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Echinolampas alexandri de Loriol, 1876 Figures 13, 14A, C, 15A, 16 1876 Echinolampas alexandri de Loriol: p. 4; pl. 1: figs. 1���3. 1948a Echinolampas Alexandri de Loriol. ���Mortensen: p. 282���286; pl. 2: figs. 20���24; pl. 3: figs. 1���3; pl. 14: figs. 1, 4, 7. Material studied. Ten denuded tests: WUSL /EI/1, from Mount-Lavinia, WUSL /EI/2 and EI/110, from Negombo 1, WUSL /EI/107 and EI/108, from Baththalangunduwa 1, WUSL /EI/109, from Silavathurai, WUSL /EI/104, EI/ 105 and EI/106, from Kirinda, WUSL /EI/111 from Hambantota, Sri Lanka. Description. Size and shape ���Test moderately large, 44.3���63.0 mm TL; broad, almost circular in outline, TW 90���99% TL; distinctly inflated, TH 58���66% TL; posterior margin rounded to indistinctly pointed. Apical system ���Monobasal with four gonopores; situated anterior of centre 34���39% TL from anterior margin of test. Ambulacra ���Petals with unequal poriferous zones; poriferous zone IIIa of petal III, usually 2���5 pore pairs longer than IIIb; poriferous zones in anterior paired petals of very different length, posterior poriferous zones (IIa, IVb) in each petal 7 to 20 pore pairs longer; difference in posterior paired petals often not as pronounced, posterior poriferous zones (Ia, Vb) in each paired petal 4 to 9 pore pairs shorter than in anterior ones; poriferous zones parallel in anterior and posterior paired petals but converging distally in anterior paired petals; transversely elongated outer pores and circular inner pores in respiratory pore pairs; petal III shortest; posterior paired petals extending c.74%, anterior paired petals 73% and petal III 60 % of corresponding test radius; at their widest point, petals bear 4 to 6 tubercles across interporiferous zone; phyllodes typically consisting of inner series with up to 3 pores per column, outer series with up to 9; phyllodes contain medial double row of up to 10 sphaeridial pits. Interambulacra ���Bourrelets indistinct; oral naked zones absent. Peristome ���Small, oval, transversely elongated; length 9���10% TL and width 12���17% TL; deeply sunken with steep-sided infundibulum; located anterior of centre, approximately 37% TL from anterior margin of test. Periproct ���Inframarginal, slightly larger than peristome, length 8���13% and width 13���19% TL; situated closer to posterior margin, 2���3% TL from posterior margin of test. Geographic range. Indo-West Pacific, from Islands of West Indian Ocean (Bell 1909; Mortensen 1948d), Mauritius (Clark 1925a), Ceylon area (Clark 1917), Bay of Bengal (Koehler 1922) and East Indies (Mortensen 1948a) to Coral Sea (Miskelly 2002) and South Pacific Islands (Clark 1952). Bathymetric range. 8���365 m (Mortensen 1948a). Observed occurrence in Sri Lanka. This species typically occurs in sandy areas between patch reefs on the western, north-western, and southern coasts of Sri Lanka, at a depth range of 9���28 m (Fig. 16). This species was first reported from Sri Lanka by Clark (1915), but has not been recorded again until the present study. Remarks. Based on general shape, two groups of Echinolampas could be distinguished in Sri Lanka. The first group is characterized by a more circular outline, elongated outer pores in the petals, parallel poriferous zones in the posterior petals, and indistinct bourrelets (Fig. 14, A and C). In the second group, the outline is more ovoid, the outer pores are circular, the posterior petal pore zones converge and the bourrelets are distinct (Fig. 14, B and D). In addition, the first group does not have conspicuous oral naked zones (Fig. 15, A), while the second group does (Fig. 15, B). The first group best matches descriptions of E. alexandri alexandri, being closer to the type material of de Loriol (1876) than to later material from the Siboga Expedition described by Mortensen (1948a) under the name E. alexandri sibogae. In comparison to the syntypes of de Loriol (illustrated in Mortensen 1948a), the present material has a distinctly higher test, but this feature appears to be variable. Mooi (1990: fig. 6b) illustrated a prominent naked median zone in E. alexandri, which is also true for one of the syntypes. This feature, while showing little variation in the present material, seems to display intraspecific variation as well, and can thus only be used in conjunction with the additional characters listed above. The second group best matches E. ovata, see below., 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 19-22, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Bell, F. J. (1909) Report on the Echinoderma (other than holothurians) collected by Mr. Stanley Gardiner in the western parts of the Indian Ocean. Transactions of the Linnaean Society of London, Zoology, 13, 12 - 22. https: // doi. org / 10.1111 / j. 1096 - 3642.1909. tb 00406. x","Mortensen, T. (1948 d) New Echinoidea (Cassiduloida, Clypeasteroida): Preliminary notice. