoronal to ambitus, narrowing distally to about half that width so that paired interambulacra only about 14% width of adjacent ambulacra at ambitus. In each paired interambulacrum, first postbasicoronal greatly elongated, nearly five times as long as wide, about twice length of corresponding second postbasicoronal. Unpaired posterior interambulacrum 5 very narrow near basicoronal, widening distally, then narrowing as it approaches ambitus inside notch (Fig. 4). All interambulacral basicoronals broadly in contact with both corresponding first postbasicoronals. Peristome circular, relatively small, about 4% TL, with distinct perradial process in each ambulacrum extending into peristome beyond slight bulge containing sphaeridium (Fig. 3B). Anterior edge of peristome 60% TL from anterior edge of test. Periproct small, about 4% TL, situated at ambitus in anterior wall of posterior notch, between second and third pair of postbasicoronals. Aboral tuberculation homogeneous. Very slight enlargement of tubercles in oral interambulacral regions relative to those in ambulacral regions. In specimens with best preservation of surface detail, distinct tube foot pores visible in food grooves. Food grooves well developed (Figs. 2, 3, 4), restricted to oral surface, with primary bifurcation near adapical ends of ambulacral basicoronal plates. After this branch point, food grooves continuously diverging as they approach ambitus. Secondary branching faint in all specimens, apparently poorly developed. Slight depressions along perradial sutures on oral surface forming extremely shallow channels reminiscent of rudimentary pressure drainage channels. Occurrence. Known from the holotype collected from the early Miocene Pirabas Formation in the town of Castelo on the island of Fortaleza, State of Pará, Brazil, and from several additional specimens from the same formation, 5 km north of Capanema, near Colônia Pedro Teixeira, Pará, Brazil. Remarks. See the description of the family for etymology of the genus name. None of Brito's (1981, 1986) descriptions illustrates oral plate patterns, but these are crucial to interpreting the taxonomic placement of Placatenella complanata. Our investigations indicate that Placatenella is very different in many respects from Abertella pirabensis. Although these two taxa are found near each other in similar stratigraphic circumstances, it is clear that Mooi et al. 2005 were incorrect in assuming that P. complanata and A. pirabensis were conspecific, and that examination of Brito's (1986) additional material of the former was not going to shed light on the morphology of the latter. It was only through re-examination of the type material of both taxa that the major differences became apparent, particularly in oral view. For example, the periproct is on the oral surface in A. pirabensis (see below), not marginal as in Placatenella. The interambulacra of A. pirabensis are all discontinuous (Fig. 4), as is typical for all adult specimens of any species of Abertella. In all specimens of Placatenella in which plate patterns can be discerned either completely (Fig. 4), or in part (including the holotype), all interambulacra are continuous. Unusual among scutelliforms, and perhaps unlike any known species of abertellid, P. complanata typically has more oral postbasicoronal plates in the anterior paired interambulacra than in the posterior paired interambulacra. In addition, the oral tuberculation of Placatenella is more strongly differentiated in the interambulacral and ambulacral regions, and the regions between the branches of the food grooves are more depressed along the perradial suture, than in Abertella. In A. pirabensis, tuberculation of the oral surface is very uniform, and the surface itself is nearly planar, without significant perradial depressions. Placatanella also differs from A. pirabensis in petal shape. In the former, the petals are more lyrate, whereas in the latter, the two columns of pore pairs are more parallel, and remain so for a great portion of the petals' lengths. In addition, the petals of A. pirabensis vary less in length, are distinctly narrower relative to test length, and have narrower interporiferous zones than in Placatenella. The posterior notch of Placatenella is remarkable for its depth and narrowness. This condition is unmatched in other South American non-lunulates, including the abertellids and the other genus of placatenellid, Camachoaster. The only species in the Americas that has a posterior notch similar to that of P. complanata is Abertella palmeri Durham, 1957, which stands out in this respect even among the abertellids. However, A. palmeri is easily assigned to the genus Abertella, having all interambulacra markedly discontinuous, and the periproct on the oral surface. The test outline of A. palmeri is also much more strongly alate (sensu Mooi et al. 2000) than in P. complanata. There can be little question that these two forms are not closely related., Published as part of Mooi, Rich, Martínez, Sergio A., Del Río, Claudia J. & Ramos, Maria Inês Feijó, 2018, Late Oligocene - Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species, pp. 301-326 in Zootaxa 4369 (3) on pages 306-310, DOI: 10.11646/zootaxa.4369.3.1, http://zenodo.org/record/1135772, {"references":["Brito, I. M. (1981) Contribuicao a paleontologia do Estado do Para. A ocorrencia de Abertella (Echinoidea, Clypeasteroidea) na Formacao Pirabas. Boletim do Museu Paraense Emilio Goeldi, Nova Serie, Geologia, 23, 1 - 8.","Brito, I. M. (1986) Abertella complanata Brito (Echinoidea, Clypeasteroida) da Formacao Pirabas (Para, Brasil). Boletim do Museu Paraense Emilio Goeldi, Nova Serie, Geologia, 28, 1 - 4.","Mooi, R. (1989) Living and fossil genera of the Clypeasteroida (Echinoidea: Echinodermata): An illustrated key and annotated checklist. Smithsonian Contributions to Zoology, 488, 1 - 51. https: // doi. org / 10.5479 / si. 00810282.488","David, B., Mooi, R. & Telford, M. (1996) The ontogenetic basis of Loven's Rule clarifies homologies of the echinoid peristome. In: Emson, R., Smith, A. B. & Campbell, A. (Eds.), Echinoderm Research 1995. A. A. Balkema, Rotterdam, pp. 155 - 164.","Durham, J. W. (1957) Notes on echinoids. Journal of Paleontology, 31 (3), 625 - 631.","Mooi, R., Martinez, S. & Parma, S. G. (2000) Phylogenetic systematics of Tertiary monophorasterid sand dollars (Clypeasteroida: Echinoidea) from South America. Journal of Paleontology, 74 (2), 263 - 281. https: // doi. org / 10.1017 / S 0022336000031486"]}

