Isostichopus maculatus subsp. phoenius

Isostichopus maculatus phoenius (Clark, 1922) Figs 1K–Y, 3A–D, 4B, 5A, 9, 10B, 11B, 15–18; Tables 1–3 Stichopus badionotus var. phoenius Clark, 1922: 60. Stichopus badionotus var. phoenius – Clark 1933: 109. — Deichmann 1930: 82. Isostichopus badionotus – Gómez-Maduro & Hernández-Ávila 2011: 223, figs k–l (at least one specimen). — Borrero-Pérez et al. 2012: 175, figs a, d. Stichopus sp. – Agudelo & Rodríguez 2015: 51–52, figs 1–3. — Vergara & Rodríguez 2015: 1022, fig. 1b. Isostichopus sp. – Invemar 2015: 115. — Vergara & Rodríguez 2015: 1022, fig. 1b; 2016: 130–137. — Martínez et al. 2016: 15, fig. 1, 22, fig. 6. — Vergara et al. 2018: 39–41, figs 4–6. — Acosta et al. 2020: 1–14, fig. 1; 2021: 1–17. Isostichopus sp. nov. – Agudelo-Martínez & Rodríguez-Forero 2017: 73. Isostichopus sp. aff. badionotus – Agudelo-Martínez & Rodríguez-Forero 2017: 6, figs 3–4. — Arias-Hernández et al. 2017: 278–289. — Fontalvo-Martínez & Rodríguez 2017: 95. — Medina-Lambrano et al. 2017: 175. Isostichopus isabellae n. sp. – Vergara et al. 2018: 40, fig. 4. Not an available name. Isostichopus maculatus phoenius – Borrero-Pérez et al. 2022: 180, 186. Isostichopus sp. ‘phoenius’ – Purcell et al. 2023: 148–149. Original name Stichopus badionotus var. phoenius Clark, 1922. Current status Isostichopus maculatus phoenius (Clark, 1922). Name-bearing type Holotype MCZ HOL-1182. Type locality Buccoo Reef, Tobago. Diagnosis Small dark spots surrounded by a clear halo and a dark thin line on white blotches on the background; semi-translucent and rugose body wall (Figs 1K–Y, 17–18); mtDNA divergence from other species of the genus> 7.7% in COI-Fr1 (barcoding region),>10.1% in COI-Fr2 and>7.9% in 16S; between subspecies 1.4%, 1,9% and 0.4%, respectively (Table 2). Material examined Holotype TRINIDAD AND TOBAGO • spec. uniform color pattern (L = 135 mm); Tobago, Buccoo Reef; 11.179396° N, 60.811479° W; 5 Apr. 1916; J. Mills leg.; living under or among rocks; MCZ HOL-1182. Other material GULF OF MEXICO – USA • 1 spec. (contracted); Texas, Stetson Bank; 28.17° N, 94.28° W; 28 Jun. 2005; depth 22 m; M. Wicksten leg.; USNM 1080469 • 1 spec. (L = 120 mm); Texas, Stetson Bank; 24 Oct. 1971; J. Teerlings leg.; USNM 1096856 • 1 spec. (L = 135 mm); Texas, North Padre Island; 26.82° N, 97.32° W; T. Shirley leg.; USNM E17227 • 2 specs (L = 95–195 mm); Florida, Tortugas, Bird Key; 24.6185° N, 82.8854° W; Jun. 1917; H. Lyman Clark, Carnegie Expedition exped.; MCZ HOL-1213. – Mexico • 2 specs; Veracruz, Bajos de Tuxpan, off Tuxpan; 21 Apr. 1994; J.D. Aghub leg.; ICML-UNAM 5.14.11 • 2 specs; Veracruz, Isla Sacrificios; 26 Jan. 1962; E. Caballero leg.; ICML-UNAM 5.14.28 • 2 specs; same locality as for preceding; 24 Jan. 1957; M.E. Caso leg.; ICML-UNAM 5.14.30 • 1 spec.; same locality as for preceding; 26 Jan. 1957; ICML-UNAM 5.14.34 • 1 spec.; Veracruz, Isla Sacrificios; 19.1749361° N, 96.09303056° W; 14 May 1997; depth 3 m; A. Medina leg.; rocky bottom; ICML-UNAM 18302 • 1 spec.; Veracruz, Isla Lobos; 21.4541667° N, 97.22500001° W; 14 Jun. 1976; depth 3 m; Acropora reef; ICML-UNAM 11459 • 1 spec.; Veracruz, Isla de Enmedio, Anton Lizardo; 19.1144444° N, 95.93583333° W; 5 Apr. 2005; depth 1–2 m; F. Solís-Marín leg.; on sandy bottom; ICML-UNAM 18303. CARIBBEAN SEA – Belize • 2 specs (L = 120–180 