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i KObenhavn, 111, 67 - 72.","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. (1917) Hawaiian and other Pacific Echini, Echinoneidae, Nucleolitidae, Urechinidae, Echinocorythidae, Calymnidae, Pourtalesiidae, Palaestomatidae, Aeropsidae, Palaeopneustidae, Hemiasteridae, and Spatangidae. Memoirs of the Museum of Comparative Zoology at Harvard College, 46, 81 - 283.","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 a) A Monograph of the Echinoidea IV. 1. Holectypoida, Cassiduloida. C. A. Reitzel, Copenhagen, 371 pp.","Miskelly, A. (2002) Sea urchins of Australia and the Indo-Pacific. Capricornica Publications, Lindfield, 179 pp.","Clark, H. L. (1915) The echinoderms of Ceylon other than holothurians. Spolia Zeylanica, 10, 83 - 102.","Mooi, R. (1990) Living cassiduloids (Echinodermata: Echinoidea): A key and annotated list. Proceedings of the Biological Society of Washington, 1990, 63 - 85."]}

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.

Vologesia rollingstones sp. nov. (Figs. 3, 4, 5 A–B, D, F) http://zoobank.org/urn:lsid:zoobank.org:act:2A9719F4-7629-41E9-9962-B0AAEF5EEDE9 2014 Vologesia sp., Saura & García p. 306, pl. 104, Figs. 3 a–c. Types. Holotype is MB.E.6224 (Fig. 3 A–C), paratype is MB.E.6225 (Fig. 3 D–F), contained in the collections of the Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung (Berlin, Germany). Locus typicus. Coastal section near the small village of Langre, northern Cantabria (Spain). Stratum typicum. Sponge Rhythmites (Scaphites hippocrepis III Zone, lower Campanian). Derivatio nominis. Named after British Rock’ n’ Roll band The Rolling Stones for the song “Gimme Shelter (1969) and all the rest. Material. 22 complete and three fragmentarily preserved specimens (MB.E.6224–6237 and MB.E.11452- 11462). Diagnosis. Small, flattened species of Vologesia with a teardrop-shaped outline. The apical disc is positioned anteriorly and tetrabasal, with a prominent madreporite. The periproct is inframarginal. The pentagonal peristome is laterally elongated. Perioral ambulacrals show flush, short phyllodes with an inner and outer series of pore pairs. Bourrelets are only weakly developed. Description. Vologesia rollingstones sp. nov. is a small species that does not exceed 25 mm in overall test length and is characterised by a flattened, ovoid test, with its widest point approximately in the posterior lower third of the test. The margins of the flattened oral surface are well rounded. The oral surface is more convex in small-sized specimens (> 13 mm). The posterior side projects into the inframarginal periproct, thus, the test is slightly teardrop shaped in dorsal view. The ambitus is positioned in the lower third of test height. The aboral surface is generally flattened. The position of the apical disc is variable, ranging between 32 and 42% of test length to the anterior margin; in general, it is closely situated to the anterior edge (35% of test length). The apical disc is tetrabasal, the madreporite being formed by a large genital plate 2, which occupies the centre of the apical disc, the remaining genital plates being reduced in size (Fig. 5 F). Adapically, the ambulacra are petaloid, having 46 pore pairs in the posterior petals and 41 in the anterior petals, respectively, at a test size of c. 19 mm and 34 in the posterior petals, 31 in the anterior petals, respectively, at a test length of c. 14 mm. Petals are relatively short, the anterior ones ranging between 17 and 26% of test length (average 22%) and posterior ones between 21 and 29% (average 26%). The shape of the petals varies from straight to slightly bowed. Petals bear conjugate anisopores. The peristome is small, laterally elongated and pentagonal in