Camachoaster maquedensis n. sp. Figures 1, 5���8. Diagnosis. As for the genus. Etymology. Named for the type locality of Punta Maqueda, Santa Cruz Province, Argentina, where the sand dollars occur in the Chenque Formation, which is exposed about 2 km south of the point. Type material studied. Material is housed at the Museo Argentino de Ciencias Naturales Bernardino Rivadavia (MACN). Holotype is MACN-Pi 5809, from shoreline deposits about 2 km south of Punta Maqueda, Santa Cruz Province, southern Argentina. These deposits form part of the Chenque Formation, lower Miocene. There are two paratypes, MACN-Pi 5859 and MACN-Pi 5860. These have the same provenience as the holotype. Description. Holotype and largest specimen (Fig. 5) 59.3 mm TL. Measurements for all known specimens given in Table 1. All ensuing percentages, calculated to facilitate comparisons with other descriptions herein, are from holotype. Aboral surface slightly domed, oral surface flat. Highest point of test 11% TL, located at apical system. Very shallow but distinct posterior notch, depth just over 5% TL, width 16% TL, notch widening significantly near ambitus. Broad, shallow marginal indentations present where perradial suture meets ambitus in each ambulacrum. Apical system monobasal, pentagonal (not star-shaped), 49% TL from ocular III to anterior edge of test, length 8% TL, numerous hydropores scattered over madreporic plate. Four gonopores, one in each of paired interambulacra and located at suture between madreporic plate and first adapical plates of interambulacral column (Fig. 8A). Ambulacra petaloid adapically. Posterior paired petals (I and V) not noticeably longer than any other petals, each extending 56% of corresponding test radius, but 30% TL; anterior paired petals (II and IV) 51% of corresponding test radius, but 29% TL; anterior unpaired (III) 56% of corresponding test radius, but 29% TL. Petal V width at widest point 12% TL, interporiferous zone 4% TL; petal IV width 12% TL, interporiferous zone 5% TL; petal III width 13% TL, interporiferous zone 6% TL. Petals lyrate, almost closed distally (Figs. 5, 8A, B), with two or perhaps as many as three trailing tube feet (sensu Mooi 1989) at distal end of each column of respiratory tube feet (Fig. 8A). Respiratory tube foot pore pairs strongly conjugated, inner pore slightly elongated, outer pore extremely elongated, comprising about half length of pore pair, apparently subdivided by stereom septae (Fig. 8A, B). Five or six occluded plates present at tips of petals (Fig. 8A). At ambitus, ambulacra greatly widened, forming strip-like ambital plates that follow contour of shallow indentation at each perradius, and curving adapically to form test wall along each side of posterior notch (Fig. 7). Ambulacra all in agreement with Lov��n���s Rule (sensu David et al. 1996). Ambulacral basicoronal plates all similar, narrow and straight with almost parallel radial sutures on each side (Figs. 6, 7). Interambulacra narrow and straight on oral surface, narrowing towards ambitus, but containing paired, zig-zag plates right up to madreporic plate. On oral surface, three or four postbasicoronal plates in each halfinterambulacrum in interambulacrum 5, four or five in in other interambulacra. Widest point of each interambulacrum about one third of way from basicoronal to ambitus, narrowing distally so that paired interambulacra about 22% width of adjacent ambulacra at ambitus. In each paired interambulacrum, first postbasicoronal greatly elongated, about four times as long as wide, about twice length of corresponding second postbasicoronal. In interambulacra 1 to 4, basicoronals broadly in contact with both corresponding first postbasicoronals. Unpaired, posterior interambulacrum 5 discontinuous, separated from basicoronal by adjacent ambulacral postbasicoronals (Figs. 6, 7). Peristome circular, relatively small, about 5% TL, with distinct perradial process in each ambulacrum extending into peristome beyond slight bulge containing sphaeridium (Fig. 8C). Anterior edge of peristome 51% TL from anterior edge of test. Periproct small, about 3% TL, on oral surface between second and third pair of postbasicoronals. Aboral tuberculation homogeneous. Very slight enlargement of tubercles in oral interambulacral regions relative to those in ambulacral regions. In specimens with best preservation of surface detail, distinct tube foot pores visible in food grooves (Fig. 8C). Food grooves well developed (Figs. 5, 7, 8C), restricted to oral surface, with primary bifurcation near adapical ends of ambulacral basicoronal plates. After this branch point, food grooves continuously diverging as they approach ambitus. Secondary branching well developed (Fig. 5). Extremely shallow depressions along perradial sutures on oral surface. Occurrence. Known only from the type locality. Geologic setting. The shallow marine deposits in which the new species was found have been described in Mooi et al. (2016). In summary, the rocks of the Chenque Formation in the Golfo San Jorge Basin (Chubut and Santa Cruz Provinces, Argentina) exposed near Punta Maqueda have been described as early Miocene (del R��o 2004). The type specimens of Monophoraster telfordi Mooi et al., 2016 were collected from the same thin bed (up to 15 cm thick) of fine-grained sandstones from which the new species of non-lunulate was recovered. Remarks. Although Camachoaster maquedensis n. sp. is found in the same strata as Monophoraster telfordi Mooi et al., 2016, the former is immediately recognizable as a different species, as well as representing a different major scutelliform clade, by its complete lack of an anal lunule. Camachoaster resembles the Abertellidae only in the possession of a shallow notch, and in the discontinuity of the posterior interambulacrum. However, the continuity of the paired interambulacra immediately invites favorable comparison with Placatenella, as do the similar shapes and dimensions of the petals. Nevertheless, Camachoaster differs from Placatenella in having the posterior interambulacrum discontinuous. Camachoaster 's combination of continuous paired interambulacra with a discontinuous posterior, unpaired interambulacrum, is unique among South American forms. A small species of Vaquerosella from California seems to have a similar oral plate pattern. However, among large scutelliforms with a posterior notch, this condition seems to be shared only with a Mexican species previously ascribed to Abertella, A. kewi Durham, 1957. The original description of A. kewi lacks figures of, or any reference to, oral plate architecture, likely due to the condition of the specimens. However, access to material from UCMP locality B8562, collected from the same locality as the types (Simojovel, Chiapas, Mexico) allows reconstruction of this architecture for comparison with Camachoaster maquedensis n. sp. (Fig. 7). This suggests that A. kewi is not an Abertella. Aspects of the oral plate pattern are reminiscent of some abertellid species, notably Abertella miskellyi Kroh et al., 2013, such as the periproct placement, the reduced posterior interambulacral basicoronal, and discontinuity of the posterior interambulacrum. However, A. kewi is unlike any abertellid in the continuity of the paired interambulacra, and the insertion of very small first postbasicoronals in either the "a" or the "b" column of each of the ambulacra. The latter situation is seen in the Pliocene Scutellaster Cragin, 1895, the Miocene and Pliocene Kewia Nisiyama, 1935, and in the Eocene Eoscutella, all from the northwest Pacific coasts of North America. The occurrence of these small postbasicoronals has not yet been analyzed in a phylogenetic context in order to determine its overall significance. The aforementioned differences between Camachoaster maquedensis n. sp. and A. kewi might suggest that they are not congeneric, but we here provisionally place A. kewi in Camachoaster as Camachoaster ? kewi pending full revision of all the relevant North American taxa associated with the Abertellidae and Placatenellidae., Published as part of Mooi, Rich, Mart��nez, Sergio A., Del R��o, Claudia J. & Ramos, Maria In��s Feij��, 2018, Late Oligocene - Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species, pp. 301-326 in Zootaxa 4369 (3) on pages 310-316, DOI: 10.11646/zootaxa.4369.3.1, http://zenodo.org/record/1135772, {"references":["Mooi, R. (1989) Living and fossil genera of the Clypeasteroida (Echinoidea: Echinodermata): An illustrated key and annotated checklist. Smithsonian Contributions to Zoology, 488, 1 - 51. https: // doi. org / 10.5479 / si. 00810282.488","David, B., Mooi, R. & Telford, M. (1996) The ontogenetic basis of Loven's Rule clarifies homologies of the echinoid peristome. In: Emson, R., Smith, A. B. & Campbell, A. (Eds.), Echinoderm Research 1995. A. A. Balkema, Rotterdam, pp. 155 - 164.","Mooi, R., Martinez, S. & del Rio, C. J. (2016) A new South American Miocene species of ' one-holed' sand dollar (Echinoidea: Clypeasteroida: Monophorasteridae). Zootaxa, 4173 (1), 45 - 54. https: // doi. org / 10.11646 / zootaxa. 4173.1.4","Durham, J. W. (1957) Notes on echinoids. Journal of Paleontology, 31 (3), 625 - 631.","Kroh, A., Mooi, R., del Rio, C. & Neumann, C. (2013) A new late Cenozoic species of Abertella (Echinoidea: Clypeasteroida) from Patagonia. Zootaxa, 3608 (5), 369 - 378. https: // doi. org / 10.11646 / zootaxa. 3608.5.5","Cragin, F. W. (1895) A new Cretaceous genus of Clypeasteridae. American Geologist, 15, 90 - 91."]}