mm); Coco-Plum Key, edge of mangrove swamp; 16.877841° N, 88.120048° W; 2 May 1972; R. Larson leg.; USNM E18639. – Panama • 1 spec. (L = 130 mm); Bocas del Toro, Cayo Adriana; 9.2405278° N, 82.1734167° W; 30 May 2013; depth 5 m; G. Borrero-Pérez and A. Castillo leg.; mixed bottom of sand, sponges and corals, IpBT5; USNM 1659468 • 1 spec. (L = 135 mm); same data as for preceding; IpBT7; MBMLP-IpBT7 • 1 spec. (L = 170 mm); same data as for preceding; IpBT21; USNM 1659473 • 1 spec. (L = 140 mm); same data as for preceding; IpBT22; USNM 1659474 • 1 spec. (L = 138 mm); same data as for preceding; IpBT23; USNM 1659475 • 1 spec. (L = 170 mm); same data as for preceding; IpBT25; USNM 1659476 • 1 spec. (L = 160 mm); same data as for preceding; IpBT43; USNM 1659477 • 1 spec. (L = 150 mm); same data as for preceding; IpBT44; USNM 1659478 • 1 spec. (L = 185 mm); same data as for preceding; IpBT53; USNM 1659479 • 1 spec. (L = 150 mm); same data as for preceding except 31 May 2013; on mixed bottom of sand, sponges and corals, completely hidden in a crevice between sponges, IpBT26; TpBT0026 • 1 spec. (L = 160 mm); same data as for preceding; MBMLP IpBT27 • 4 specs (L = 135–180 mm); same data as for preceding; Ip8, 24, 28, 29 BT; TpBT0008, TpBT0024, TpBT0028, TpBT0029 • 1 spec. (L = 130 mm); Bocas del Toro, STRI research station; 9.3488611° N, 82.2585° W; 4 Jun. 2013; depth 5 m; G. Borrero-Pérez leg.; mixed bottom of sand, corals and isolated seagrass, exposed, between coral colonies, IpBT131; USNM 1659469 • 1 spec. (L = 130 mm); Bocas del Toro, Popa Island Cay; 9.2495° N, 82.1815279° W; 6 Jun. 2013; depth 7 m; G. Borrero-Pérez and A. Castillo leg.; mixed bottom of sand, corals and sponges at the reef fringe, completely hidden, under a loose coral head and among rubble coral, IpBT190; USNM 1659470 • 1 spec. (L = 140 mm); Bocas del Toro, near Popa Island Cay; 9.2541944° N, 82.1829444° W; 6 Jun. 2013; depth 2 m; G. Borrero-Pérez and A. Castillo leg.; mixed bottom dominated by Porites porites, with isolated seagrasses, octocorals and sponges, exposed, IpBT191; USNM 1659471 • 1 spec. (L = 125 mm); same data as for preceding; depth 3 m; IpBT192; USNM 1659472 • 1 spec.; Galeta; 9.4018444° N, 79.86053056° W; 15 Dec. 2015; depth 2 m; G. Borrero-Pérez leg.; mixed bottom of rocky, corals and isolated seagrass, exposed, IpGa8; Tiss-IpGa8 • 1 spec.; same data as for preceding; IpGa11; Tiss-IpGa11 • 1 spec.; same data as preceding, IpGa209; Tiss-IpGa209 • 1 spec. (L = 100 mm); Galeta; 9.4018444° N, 79.86053056° W; 17 Sep. 2016; depth 3 m; G. Borrero-Pérez leg.; mixed bottom of corals and rocks, exposed but camouflaged between Porites porites, IpGa269; Tiss-IpGa269 • 1 spec. (L = 158 mm); same data as for preceding; depth 1 m; A. Calderón leg.; completely hidden inside rocks, IpGa223; USNM 1659480 • 1 spec. (L = 90 mm); same data as for preceding; IpGa224; USNM 1659481. – Colombia • 1 spec. (L = 120 mm); Magdalena, Banco de las Ánimas; 11.0397778° N, 74.39919445° W; 1 Dec. 2013; depth 15 m; G. Borrero-Pérez and N. Ardila leg.; rocky bottom, anterior part exposed, IpBA313; INV EQU4144 • 1 spec. (L = 130 mm); same data as for preceding; IpBA314; INV EQU4146 • 1 spec. (L = 140 mm); same data as for preceding; rocky bottom, partially exposed within crevices between rocks, concealed by spines of Diadema antillarum, IpBA315; INV