outline. The distance from the peristome to the anterior margin is 25–35% of test length (average 30%). The bourrelets are only weakly developed, which is why only an imperfect floscelle structure is developed. Short phyllodes, which are flush to the oral surface, are simple, being composed of an outer series with 7 pores and an inner series of 2 pores (see Fig. 5 D); the inner pores are carried by occluded plates; small buccal pores are present. The laterally elongated, oval periproct has an inframarginal position. The tuberculation is fine and dense, with tubercles decreasing in size in aboral direction. A narrow, pitted band without tubercles extends from the peristome to the periproct. Discussion. So far, the genus Vologesia comprised only three species; V. tataosi, V. rawdahensis and V. toucasi. From V. rawdahensis (for a detailed description, see Smith, 1995), V. rollingstones sp. nov. can be distinguished by its smaller size, a more rounded ambitus, a more anteriorly situated apical disc and shorter petals, while the peristome is further to the anterior margin (21–27% of test length in V. rawdahensis). In addition, a third inner series of pores in the phyllodes is partially developed in V. rawdahensis, while this is absent from V. rollingstones sp. nov.; the inner pores in the phyllodes of the latter are dispersed, in contrast to V. rawdahensis. Ali (1989) noted that V. rawdahensis was similar to V. tataosi and could only be distinguished from the latter by a more elongate and pointed posterior test with a smaller peristomal opening. Thus, V. rawdahensis and V. tataosi might turn out to be conspecific (see discussion in Smith, 1995). Other species that had previously been placed in Vologesia by Lambert and Thiéry (1909 –1925), including Conoclypus rhothomagensis d’Orbigny, 1855, Clypeolampas ovum de Grateloup, 1836, Clypeolampas conicus Arnaud, 1877, Clypeolampas gossaviensis Lambert, 1907, Clypeolampas helios Noetling, 1897 and Echinolampas acuta Des Moulins, 1837), are better assigned to Hungaresia or Clypeolampas (compare Smith, 1995). They can all be distinguished from V. rollingstones sp. nov. by their inflated tests, longer petals, more centrally positioned peristome and swollen bourrelets. The degree of bourrelet development cannot be identified in C. gossaviensis [treated by Smith (1995) as a nomen dubium] and C. conicus on the basis of their original descriptions.

Genus Vologesia Cotteau & Gauthier, 1895 Type species. Vologesia tataosi Cotteau & Gauthier, 1895, p. 66, by monotypy. Diagnosis. Apical disc tetrabasal with large madreporite, other genital plates reduced in size; peristome close to anterior margin, 20–35% of test length away from it; bourrelets only very weakly developed; phyllodes with 2–3 series of pores; peristome pentagonal and laterally elongated (modified from Smith, 1995, p. 190). Remarks. Vologesia, Hungaresia Szörényi, 1955 and Clypeolampas Pomel, 1869 are closely similar; according to Smith (1995), there are few stable characters to differentiate between these genera. For example, Clypeolampas and Hungaresia have subcentrally positioned peristomes, while in Vologesia it is in a more anterior position. Smith (1995) also indicated the peristome to be placed at 20–30% of test length from the anterior border as diagnostic feature in Vologesia. These values were based on V. toucasi (Lambert, 1911) and V. rawdahensis Ali, 1989. This range can now be extended to 20–35% on the basis of our description of Vologesia rollingstones sp. nov., illustrating the position of the peristome to be more variable than previously suggested by Smith (1995). The presence of bourrelets (i.e., swollen perioral interambulacral plates) appears to be a more reliable character for differentiating between Clypeolampas, Hungaresia and Vologesia; in the last-named genus bourrelets are missing. Furthermore, aboral pustular granules are present only in Clypeolampas (see Smith, 1995). Occurrence. Vologesia toucasi is known from the upper Santonian of Aude, southern France (Lambert, 1911) and the upper Santonian of south central Pyrenees, Lleida province, north-eastern Spain (Gallem, personal communication), V. tataosi from the “Senonian of Louristan, Iran (Cotteau & Gauthier, 1895) and V. rawdahensis from the mid-Maastrichtian of the United Arab Emirates/Oman border region (Ali, 1989).