Abertella gualichensis Mart��nez et al., 2005 Figures 1, 4. 2005 Abertella gualichensis Mart��nez et al.: 1230���1232, figs. 2���3. Diagnosis. Abertella with pronounced, marginally-directed, curved extensions of oral ambulacral plates joining proximal edges of first interambulacral postbasicoronal plates in posterior interambulacrum; posterior notch shallow but acute; paired interambulacra narrower, than in other Abertella, not narrowing near the ambitus. Type material studied. Holotype MACN-Pi 4714, paratypes MACN-Pi 4705, 4706, 4709. Description. See Mart��nez et al. (2005). Occurrence. A. gualichensis is recorded only from the earliest middle Miocene of Salina del Gualicho, R��o Negro Province, Argentina in the lower part of the Gran Bajo del Gualicho Formation (Fig. 1). Remarks. In the original description, A. gualichensis was distinguished from other Abertella largely by its broad but shallow posterior notch and its relatively narrow, non-alate test compared to other members of the genus. In addition, the separation of the oral interambulacral postbasicoronals from the basicoronal plates is less pronounced in A. gualichensis than in most other Abertella, with the exception of A. pirabensis, and the oral interambulacra of A. gualichensis do not widen and then markedly attenuate as they approach the ambitus., Published as part of Mooi, Rich, Mart��nez, Sergio A., Del R��o, Claudia J. & Ramos, Maria In��s Feij��, 2018, Late Oligocene - Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species, pp. 301-326 in Zootaxa 4369 (3) on pages 317-318, DOI: 10.11646/zootaxa.4369.3.1, http://zenodo.org/record/1135772

Genus Camachoaster n. gen. Diagnosis. As for the family, but with interambulacrum 5 discontinuous, the basicoronal widely separated from the first postbasicoronals by adjacent first postbasicoronal plates I and V. Etymology. Named in recognition of Dr. Horacio Homero Camacho (1922-2015), the first geologist and paleontologist to revisit the early 20th century work of H. von Ihering on Patagonian Mesozoic and Cenozoic faunas. Camacho worked and taught at the University of Buenos Aires for over 65 years, and was among the first to recognize the importance of the study of Patagonian echinoids in stratigraphic correlation and paleoenvironmental reconstruction. Type species. Camachoaster maquedensis n. sp., Published as part of Mooi, Rich, Mart��nez, Sergio A., Del R��o, Claudia J. & Ramos, Maria In��s Feij��, 2018, Late Oligocene - Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species, pp. 301-326 in Zootaxa 4369 (3) on page 310, DOI: 10.11646/zootaxa.4369.3.1, http://zenodo.org/record/1135772

Genus Iheringiella Berg, 1898 1898 Iheringia Lahille: 437. 1898 Iheringiella Berg: 16. 1898 Iheringiana Berg: 41. 1899 Iheringina Lahille: 395. Diagnosis. As for the family. Type species. Scutella patagonensis Desor in Agassiz & Desor, 1847, by original designation (Desor 1847: 135). Material studied. Holotype of Echinarachnius juliensis Desor in Agassiz & Desor, 1847, MCZ 102431, plus specimens attributed to I. patagonensis, including CPBA 16493-95, MACN-Pi 4586, MACN-Pi 5144, ROM 5433M, 5468M, 5469M, and two specimens sent to R.M. for identification by A. Wyss (University of California, Santa Barbara)., Published as part of Mooi, Rich, Mart��nez, Sergio A., Del R��o, Claudia J. & Ramos, Maria In��s Feij��, 2018, Late Oligocene - Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species, pp. 301-326 in Zootaxa 4369 (3) on page 322, DOI: 10.11646/zootaxa.4369.3.1, http://zenodo.org/record/1135772, {"references":["Berg, A. (1898) Substitucion de nombres genericos. Comunicaciones del Museo Nacional de Buenos Aires, 1, 16.","Agassiz, L. & Desor, P. J. E. (1847) Catalogue raisonne des especes, des genres, et des familles d'echinides. Annales des Sciences Naturelles, Troisieme Serie, Zoologie, 7, 129 - 168."]}

Genus Placatenella n. gen. Diagnosis. As for the family, but with interambulacrum 5 continuous, basicoronal 5 contiguous with both postbasicoronals; postbasicoronals in interambulacrum 5 greatly elongated and arrow-shaped, at least three times the length of more distal oral postbasicoronals in the same interambulacrum; periproct marginal inside posterior notch. Type species. Abertella complanata Brito, 1981., Published as part of Mooi, Rich, Mart��nez, Sergio A., Del R��o, Claudia J. & Ramos, Maria In��s Feij��, 2018, Late Oligocene - Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species, pp. 301-326 in Zootaxa 4369 (3) on page 306, DOI: 10.11646/zootaxa.4369.3.1, http://zenodo.org/record/1135772, {"references":["Brito, I. M. (1981) Contribuicao a paleontologia do Estado do Para. A ocorrencia de Abertella (Echinoidea, Clypeasteroidea) na Formacao Pirabas. Boletim do Museu Paraense Emilio Goeldi, Nova Serie, Geologia, 23, 1 - 8."]}