EQU4147 • 1 spec.; same data as for preceding; depth 4.5 m; IpBA319; INV TEJ1336 • 1 spec. (L = 25 mm); Magdalena, Bahía del Rodadero; 11.2070556° N, 74.2303916° W; 23 Jul. 2017; depth 1 m; G. Borrero leg.; hidden under a rock in a mixed bottom of sediment and small rocks, IpRod; INV EQU4565 • 1 spec. (L = 125 mm); La Guajira, El Pájaro-Tawaya; 11.7308333° N, 72.70436111° W; 9 Sep. 2013; depth 4.5 m; E. Ortíz and J. López leg., IpGV82; INV TEJ1257 • 1 spec. (L = 100 mm); same data as for preceding; IpGV83; INV TEJ1258 • 1 spec. (L = 125 mm); same data as for preceding; IpGV84; INV TEJ1259 • 1 spec. (L = 160 mm); same data as for preceding; IpGV86; INV TEJ1261 • 1 spec. (L = 170 mm); same data as for preceding; IpGV90; INV TEJ1265 • 1 spec. (L = 120 mm); same data as for preceding; IpGV108; INV TEJ1269 • 1 spec. (L = 180 mm); same data as for preceding; IpGV117; INV TEJ1272 • 1 spec.; La Guajira, Cabo de la Vela; 12.2086422° N, 72.1674894° W; 28 Oct. 2011; depth 2 m; G. Borrero-Pérez and C. Díaz leg., IpGV1; INV-TEJ1113 • 1 spec.; same data as for preceding; IpGV3; INV-TEJ1115. – Trinidad and Tobago • 4 specs (L = 90–115 mm); Tobago, Buccoo Bay; 11.177882° N, 60.811479° W; Apr. 1916; J. Mills leg.; MCZ HOL-1181. – Jamaica • 1 spec. (L = 120 mm); Kingston Harbor, Port Royal, Drunken Man’s Cay; 17.905085° N, 76.845067° W; 3 Sep. 1959; H. Lyman Clark leg.; MCZ HOL-4282. – Antigua and Barbuda • 2 specs (L = 90–120 mm); Antigua Island, English Harbor; 1 Jul. 1918; C. Nutting leg.; USNM 1283367 • 1 spec. (L = 170 mm); Antigua Island, Falmouth Bay; 17° N, 61.7667° W; 1918; J. Henderson leg.; USNM E41458. Description EXTERNAL APPEARANCE. Medium sized species, preserved and live ex situ specimens examined range from 25 to 260 mm in length (n = 68, holotype: 130 mm long and 40 mm wide). Body loaf-like, length/ width ratio 3.7± 0.8 (n = 38, 3.2–5.5, holotype 3.2). Rounded posteriorly and anteriorly (Fig. 15A), quadrangular in cross section (Figs 17–18). Live specimens also tapering slightly posteriorly and widened anteriorly when the animal is relaxed (Fig. 17E, O), or when it is coming out of its crevice, especially when only the anterior part of the body is exposed (Fig. 18O). Body wall firm and thick (0.5 to 4 mm thick in the holotype). Anus supra-terminal, circular and surrounded by large papillae. Mouth directed ventrally in live and preserved animals, encircled by a collar of large papillae (2–4 mm long in the holotype). Large peltate tentacles 20, about 7–9 mm long, 6 mm shield width, with indentations 0.5–3 mm deep. Dorsal papillae medium to large, conical, and spiky (although some of them somewhat rounded) ending in a sharply pointed tip (5 to 10 mm high; 5 to 8 mm wide); some smaller papillae rounded and scattered irregularly; most of the largest dorsal papillae arranged in two dorsal irregular rows along each edge, defining the quadrangular shape. Holotype covered with medium size conical papillae (up to 4 mm high and 5 mm wide at the base), most of them flattened and not very conspicuous. Lateral papillae large and spiky, sharply defining the dorsal and ventral surface (Figs 17–18); in the holotype they are larger than dorsal ones (up to 5 mm high and 5 mm wide at the base) (Fig. 15A). Ventral surface covered with cylindrical, large pedicels (2–4 mm long and 0.5 mm wide in the h