Vologesia toucasi (Lambert, 1911) (Figs. 5 C, E, G, 7) 1911 Clypeolampas toucasi Lambert, p. 155, pl. 3, Figs. 1–3. 2014 Hungaresia toucasi (Lambert), Saura & García p. 306, pl. 104, Figs. 1 a–c. Type. Holotype, by monotypy, is the specimen illustrated by Lambert (1911, pl. 3, Figs. 1–3), in the Lambert Collection at the Muséum national d’Histoire naturelle, Paris ( MNHN.F.J01723) (not seen). It is from Sougraigne in the Corbières (Aude, France), having been collected from the “couches Hippurites... avec Actinocamax Grossouvrei. Lambert (1911) considered those strata to be of Campanian age. According to the revised stratigraphy by Christensen et al. (1993), these levels are of late Santonian age (Placenticeras polyopsis / Texanites gallicus Zone). Material. A single near-complete and four fragmentarily preserved specimens (MB.E.11463-11467) from the top of Bed 200, upper Santonian (Placenticeras polyopsis Zone), deposited in the Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung (Berlin, Germany). Diagnosis. Small species with a domed to conical test and a flattened oral surface. The tetrabasal apical disc, with a prominent madreporite, is in a central position. The pentagonal, laterally elongated peristome is positioned close to the anterior margin. The phyllodes are short and slightly sunken, with an inner and an outer series of pore pairs; bourrelets are weakly developed. Description. Vologesia toucasi is a small species with a maximum test length of c. 35 mm. The oval tests are domed to conical in profile, with its longest axis along the anterior-posterior direction in cross section. The ambitus is positioned close to the oral surface, in the lower quarter of test height. The apical disc is positioned at 47% of test length to the anterior margin. The apical disc is not completely preserved; it was presumably tetrabasal, with the madreporite formed by the large genital plate 2, occupying the centre of the apical disc, while other genital plates are much reduced in size (Fig. 5 G). Petaloid ambulacra are present adapically; pore counts are not available due to poor preservation. The straight petals are relatively short, not reaching below the upper third of test height. Conjugate anisopores can be recognised within the petals. The small peristome is laterally elongated and has a pentagonal outline. It is positioned anteriorly, with a distance to the anterior margin of 29% of test length (at a length of 29.8 mm). A weakly pronounced floscelle structure can be seen; the bourrelets are only weakly swollen, in combination with only slightly sunken phyllodes. The phyllodes are short, with an outer series of 10–11 pores, including small buccal pores, and an inner series of 3–4 pores (see Fig. 4 C). The oval periproct is inframarginal. The tubercules generally are small, largest on the oral surface and decreasing in size in aboral direction. A narrow, granular band without tubercles extends from the peristome to the periproct. Discussion. Our record of V. toucasi from the upper Santonian of Langre is in accordance with the stratigraphic position of the type series from the Corbières, southern France and extends the known geographic range of the species to Spain. Vologesia toucasi differs from V. rollingstones sp. nov. by its subconical to conical profile and in phyllode structure that generally reveals a floscelle on account of more pronounced bourrelets. This species has a more inflated and circular shape in comparison to V. rawdahensis and V. tataosi. In addition, the apical disc is more central. Lambert (1919) mentioned Vologesia acuta (Des Moulins, 1837) from the Santander region. This is better referred to Clypeolampas (see Smith, 1995) and differs to Vologesia by a more centrally positioned peristome and better-developed bourrelets., Published as part of Schlüter, Nils & Wiese, Frank, 2017, Late Cretaceous species of Vologesia (Echinoidea, Cassiduloida) from northern Spain, pp. 261-270 in Zootaxa 4306 (2) on page 267, DOI: 10.11646/zootaxa.4306.2.6, http://zenodo.org/record/843798, {"references":["Lambert, J. (1911) Etude sur les Echinides cretaces de Rennes les Bains et des Corbieres. Bulletin de la Societ d'Etudes scientifiques de l'Aude, 22, 1 - 120.","Christensen, W. K., Bilotte, M. & Hansotte, M. (1993) Additional Late Cretaceous belemnitellids from the Corbieres, French Pyrenees and ammonite biostratigraphy of the Santonian. Cretaceous Research, 14, 101 - 106. https: // doi. org / 10.1006 / cres. 1993.1008","Lambert, J. (1919) Echinides fossiles des environs de Santander recueillis par M. L. Mengaud. Annales de la Societ Linneenne de Lyon, Nouvelle Serie, 66, 1 - 32.","Smith, A. B. (1995) Late Campanian-Maastrichtian echinoids from the United Arab Emirates - Oman border region. Bulletin of the British Museum (Natural History), Geology, 51, 121 - 240."]}

A new species of the echinolampadid genus Vologesia, V. rollingstones sp. nov., is recorded from the lower Campanian of northern Cantabria (Spain). Additionally, material of Vologesia toucasi, of late Santonian age, is described from the same area; this species had previously been noted only for southern France.