Iheringiella patagonensis (Desor in Agassiz & Desor, 1847) Figures 1, 4. Diagnosis. As for the family and genus. We note, in addition, that I. patagonensis is the only South American scutelliform clypeasteroid in which test length and width are nearly equal (all other such species being significantly wider than long). Occurrence. Arguably the most commonly encountered species of clypeasteroid in South America, I. patagonensis was evidently widespread during the late Oligocene and early Miocene in the southern Patagonian region of Argentina from the southern edge of Golfo San Jorge south to the northern edge of Tierra del Fuego (Fig. 1, inset). The recent discovery of specimens from the early Miocene Guadal Formation extends the known range of I. patagonensis westward to the Pampa Castillo region of southern Chile (Frassinetti & Covacevich, 1999; Andre Wyss, pers. comm.). Remarks. We recommend usage of the original spelling of the species name used in Desor (1847) and Agassiz & Desor (1847). Desor (1847) only listed names of taxa from Patagonia, without accompanying figures or descriptions, and these names are therefore here considered nomina nuda. In the same year, Desor (in Agassiz & Desor, 1847) provided cursory, but valid descriptions of two taxa, with locality information from the Patagonian region. Lacking figures, these descriptions have been the source of great confusion ever since. The type of one of these forms, Echinarachnius juliensis, from "Port Saint Julien" (Desor in Agassiz & Desor, 1847: 134), is known to exist in the collections of the Museum of Comparative Zoology, in Cambridge, Massachusetts. All known specimens presently identified as Iheringiella patagonensis compare favorably with this type. The types of the second form, Scutella patagonensis, reported to be from "Port Desire" (Desor in Agassiz & Desor, 1847: 135), have not been yet been found. The situation is made more difficult by our inability to locate the original material upon which Lahille (1898) based his revision. Given the complex taxonomic history of the species involved, along with the provenance of the material, a full revision of the nomenclature is warranted and pending. For comparison with other taxa described herein, we provide a map of plate architecture of the oral surface of a typical specimen from the Puerto Santa Cruz region of Argentina. This specimen compares most favorably with the type of Scutella juliensis. However, as is common for all such material, it is identified as Iheringiella patagonensis until the situation concerning the relationship between the types of S. juliensis and I. patagonensis can be resolved. What seems clear from the present evidence is that these two named entities are congeneric. Iheringiella does not have any indication of a posterior notch, making it the only described species of South American scutelliform to lack this feature., Published as part of Mooi, Rich, Martínez, Sergio A., Del Río, Claudia J. & Ramos, Maria Inês Feijó, 2018, Late Oligocene - Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species, pp. 301-326 in Zootaxa 4369 (3) on pages 322-323, DOI: 10.11646/zootaxa.4369.3.1, http://zenodo.org/record/1135772, {"references":["Agassiz, L. & Desor, P. J. E. (1847) Catalogue raisonne des especes, des genres, et des familles d'echinides. Annales des Sciences Naturelles, Troisieme Serie, Zoologie, 7, 129 - 168.","Frassinetti, D. & Covacevich, V. (1999) Invertebrados fosiles marinos de la Formacion Guadal (Oligoceno Superior-Mioceno Inferior) en Pampa Castillo, Region de Aisen, Chile. Servicio Nacional de Geologia y Mineria, Chile, Boletin, 51, 1 - 96.","Lahille, F. (1898) Notes sur le nouveau genre Iheringia. Revista del Museo de la Plata, 8, 439 - 451."]}

Genus Camachoaster n. gen. Diagnosis. As for the family, but with interambulacrum 5 discontinuous, the basicoronal widely separated from the first postbasicoronals by adjacent first postbasicoronal plates I and V. Etymology. Named in recognition of Dr. Horacio Homero Camacho (1922-2015), the first geologist and paleontologist to revisit the early 20th century work of H. von Ihering on Patagonian Mesozoic and Cenozoic faunas. Camacho worked and taught at the University of Buenos Aires for over 65 years, and was among the first to recognize the importance of the study of Patagonian echinoids in stratigraphic correlation and paleoenvironmental reconstruction. Type species. Camachoaster maquedensis n. sp.

Camachoaster maquedensis n. sp. Figures 1, 5–8. Diagnosis. As for the genus. Etymology. Named for the type locality of Punta Maqueda, Santa Cruz Province, Argentina, where the sand dollars occur in the Chenque Formation, which is exposed about 2 km south of the point. Type material studied. Material is housed at the Museo Argentino de Ciencias Naturales Bernardino Rivadavia (MACN). Holotype is MACN-Pi 5809, from shoreline deposits about 2 km south of Punta Maqueda, Santa Cruz Province, southern Argentina. These deposits form part of the Chenque Formation, lower Miocene. There are two paratypes, MACN-Pi 5859 and MACN-Pi 5860. These have the same provenience as the holotype. Description. Holotype and largest specimen (Fig. 5) 59.3 mm TL. Measurements for all known specimens given in Table 1. All ensuing percentages, calculated to facilitate comparisons with other descriptions herein, are from holotype. Aboral surface slightly domed, oral surface flat. Highest point of test 11% TL, located at apical system. Very shallow but distinct posterior notch, depth just over 5% TL, width 16% TL, notch widening significantly near ambitus. Broad, shallow marginal indentations present where perradial suture meets ambitus in each ambulacrum. Apical system monobasal, pentagonal (not star-shaped), 49% TL from ocular III to anterior edge of test, length 8% TL, numerous hydropores scattered over madreporic plate. Four gonopores, one in each of paired interambulacra and located at suture between madreporic plate and first adapical plates of interambulacral column (Fig. 8A). Ambulacra petaloid adapically. Posterior paired petals (I and V) not noticeably longer than any other petals, each extending 56% of corresponding test radius, but 30% TL; anterior paired petals (II and IV) 51% of corresponding test radius, but 29% TL; anterior unpaired (III) 56% of corresponding test radius, but 29% TL. Petal V width at widest point 12% TL, interporiferous zone 4% TL; petal IV width 12% TL, interporiferous zone 5% TL; petal III width 13% TL, interporiferous zone 6% TL. Petals lyrate, almost closed distally (Figs. 5, 8A, B), with two or perhaps as many as three trailing tube feet (sensu Mooi 1989) at distal end of each column of respiratory tube feet (Fig. 8A). Respiratory tube foot pore pairs strongly conjugated, inner pore slightly elongated, outer pore extremely elongated, comprising about half length of pore pair, apparently subdivided by stereom septae (Fig. 8A, B). Five or six occluded plates present at tips of petals (Fig. 8A). At ambitus, ambulacra greatly widened, forming strip-like ambital plates that follow contour of shallow indentation at each perradius, and curving adapically to form test wall along each side of posterior notch (Fig. 7). Ambulacra all in agreement with Lovén’s Rule (sensu David et al. 1996). Ambulacral basicoronal plates all similar, narrow and straight with almost parallel radial sutures on each side (Figs. 6, 7). Interambulacra narrow and straight on oral surface, narrowing towards ambitus, but containing paired, zig-zag plates right up to madreporic plate. On oral surface, three or four postbasicoronal plates in each halfinterambulacrum in interambulacrum 5, four or five in in other interambulacra. Widest point of each interambulacrum about one third of way from basicoronal to ambitus, narrowing distally so that paired interambulacra about 22% width of adjacent ambulacra at ambitus. In each paired interambulacrum, first postbasicoronal greatly elongated, about four times as long as wide, about twice length of corresponding second postbasicoronal. In interambulacra 1 to 4, basicoronals broadly in contact with both corresponding first postbasicoronals. Unpaired, posterior interambulacrum 5 discontinuous, separated from basicoronal by adjacent ambulacral postbasicoronals (Figs. 6, 7). Peristome circular, relatively small, about 5% TL, with distinct perradial process in each ambulacrum extending into peristome beyond slight bulge containing sphaeridium (Fig. 8C). Anterior edge of peristome 51% TL from anterior edge of test. Periproct small, about 3% TL, on oral surface between second and third pair of postbasicoronals. Aboral tuberculation homogeneous. Very slight enlargement of tubercles in oral interambulacral regions relative to those in ambulacral regions. In specimens with best preservation of surface detail, distinct tube foot pores visible in food grooves (Fig. 8C). Food grooves well developed (Figs. 5, 7, 8C), restricted to oral surface, with primary bifurcation near adapical ends of ambulacral basicoronal plates. After this branch point, food grooves continuously diverging as they approach ambitus. Secondary branching well developed (Fig. 5). Extremely shallow depressions along perradial sutures on oral surface. Occurrence. Known only from the type locality. Geologic setting. The shallow marine deposits in which the new species was found have been described in Mooi et al. (2016). In summary, the rocks of the Chenque Formation in the Golfo San Jorge Basin (Chubut and Santa Cruz Provinces, Argentina) exposed near Punta Maqueda have been described as early Miocene (del Río 2004). The type specimens of Monophoraster telfordi Mooi et al., 2016 were collected from the same thin bed (up to 15 cm thick) of fine-grained sandstones from which the new species of non-lunulate was recovered. Remarks. Although Camachoaster maquedensis n. sp. is found in the same strata as Monophoraster telfordi Mooi et al., 2016, the former is immediately recognizable as a different species, as well as representing a different major scutelliform clade, by its complete lack of an anal lunule. Camachoaster resembles the Abertellidae only in the possession of a shallow notch, and in the discontinuity of the posterior interambulacrum. However, the continuity of the paired interambulacra immediately invites favorable comparison with Placatenella, as do the similar shapes and dimensions of the petals. Nevertheless, Camachoaster differs from Placatenella in having the posterior interambulacrum discontinuous. Camachoaster 's combination of continuous paired interambulacra with a discontinuous posterior, unpaired interambulacrum, is unique among South American forms. A small species of Vaquerosella from California seems to have a similar oral plate pattern. However, among large scutelliforms with a posterior notch, this condition seems to be shared only with a Mexican species previously ascribed to Abertella, A. kewi Durham, 1957. The original description of A. kewi lacks figures of, or any reference to, oral plate architecture, likely due to the condition of the specimens. However, access to material from UCMP locality B8562, collected from the same locality as the types (Simojovel, Chiapas, Mexico) allows reconstruction of this architecture for comparison with Camachoaster maquedensis n. sp. (Fig. 7). This suggests that A. kewi is not an Abertella. Aspects of the oral plate pattern are reminiscent of some abertellid species, notably Abertella miskellyi Kroh et al., 2013, such as the periproct placement, the reduced posterior interambulacral basicoronal, and discontinuity of the posterior interambulacrum. However, A. kewi is unlike any abertellid in the continuity of the paired interambulacra, and the insertion of very small first postbasicoronals in either the "a" or the "b" column of each of the ambulacra. The latter situation is seen in the Pliocene Scutellaster Cragin, 1895, the Miocene and Pliocene Kewia Nisiyama, 1935, and in the Eocene Eoscutella, all from the northwest Pacific coasts of North America. The occurrence of these small postbasicoronals has not yet been analyzed in a phylogenetic context in order to determine its overall significance. The aforementioned differences between Camachoaster maquedensis n. sp. and A. kewi might suggest that they are not congeneric, but we here provisionally place A. kewi in Camachoaster as Camachoaster ? kewi pending full revision of all the relevant North American taxa associated with the Abertellidae and Placatenellidae.

