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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"]}

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).