Echinolampas crassa (Bell, 1880) Fig. 55 A–C. Palaeolampas crassa, nov. sp.: Bell, 1889: 43–45. Pl. IV. Palaeolampas crassa: Bell, 1904: 172: Döderlein, 1906: 236; Döderlein, 1910: 248; Mortensen, 1948d: 293 –295. Pl. III, Figs 4–7. Pl. IV, Figs 11–12, Pls. IX, X, XIV, Figs 8–9. Echinolampas crassa: H.L. Clark, 1923: 397; H.L. Clark, 1924: 11 –12; H.L. Clark, 1925a: 182 –185; Branch et al., 2010: 236, Fig. 106.7; Schultz, 2009: 521, Figs 877–879. Echinolampas (Palaeolampas) crassa: Clark & Courtman-Stock, 1976: 244. Material examined. A22340; MBC-A 022340; MBC-A 022341; MBC-A023373; MBC-A077887; MBC- A077885; MBC-A077886; A22267; A22268; A22272; A22270; A23433; A22292. Identification. Test moderate to large, aborally raised, and slightly elongated. Apical disc anteriorly positioned, monobasal, with four gonopores. Peristome wider than long, sub-centrally positioned; bourrelets and phyllodes distinct; five well-developed petals, extending to edge, distally opened; anterior-lateral petals being the shortest, sub-equal. Posterior interambulacrum sometimes having a naked space, not raised towards peristome, tuberculation and spines dense, uniform, and becoming sparse towards peristomial area. Test and spine both brown. Global maximum size. Maximum test length 125 mm. Global distribution. Endemic to South coast of South Africa, at 15–500 m depth (Döderlein 1910; Clark & Courtman-Stock 1976). Remarks. The most stricking feature that distinguishes Echinolampas crassa and Tropholampas loveni, is the lack of obvious sexual dimorphism in Echinolampas crassa., 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 49-50, DOI: 10.11646/zootaxa.4296.1.1, http://zenodo.org/record/843325, {"references":["Bell, F. J. (1880) On Palaeolampas, a new genus of the Echinoidea. Proceedings of the Zoological Society, London, 1880, 43 - 49.","Bell, F. J. (1904) The Echinodermata found off the coast of South Africa. I. Echinoidea. Marine Investigations in South Africa, 3, 167 - 175.","Doderlein, L. (1906) Die Echinoiden der deutschen Tiefsee-Expedition. Wissenschaftliche Ergebnisse der Deutschen Tiefsee- Expedition, 5, 61 - 290.","Doderlein, L. (1910) Asteroidea, Ophiuroidea, Echinoidea. In: Schultze, L. (Ed.), Zoologische und Anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Sudafrika ausgefuhrt 1903 - 1905 (Jenaer Denkschrift 16), 4 (1), pp. 246 - 258.","Mortensen, T., (1948 d) A Monograph of the Echinoidea. IV. 2. Clypeasteroidea. Arachnoididae, Fibulariidae, Laganidae and Scutellidae. Reitzel, Copenhagen, pp. 471.","Clark, H. L. (1923) The echinoderm fauna of South Africa. Annals of the South African Museum, 13 (7), 221 - 435.","Clark, H. L. (1924) Echinoderms from the South African Fisheries and Marine Biology Survey. Part I. Sea-Urchins (Echinoidea). Report, Fisheries and Marine Biological Survey, Union of South Africa, 4 (1), 1 - 33.","Clark, H. L. (1925 a) A Catalogue of the Recent Sea-urchins (Echinoidea) in the Collection of the British Museum (Natural History). Trustees of the British Museum, London, 250 pp.","Branch, G. M., Griffiths, C. L., Branch, M. L. & Beckley, L. (2010) Two Oceans: a Guide to the Marine Life of southern Africa. Struik Nature, Cape Town, 456 pp.","Schultz, H. (2009) Sea-Urchins II, Worldwide Irregular Deep Water Species. 1 st Edition. Scientific Publications, Hemdingen, 849 pp.","Clark, A. & Courtman-Stock, J. (1976) The Echinoderms of Southern Africa. Publication no. 766. British Museum of Natural History, London, 277 pp."]}

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