Abertella miskellyi Kroh et al., 2013 2013 Abertella miskellyi Kroh et al.: 371���376, figs. 2���6. Diagnosis. Abertella with strongly heterogeneous interambulacral basicoronal plates (small in interambulacrum 5, largest in interambulacra 2 and 3); oral interambulacra discontinuous by involvement of two adjacent ambulacral plates (rather than one as in other Abertella) in at least one of the oral interambulacra; apparent "violation" of Lov��n's Rule in oral ambulacrum III. Type material studied. Holotype MB E.7463, paratype MB E.7462. Description. See Kroh et al. (2013). Occurrence. A. miskellyi is recorded only from the (possibly early) Miocene of Chubut Province, southern Argentina in the Camarones Formation (Fig. 1). Remarks. This species is unique among Abertella in that the posterior interambulacral basicoronal plate is much smaller than that in each of the paired interambulacral basicoronals. It is also the only Abertella in which specimens are known to have a discontinuity involving more than one postbasicoronal plate in a given halfambulacrum. An unusual feature of one of the known specimens of A. miskellyi is that ambulacral basicoronal III at first seemed longer in the "a" column than in the "b" column, suggesting violation of Lov��n's Rule (sensu David et al. 1996) until a small plate was detected, wedged between the basicoronal plates of ambulacrum III to re-establish the Lov��nian pattern (Kroh et al. 2013)., Published as part of Mooi, Rich, Mart��nez, Sergio A., Del R��o, Claudia J. & Ramos, Maria In��s Feij��, 2018, Late Oligocene - Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species, pp. 301-326 in Zootaxa 4369 (3) on page 318, DOI: 10.11646/zootaxa.4369.3.1, http://zenodo.org/record/1135772, {"references":["Kroh, A., Mooi, R., del Rio, C. & Neumann, C. (2013) A new late Cenozoic species of Abertella (Echinoidea: Clypeasteroida) from Patagonia. Zootaxa, 3608 (5), 369 - 378. https: // doi. org / 10.11646 / zootaxa. 3608.5.5","David, B., Mooi, R. & Telford, M. (1996) The ontogenetic basis of Loven's Rule clarifies homologies of the echinoid peristome. In: Emson, R., Smith, A. B. & Campbell, A. (Eds.), Echinoderm Research 1995. A. A. Balkema, Rotterdam, pp. 155 - 164."]}

