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Metalia sp. Figures 68–71 Material studied. Four denuded tests: WUSL /EI/85 and EI/86, from Kamburugamuwa and WUSL /EI/138 and EI/ 139 from Negombo 1, Sri Lanka; one complete specimen, WUSL /EI/85 and three broken specimens. Description. Shape and size —Test large, robust 121.0– 144.3 mm TL; width 89–92% TL; height approximately 50% TL; test oval in outline when viewed aborally; no ambital notch in ambulacrum III. Apical system —Ethmolytic, with four large circular gonopores; madreporite expanding beyond posterior oculars; length 4–5% TL; situated anteriorly, 40–43% TL away from anterior margin. Ambulacra —Ambulacrum III narrow and flush with test, with pore pairs not differentiated to support respiratory tube feet; paired ambulacra distinctly petaloid; petals deeply sunken towards ambitus, straight; each anterior paired petal extending 92–96% of corresponding test radius; anterior paired petals forming angle of 135–145°; anterior and posterior paired petals about equal, length 39–42% TL, width 5–6% TL; angle between anterior paired petals slightly larger than angle between posterior paired petals; interporiferous zones narrow, of equal width, c. 1% TL; pores large, about equal in size, connected by deep groove; adorally well-developed phyllodes occur, featuring 17–18 pores in the anterior paired ambulacra, 8 in posterior paired ambulacra, and 10–11 in ambulacrum III. Tuberculation —Aboral surface densely packed with primary tubercles; larger tubercles inside peripetalous fasciole, bordering ambulacrum III and scattered over posterior interambulacra, usually arranged in short, oblique bands; primary tubercles on oral side larger than aboral ones; primary tubercles in plastron much smaller than those of oral interambulacra 1, 2, 3 and 4. Fasciole —Peripetalous fasciole well-developed, closely outlining petals; fasciole strongly indented between petals; subanal fasciole shield-shaped, wider than long; length 16–18% TL, width 22–27% TL; tapering to one point at posterior edge of plastron; upper part shallowly indented; usually four pore pairs enclosed on each side of subanal fasciole; anal branches extending aborally to points even with midpoint of periproct. Plastron —Narrow, elongate, width 19–20% TL, slightly keeled. Peristome —Kidney-shaped; situated anteriorly, c. 21% TL from anterior margin of test; about twice as wide as long; moderately large, length c. 6% TL, width c. 14% TL; from labrum pointed and overhanging peristome. Periproct —Vertically elongate; pointed aborally and orally; large, length 10–11% TL, width 5–7% TL; region just oral to periproct concave. Observed occurrence in Sri Lanka. Specimens were collected at 20–25 m from sand patches between patch reefs in Kamburugamuwa, southern coast, and from 27–28 m at Negombo, western coast of Sri Lanka (Fig. 71). Remarks. As noted by one of us (R. Mooi, unpublished and Askin & Mooi 2017), these specimens differ from the 10 known, extant congeners, M. angustus de Ridder, 1984, M. dicrana H.L. Clark, 1917, M. kermadecensis Baker & Rowe, 1990, M. latissima H.L. Clark, 1925, M. nobilis Verrill, 1867, M. persica (Mortensen, 1940), M. robillardi (de Loriol, 1876), M. spatagus (Linnaeus, 1758), M. sternalis (Lamarck, 1816), and M. townsendi (Bell, 1904). The new species lacks a notch in ambulacrum III, a condition that strongly differs from that in the following species: M. angustus, M. kermadecensis, M. latissima, M. persica, and M. robillardi. The lack of coalescence between the aboralmost regions of the posterior paired petals is very different from the situation seen in M. sternalis. M. townsendi differs in having a subcentral position of the apical system, and much lower anterior end of the test. Forms that lack the anterior notch, lack the coalescence, and have the apical system at least somewhat anterior, as in the new species, include M. dicrana, M. nobilis, and M. spatagus. M. nobilis, the only Metalia known from the Americas, has small petals relative to its congeners. In addition, the periproct in Metalia n. sp. is directed downwards and is not fully visible from the aboral side. This is not only unlike M. nobilis, it is not like M. spatagus either, which has a more vertical posterior region with at least part of the periproct visible in aboral view. This is also true for M. dicrana, which is distinguished from all other Metalia by its peculiar double point along the sternum just adoral to the subanal fasciole (Mooi, unpubl. data). Therefore, the specimens are very unlike M. dicrana. The Sri Lankan specimens of this new form have a very distinctive, concave region below the periproct which has not been noted in other Metalia species (Fig. 70), although it does occur to some degree in some of the species (Mooi, unpubl. data). As discovered by Askin & Mooi (2017) using molecular techniques, it would appear that this is a new species. It was depicted by van Noordenburg (2008), but remains unnamed pending further work on the abundant material at the California Academy of Sciences and the Natural History Museum, Vienna. Variation in the path of the peripetalous fasciole was noted in collected specimens. WUSL/IE/E85 shows shallow aborally-curved segments in interambulacrum 4, and one sharply defined indentation in interambulacrum 1 (Fig. 68, A)., 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 78-83, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Askin, R. & Mooi, R. (2017) The heart of the problem: Phylogenetic systematics of burrowing sea urchins in the genus Metalia (Echinoidea: Spatangoida). Integrative and Comparative Biology, 57 (Supplement 1), e 7.","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.","Mortensen, T. (1940) Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea, and Echinoidea. Danish Scientific Investigations in Iran, Part 2, 55 - 112.","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.","van Noordenburg, H. (2008) Sea Urchins of the Philippines. Part I. The Irregulars. Artificial Harmonics, Utrecht, 153 pp."]}

Metalia sternalis (Lamarck, 1816) Figures 66, 67 1816 Spatangus sternalis Lamarck: p. 31. 1951 Metalia sternalis (Lamarck).—Mortensen: p. 535–537; pl. 64: figs. 13, 17, 19, 22. Material studied. Seven denuded tests: WUSL /EI/70 and EI/71, from Baththalangunduwa 2 and WUSL /EI/122, EI/123, EI/124, EI/125, and EI/126 from Silavathurai, Sri Lanka. Description. Shape and size —Test large, 60.6–84.5 mm TL; width 89–92% TL; height 55–59% TL; outline of test elliptical to slightly angular in aboral view, with anterior slope more steeply angled forward than the posterior slopes to the periproct; slightly truncated posteriorly; anterior notch deep and broad. Apical system —Ethmolytic, with four large circular gonopores; posterior paired pores about 1.5 times larger than anterior ones; madreporite expanding beyond posterior oculars; situated anteriorly, 30–37% TL from anterior margin. Ambulacra —Ambulacrum III narrow and depressed aborally; paired ambulacra distinctly petaloid, deeply sunken towards margin, broad, elongate, somewhat rounded and closed distally; anterior paired petals extend laterally forming an angle of c. 175° from each other, c. 65% of TW; petal III length c. 33% TL (SD=2.3, N=7); interporiferous zone broad, of equal width in each petal; respiratory pores large, pores within one pair about equal in size, connected by deep groove; posterior paired petals 1.25 times longer than anterior ones; adapically posterior paired petals narrow; at about one quarter petal length, inner pore pairs rudimentary; posterior paired petals coalescing, elongate, slightly diverging posteriorly; at about half petal length, two poriferous zones are equal width, up to petal tip. Interambulacra —Aboral interambulacrum 5 raised, slightly convex to flat; but not keeled, not projecting above periproct; posterior end almost vertically truncated; interambulacra densely packed with perforate crenulate primary tubercles, larger tubercles inside peripetalous fasciole, particularly along petals; adapically no primary tubercles in interambulacrum 5 between posterior paired petals. Fasciole —Peripetalous fasciole well-developed, closely outlining petals, with indentations between petals; subanal fasciole broad, shield-shaped, wider than long; length 26–29% TL and width 31–39% TL; left and right halves joining at point near posterior edge of plastron; five to six pore pairs inside subanal fasciole on each side; anal branches present, extending aborally to points even with adapical margin of periproct. Plastron —Narrow, elongate, width 25–31% TL, slightly keeled. Peristome —Kidney-shaped; very close to anterior edge, 19–27% TL from anterior margin of test; about twice as wide as long; moderately large, length 8–10% TL, width 17–19% TL; labrum narrow and wide. Periproct —Elongate along anterior-posterior axis; pointed above and below; large, length 15–20% TL and width 8–11% TL. Geographic range. Indo-West Pacific, from Islands of West