Abertella pirabensis (Marchesini Santos, 1958) Figures 1, 4, 9. 1958 Karlaster pirabensis Marchesini Santos: 16–19, pl. 5: figs. 1–3. 1979 Karlaster pirabensis Santos—Brito: 736–738, pl. 3: fig. 1, pl. 4: fig. 6. 1997 Abertella pirabensis (Marchesini Santos) —Martínez & Mooi: 61. 2000 Abertella pirabensis (Marchesini Santos) (as senior synonym of Abertella complanata Brito 1986)—Mooi et al.: 266. Diagnosis. An Abertella in which the posterior notch is deep and sharply defined, the periproct just barely between the second pair of posterior interambulacral postbasicoronals, and the length of the disjunction between the interambulacral basicoronal and first postbasicoronals much less than half the length of the corresponding basicoronal in paired interambulacra. Type material studied. The holotype, DNPM 4493, is the only known specimen. Description. In general, the original description by Marchesini Santos (1958) is accurate and detailed, except for the instances discussed below. Her images do not satisfy modern standards, and we provide additional images of both surfaces, detailed views of the oral surface and wall of the notch, and a reconstructed plate map (Figs. 4, 9). The following description is of the same format as for placatenellid species discussed above, to facilitate comparisons. Holotype (Fig. 9A, B) approximately 51 mm TL (as defined in Table 1). Ratio of width to length including lobes on either side of notch 1.21. Aboral surface slightly domed, oral surface extremely flat, nearly planar and without significant sculpting or radial depressions. Highest point of test approximately 11% TL, located at apical system. Well-defined posterior notch, depth estimated to be approximately 11% TL, width approximately 10% TL at ambitus, notch widening near ambitus. Marginal indentations extremely shallow where perradial suture meets ambitus in posterior paired ambulacra, nearly absent in anterior paired ambulacra. Apical system monobasal, star-shaped, 54% TL from ocular III to anterior edge of test, length 9% TL, numerous hydropores scattered over madreporic plate. Four gonopores, one in each of paired interambulacra and located at suture between madreporic plate and first adapical plates of interambulacral column. Ambulacra petaloid adapically. Posterior paired petals (I and V) longest, but only slightly so, each extending 65% of corresponding test radius, but 34% TL; anterior paired petals (II and IV) 62% of corresponding test radius, but 33% TL; anterior unpaired (III) shortest, 61% of corresponding test radius, but 32% TL. Petal V width at widest point 13% TL, interporiferous zone 4% TL; petal IV width 13% TL, interporiferous zone 5% TL; petal III width 15% TL, interporiferous zone 6% TL. Petals not obviously lyrate, but have outer edges of each pore pair column in a given petal parallel for most of its length, petals almost closed distally, with three or four trailing tube feet at distal end of each column of respiratory tube feet. Respiratory tube foot pore pairs strongly conjugated, inner pore slightly elongate or almost circular, outer pore extremely elongated, comprising about two thirds length of pore pair, apparently subdivided by stereom septae. Five or six occluded plates present at tips of petals. At ambitus, ambulacra greatly widened, forming strip-like ambital plates, curving strongly adapically to form test wall along each side of posterior notch (Figs. 4, 9C, D). Ambulacra all in agreement with Lovén’s Rule (sensu David et al. 1996). Ambulacral basicoronal plates all similar, narrow and straight with almost parallel radial sutures on each side (Fig. 4). Interambulacra narrow and straight on oral surface, narrowing towards ambitus, but containing paired, zig-zag plates right up to madreporic plate. On oral surface, three or four postbasicoronal plates in each halfinterambulacrum in interambulacrum 5, four or five in the other interambulacra. Widest point of interambulacra 1 and 4 at first or second postbasicoronals, about one third of way to ambitus, narrowing distally to about two thirds that width so that paired interambulacra about 21% width of adjacent ambulacra at ambitus. In each paired interambulacrum, first postbasicoronal slightly elongated, two to three times as long as wide in posterior paired interambulacra, three to four times as long as wide in anterior paired interambulacra. Unpaired, posterior interambulacrum narrowing as it approaches ambitus inside notch (Fig. 4). All interambulacral basicoronals discontinuous, separated from first postbasicoronals by adjacent ambulacral first postbasicoronals, very widely so in interambulacrum 5 (Fig.4), but by far less than half length of a corresponding basicoronal in paired interambulacra. Peristome circular, relatively small, about 4% TL, with distinct perradial process in each ambulacrum extending into peristome beyond slight bulge containing sphaeridium. Anterior edge of peristome 54% TL from anterior edge of test. Periproct small, about 4% TL, situated 90% TL from anterior edge of test, just barely between second pair of postbasicoronals, with slight apparent contact with one of the first pair of postbasicoronals (Fig. 4). Aboral tuberculation homogeneous, oral tuberculation nearly so, without discernible enlargement of tubercles in oral interambulacral regions. Tube foot pores visible in food grooves (Fig. 9C). Food grooves well developed (Figs. 4, 9B, C), restricted to oral surface, with primary bifurcation near adapical ends of ambulacral basicoronal plates. After this branch point, food grooves continuously diverging as they approach ambitus. Secondary branching faint or non-existent. No significant depressions along perradial sutures on oral surface, no evidence of pressure drainage channels. Occurrence. A. pirabensis is known only from the early Miocene Pirabas Formation, Ponta de Pirabas, Ilha de Fortaleza, State of Pará, Brazil. Remarks. We suggest that all previous attempts to allocate this taxon to a genus were incorrect to some degree. We were able to confirm the original suggestion, indicated by the figures of Marchesini Santos (1958) and Brito (1979), that all interambulacra are discontinuous. This alone suggests that the attempts to place A. pirabensis in the Monophorasteridae Lahille, 1896 are misguided because all the interambulacra of all species in that family are broadly continuous, both postbasicoronal plates being in contact with the basicoronal, with a minor exception in a specimen of M. telfordi (see Mooi et al. 2016). Martínez and Mooi (1997) and Mooi et al. (2000) compared A. pirabensis with what was then known as Abertella complanata (here placed in the new genus Placatenella), and became convinced that A. complanata was a junior synonym of A. pirabensis. However, this was before the oral surface plate pattern of A. pirabensis could be confirmed to be of the Abertella configuration. Moreover, the hitherto unknown plate architecture of the oral surface of what we here recognize as Placatenella complanata has turned out to be unique among all taxa with a posterior notch because all the oral interambulacra are continuous. This is clearly unlike any Abertella, and undermines any relation between A. pirabensis and P. complanata. The conclusion is that the only known specimen of A. pirabensis is the holotype, described as the type specimen of a new genus, Karlaster, by Marchesini Santos (1958). Karlaster Marchesini Santos (1958) is not recognized by the present revision, and in this respect, we follow the work of Mooi et al. (2000) in regarding Karlaster a junior synonym of Abertella. We have been unable to support the assertion by Marchesini Santos (1958) that A. pirabensis possessed an anal lunule. Breakage along the posterior edge of the only known specimen would have rendered it impossible to make the determination that the distal parts of the lobes meet again at the ambitus. However, the curvature of the ambulacral plate sutures on either side of the posterior indentation is not consistent with that observed in all other scutelliforms that possess an anal lunule. The anal lunule of all mellitids and monophorasterids possesses walls constructed of interambulacral plates in the cross-linked pattern (sensu Seilacher 1979: Fig. 8B, Mooi et al. 2000: Fig. 4.4), a situation than can be considered diagnostic for the anal lunule. In A. pirabensis, there are no traces of the cross-linked pattern, leading us to reconstruct A. pirabensis with a notch of moderate depth through extrapolation of the aforementioned ambulacral plate curvatures (Fig. 4). Examination of the anterior wall of the notch reveals no evidence for the existence of an opening that could be interpreted as a periproct (Fig. 9D). Marchesini Santos (1958) appears to have been correct in interpreting the aperture on the oral surface as the periproct. It is neither a trace of an anal lunule, nor damage to the fossil. However, there does appear to be some damage to the test around the edges of the aperture, causing enlargement that could explain why the periproct appears to be in slight contact with the first interambulacral postbasicoronals (Fig. 9C). The periproct of A. pirabensis is between the second pair of interambulacral postbasicoronals, not solely enclosed within the first pair as in all monophorasterids. Therefore, the periproct position of A. pirabensis is like that of abertellids, but not of monophorasterids. The unequal development of the interambulacral basicoronals is also very similar to the condition seen in other Abertella, notably A. miskellyi, further undermining placement of A. pirabensis in the monophorasterids., Published as part of Mooi, Rich, Martínez, Sergio A., Del Río, Claudia J. & Ramos, Maria Inês Feijó, 2018, Late Oligocene - Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species, pp. 301-326 in Zootaxa 4369 (3) on pages 318-321, DOI: 10.11646/zootaxa.4369.3.1, http://zenodo.org/record/1135772, {"references":["Marchesini Santos, M. E. (1958) Equinoides Miocenicos da Formacao Pirabas. Departamento Nacional da Producao Mineral, Divisao de Geologia e Mineralogia, Boletim, 179, 1 - 24.","Brito, I. M. (1986) Abertella complanata Brito (Echinoidea, Clypeasteroida) da Formacao Pirabas (Para, Brasil). Boletim do Museu Paraense Emilio Goeldi, Nova Serie, Geologia, 28, 1 - 4.","David, B., Mooi, R. & Telford, M. (1996) The ontogenetic basis of Loven's Rule clarifies homologies of the echinoid peristome. In: Emson, R., Smith, A. B. & Campbell, A. (Eds.), Echinoderm Research 1995. A. A. Balkema, Rotterdam, pp. 155 - 164.","Brito, I. M. (1979) Clipeasteroides Cenozoicos do Brasil (Echinodermata, Echinoidea). Anais da Academia Brasileira de Ciencias, 54, 729 - 740.","Lahille, F. (1896) Variabilite et affinites du Monophora darwini. Revista del Museo de La Plata, 7, 411 - 444.","Mooi, R., Martinez, S. & del Rio, C. J. (2016) A new South American Miocene species of ' one-holed' sand dollar (Echinoidea: Clypeasteroida: Monophorasteridae). Zootaxa, 4173 (1), 45 - 54. https: // doi. org / 10.11646 / zootaxa. 4173.1.4","Martinez, S. & Mooi, R. (1997) \" Karlaster \" pirabensis from the Brazilian Miocene is a species of Abertella (Scutellina, Echinoidea), not a monophorasterid. 158 Congresso Brasileiro de Paleontologia, Sao Paulo, 1997, 61.","Mooi, R., Martinez, S. & Parma, S. G. (2000) Phylogenetic systematics of Tertiary monophorasterid sand dollars (Clypeasteroida: Echinoidea) from South America. Journal of Paleontology, 74 (2), 263 - 281. https: // doi. org / 10.1017 / S 0022336000031486","Seilacher, A. (1979) Constructional morphology of sand dollars. Paleobiology, 5, 191 - 221. https: // doi. org / 10.1017 / S 0094837300006527"]}

Genus Abertella Durham, 1953 1958 Karlaster Marchesini Santos, 1958, p. 16 Emended diagnosis. As for the family, see above. Type species. Scutella aberti Conrad, 1842, by original designation (Durham 1 953, p. 350). Included species. A. aberti (Conrad, 1842); A. cazonesensis Kew in Dickerson & Kew, 1917; A. ? habanensis (Sánchez-Roig, 1949); A. palmeri Durham, 1957; A. pirabensis (Marchesini Santos, 1958); A. gualichensis Martínez et al., 2005; A. dengleri Osborn & Ciampaglio, 2010; A. miskellyi Kroh et al., 2013. Remarks. Of the above included species, only A. pirabensis, A. gualichensis, and A. miskellyi are known to occur in South American outcrops. Abertella ? habanensis is provisionally listed as an Abertella until further information comes to light on this species. Abertella kewi Durham, 1957, as listed by Kroh et al. (2013), is hereby formally removed from Abertella and moved provisionally to Camachoaster as Camachoaster ? kewi, as it no longer satisfies the emended diagnosis provided in this paper., Published as part of Mooi, Rich, Martínez, Sergio A., Del Río, Claudia J. & Ramos, Maria Inês Feijó, 2018, Late Oligocene - Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species, pp. 301-326 in Zootaxa 4369 (3) on page 317, DOI: 10.11646/zootaxa.4369.3.1, http://zenodo.org/record/1135772, {"references":["Durham, J. W. (1953) Type species of Scutella. Journal of Paleontology, 27, 347 - 352.","Marchesini Santos, M. E. (1958) Equinoides Miocenicos da Formacao Pirabas. Departamento Nacional da Producao Mineral, Divisao de Geologia e Mineralogia, Boletim, 179, 1 - 24.","Conrad, T. A. (1842) Observations on a portion of the Atlantic Tertiary region, with a description of a new species of organic remains. Proceedings of the National Institute for the Promotion of Science, 2, 171 - 194.","Dickerson, R. E. & Kew, W. S. W. (1917) The fauna of a medial Tertiary formation and the associated horizons of northeastern Mexico. Proceedings of the California Academy of Sciences, Series 4, 7, 125 - 156.","Durham, J. W. (1957) Notes on echinoids. Journal of Paleontology, 31 (3), 625 - 631.","Osborn, A. S. & Ciampaglio, C. N. (2010) A new species of Abertella (Echinoidea, Scutellina) from the Late Miocene (Tortonian) Peace River Formation of Hardee County, Florida. Southeastern Geology, 47, 207 - 218.","Kroh, A., Mooi, R., del Rio, C. & Neumann, C. (2013) A new late Cenozoic species of Abertella (Echinoidea: Clypeasteroida) from Patagonia. Zootaxa, 3608 (5), 369 - 378. https: // doi. org / 10.11646 / zootaxa. 3608.5.5"]}

Genus Placatenella n. gen. Diagnosis. As for the family, but with interambulacrum 5 continuous, basicoronal 5 contiguous with both postbasicoronals; postbasicoronals in interambulacrum 5 greatly elongated and arrow-shaped, at least three times the length of more distal oral postbasicoronals in the same interambulacrum; periproct marginal inside posterior notch. Type species. Abertella complanata Brito, 1981.