Indian Ocean (Bell 1884), Mauritius (De Loriol 1883), East Africa & Madagascar (Mortensen 1948c), Red Sea (Fourtau 1904), India (Bell 1888), Bay of Bengal (Clark 1925a), East Indies (de Meijere 1904), North Australia (Clark 1925a) to NSW, Australia (Miskelly 2002), Philippine Islands (Mortensen 1948e; Mooi & Munguia 2014), China & South Japan (Döderlein 1885), South Pacific Islands (Agassiz 1872) and Hawaiian Islands (Agassiz 1872). Bathymetric range. Littoral zone to 90 m (Mortensen 1951). Observed occurrence in Sri Lanka. Specimens were collected from a sandy bottom with sea grass beds in Silavathurai and Baththalangunduwa, north-western coast of Sri Lanka, at depth range of 10–20 m (Fig. 67). This species was first recorded from Sri Lanka by Clark (1915). Remarks. M. sternalis can be distinguished from Metalia sp. by having a coalescence between the aboralmost regions of the posterior paired petals., 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 75-78, DOI: 10.11646/zootaxa.4541.1.1, http://zenodo.org/record/2617460, {"references":["Bell, F. J. (1884) Echinodermata. Report on the zoological collections made in the Indo-Pacific Ocean during the voyage of H. M. S. \" Alert \", London, 1881 - 2, 117 - 177, 509 - 512, pls. 8 - 17 & 45.","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.","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.","Fourtau, R. (1904) Contribution a l'etude des Echinides vivant dans le Golfe de Suez. Bulletin de l'Institut Egyptien, IV, Serie, 5, 407 - 446.","Bell, F. J. (1888) Report on a collection of echinoderms made at Tuticorin, Madras, by Mr. Edgar Thurston, C. M. Z. S., Superintendent, Government Central Museum, Madras. Proceedings of the Zoological Journal of London, 1888, 383 - 389. https: // doi. org / 10.1111 / j. 1469 - 7998.1888. tb 06719. x","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 Meijere, J. C. H. (1904) Die Echinoidea der Siboga-Expedition. Siboga Expeditie, 43, 1 - 252.","Miskelly, A. (2002) Sea urchins of Australia and the Indo-Pacific. Capricornica Publications, Lindfield, 179 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.","Doderlein, L. (1885) Seeigel von Japan und den Liu-Kiu Inseln. Archiv fur Naturgeschichte, 51, 73 - 112. https: // doi. org / 10.5962 / bhl. part. 1569","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. (1951) A Monograph of the Echinoidea V. 2. Spatangoida. 2. Amphisternata. II. Spatangidae, Loveniidae, Pericosmidae, Schizasteridae, Brissidae. C. A. Reitzel, Copenhagen, 593 pp.","Clark, H. L. (1915) The echinoderms of Ceylon other than holothurians. Spolia Zeylanica, 10, 83 - 102."]}

* Metalia robillardi (de Loriol, 1876) Fig. 66 A–C. Metalia robillardi: Clark & Rowe, 1971: 166; Schultz, 2010: 394, Fig. 742; Filander & Griffiths, 2014: 55. Pl. II, Fig. H. Material examined. SAMC-A28231. Identification. Large, oval test; anterior side steep, with deep notch. Apical disc anteriorly positioned, with four gonopores. Anterior ambulacra narrow, somewhat sunken, simple pore-pairs; anterior petals directed posteriorly, posterior petals more flexed. Oral side with short, narrow labrum, slightly covering peristome; large periproct at short posterior end. Subanal fasciole with five (to six) pores on either side. Preserved specimen white, with short spines. Global maximum size. Maximum test length 125 mm. Global distribution. East coast of South Africa, Australia, Mauritius, Réunion and Madagascar (Schultz 2010); at 2–50 m depth. Remarks. Records collected in this study show that this species inhabits deeper waters than the previously reported 14 m (Filander & Griffiths 2014). Distinguished from Brissopsis lyrifera capensis by anterior part of test distinctively raised; by anterior petals being directed downwards and by white color., 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 page 58, DOI: 10.11646/zootaxa.4296.1.1, http://zenodo.org/record/843325, {"references":["Loriol, P. de. (1876) Echinodermes. Memoires de la Societe de physique et d'histoire naturelle de Geneve, 24, 59 - 673.","Clark, A. M. & Rowe, F. W. E. (1971) Monograph of Shallow-water Indo-West Pacific Echinoderms. Trustees of the British Museum, London, pp. 238.","Schultz, H. (2010) Sea-Urchins, a Guide to Worldwide Shallow Water Species. 3 rd Edition. Scientific Publication, Hemdingen, 484 pp.","Filander, Z. & Griffiths, C. L. (2014) Additions to and revision of the South African echinoid fauna (Echinodermata: Echinoidea). African Natural History, 10, 47 - 56."]}