Family Abertellidae Durham, 1955 Emended diagnosis. Scutelliforms with shallow to very deep, usually well-defined notch at ambitus in posterior interambulacrum; all oral interambulacra discontinuous, sometimes widely disjunct; interambulacral basicoronals large, usually more than twice length of ambulacrals. In addition, the following plesiomorphic features that are useful in that they are found in virtually all members of the family: two to five distinct trailing tube feet at end of each column of respiratory tube feet; pressure drainage channels absent, but ambulacral regions between main branches of food grooves populated by spines slightly shorter than interambulacral basicoronals; posterior interambulacral column typically narrowing towards ambitus and into posterior notch; periproct on oral surface between second, or second and third postbasicoronals. Description and remarks. Previous diagnoses of the family (Durham 1953, 1955, 1966) relied on combinations of easily discerned features otherwise not unique within the scutellines. Durham (1955) provided the following description for his new family: "Medium-sized to large, flattened; internal supports well developed; with broad ambulacral and anal indentations of margin; petals well defined, nearly closed; outer member of pore-pair greatly elongated, few primary pore-pairs outside petals; all interambulacra discontinuous on oral surface; basicoronal interambulacral plates considerably larger than ambulacral plates; periproct on oral surface; ambulacral food grooves bifurcating just outside basicoronal row; 4 genital pores." Unfortunately, not a single one of these features, taken on their own, is unique to the Abertellidae, and do not provide an unequivocal diagnosis in the modern concept of a diagnosis, which should consist of autapomorphies and, in some cases, distinctive plesiomorphic features unique to the taxon being diagnosed. In part because it is an entirely extinct group, this remains a challenge for the Abertellidae, and even the diagnosis provided above does not contain synapomorphies entirely unique to the abertellids. However, when all these features are taken in combination, the family is well circumscribed. Durham (1955) noted both the discontinuous oral interambulacra and the pronounced posterior notch. Some other South American non-lunulate taxa possess this notch as well, but are not like abertellids in other ways, including the pattern of interambulacral discontinuity. Therefore, in considering such taxa, we are left with a choice of considerably broadening the concept of the Abertellidae to include these species, or establish new supraspecific taxa. We decided to restrict the concept of the Abertellidae to include only those forms that have all the oral interambulacra discontinuous, as discussed above in the sections dealing with the excluded forms such as Placatenella and Camachoaster. The diagnosis of Abertellidae is therefore emended here to include scutellines with a shallow to very deep notch at the ambitus in the posterior interambulacrum, in combination with having all oral interambulacra strongly discontinuous in all adult specimens. Studies of specimens of the type species, A. aberti, from the Smithsonian Institution (NMNH 438168, 438169) permits inclusion of characters concerning spine morphology and distribution that seem unique to the family. Although the spines are differentiated and distributed in fields of aboral club-shaped and miliary types as in most scutelliforms, the oral surface is populated by fields of locomotory and geniculate spine types similar to those seen in mellitids, but not as strongly differentiated. There are no well-differentiated pressure drainage channels, but ambulacral regions between the main branches of the food grooves are populated by spines slightly shorter than interambulacral basicoronals. Each main branch of the food grooves beyond the primary bifurcation at the ends of the ambulacral basicoronals is always strongly developed in abertellids, but the degree of secondary branching is variable. The periproct is always situated distinctly on the flat portion of the oral surface, surrounded by the second pair of postbasicoronal interambulacral plates, or sometimes in contact with one (rarely both) of the third postbasicoronals. Features of the petals, which are lyrate, almost closed, but large (one half to three-quarters the length of the corresponding aboral ambulacrum), do not serve to distinguish abertellids from other scutelliforms. Trailing tube feet at the end of each column of respiratory tube feet are always large and distinct in abertellids, and can number up to five, but this also does not separate abertellids from other forms. The type species, A. aberti, can attain a TL of well over 150 mm, but the smallest known species, such as A. palmeri, are not known to exceed 60 mm TL. As noted and figured by Clark & Twitchell (1915), the microcanal system is well developed, as it is in several other scutelliform groups including lunulates such as the monophorasterids. Type genus. Abertella Durham, 1953, Published as part of Mooi, Rich, Mart��nez, Sergio A., Del R��o, Claudia J. & Ramos, Maria In��s Feij��, 2018, Late Oligocene - Miocene non-lunulate sand dollars of South America: Revision of abertellid taxa and descriptions of two new families, two new genera, and a new species, pp. 301-326 in Zootaxa 4369 (3) on pages 316-317, DOI: 10.11646/zootaxa.4369.3.1, http://zenodo.org/record/1135772, {"references":["Durham, J. W. (1955) Classification of clypeasteroid echinoids. California University Publications in Geological Science, 31, 73 - 198.","Durham, J. W. (1953) Type species of Scutella. Journal of Paleontology, 27, 347 - 352.","Durham, J. W. (1966) Clypeasteroids. In: Moore, R. C. (Ed.), Treatise on Invertebrate Paleontology, Part U. Echinodermata. 3 (2). Geological Society of America and The University of Kansas Press, Boulder, CO & Lawrence, KS, pp. 450 - 491. [pp. U 450 - U 491]","Clark, H. L. & Twitchell, M. W. (1915) The Mesozoic and Cenozoic Echinodermata of the United States. United States Geological Survey Monograph, 54, 1 - 341."]}

RESUME Ce travail est un recapitulatif de nos connaissances sur les diverses especes de collemboles littoraux interstitiels terrestres de la cote Est de Madagascar. Il est complete par l'etude pendant deux ans des repartitions horizontale et verticale du peuplement de collemboles de la plage d'Antalaha sur cette cote.

Echinocyamus elegans Mazzetti, 1893 Fig. 60 A���D. Echinocyamus elegans: H.L. Clark, 1923 *: 393���394; Mortensen, 1948d: 184���185. Pl. XLVI, Figs 29���31; Clark & Courtman- Stock, 1976: 242; Schultz, 2009: 558, Fig. 954.d. Material examined. MBC-A 022304; MBC-A 022305; MBC-A 022306; MBC-A 022307; MBC-A 022308; MBC- A 022309; MBC-A 022310; MBC-A 022311; MBC-A 022312; MBC-A 022313; MBC-A 022314. Identification. Test convex aborally, concave orally; petals well-developed, almost extending to ambitus, pores large, maximum of eight pore-pairs, conspicuous, pore zones as wide as interporiferous zones, not meeting distally. Specimen white. Global maximum size. Maximum test length 6 mm. Global distribution. Red Sea to West and East coast regions of South Africa; at 110 ��� 275 m depth (Mortensen 1948d; Schultz 2009). Remarks. Samples from the Iziko Museum collection extend distribution westwards towards Saldanha Bay., Published as part of Filander, Zoleka & Griffiths, Charles, 2017, Illustrated guide to the echinoid (Echinodermata: Echinoidea) fauna of South Africa, pp. 1-72 in Zootaxa 4296 (1) on pages 53-54, DOI: 10.11646/zootaxa.4296.1.1, http://zenodo.org/record/843325, {"references":["Mazzetti, (1893) Catalogo degli Echinidi del Mar Rosso, Atti della Societa dei Naturalisti e Matematici di Modena, 12 (3), 238 - 243.","Clark, H. L. (1923) The echinoderm fauna of South Africa. Annals of the South African Museum, 13 (7), 221 - 435.","Mortensen, T., (1948 d) A Monograph of the Echinoidea. IV. 2. Clypeasteroidea. Arachnoididae, Fibulariidae, Laganidae and Scutellidae. Reitzel, Copenhagen, pp. 471.","Schultz, H. (2009) Sea-Urchins II, Worldwide Irregular Deep Water Species. 1 st Edition. Scientific Publications, Hemdingen, 849 pp."]}