Ophioderma sp. Granja-Fernández et al . 2014: 135–137 Ophioderma hendleri sp. nov. (Echinodermata: Ophiuroidea: Ophiodermatidae) and its congeners from the Eastern Pacific Granja-Fernández, Rebeca Pineda-Enríquez, Tania Solís-Marín, Francisco Alonso Laguarda-Figueras, Alfredo European Journal of Taxonomy 2020 2020-12-14 729 11 41 Granja-Fernández & Pineda-Enríquez & Solís-Marín & Laguarda-Figueras, 2020 Granja-Fernández & Pineda-Enríquez & Solís-Marín & Laguarda-Figueras 2020 [615,875,1298,1325] Ophiuroidea Ophiodermatidae Ophioderma Animalia Ophiurida 3 14 Echinodermata species hendleri sp. nov.   urn:lsid:zoobank.org:act: 3455BEAC-DF5D-460A-89A8-A70B9A080BB4  Figs 1–4, 5A–D; Table 1     Ophiodermasp.–  Granja-Fernández et al. 2014: 135–137, fig. 7A–F; 2015a: 41–42; 2017: 172–175.    Diagnosis Radial shields covered by granulation. Adoral shields naked, as long as broad. Up to 11 arm spines, dorsalmost the shortest and blunt, ventralmost the largest and robust. In vivo coloration, disc dark brown and mottled with small white spots, ventral margin beige and brown; arms dark brown with transverse white bands.    Etymology This species is named after Dr. Gordon Hendler, an eminent ophiuroid taxonomist, curator of echinoderms at the Natural History Museum of Los Angeles County.    Material examined   Holotype   MEXICO• 1 spec.; Chipehua, Oaxaca, 16°1´49.90˝ N, 95°21´33.70˝ W;  21 m;  5 Nov. 2011; R. Granjaleg.; under rocks; ICML-UNAM 18315.   Paratypes   COSTA RICA• 6 specs; Coco Island;  19–28 Feb. 1994; Gordon Hendlerleg.; MZUCR-ECH 321.   MEXICO• 2 specs; Zacatoso, Ixtapa, Guerrero; 17°39´15.00˝ N, 101°37´19.00˝ W;  9.1 m;  1 Jun. 2012; R. Granjaleg.; under rocks; ICML-UNAM 10582•  1 spec.; Chahué, Oaxaca; 15°45´02.90˝ N, 96°07´23.43˝ W;  6.5 m;  14 Jan. 2017; R. Granjaleg.; under rocks, 27°C; ICML-UNAM 18316•  2 specs; Tijera, Oaxaca; 15°41´15.12˝ N, 96°26´31.47˝ W;  9 m;  29 Jan. 2016; D. Rangeland R. Granjaleg.; under rocks, 27°C; ICML-UNAM 18317•  1 spec.; Frente a Llano Grande, Oaxaca; 16°14´30.26˝ N, 98°27´04.64˝ W;  20 m;  1 Feb. 2016; Rebeca Granjaleg.; sediment under rocks, 27°C; ICML-UNAM 18318•  1 spec.; La Jeringa, Cacaluta Island, Oaxaca; 15°43´10.92˝ N, 96°09´39.91˝ W;  9–14 m;  26 Jan. 2016; D.Rangeland R. Granjaleg.; sediment under rocks; ICML-UNAM 18319•  2 specs; Punta Lobos, Espíritu Santo Island, Gulf of California; 24°28´29.06˝ N, 110°17´20.65˝ W;  15–17 m;  28 Oct. 2016; R. Granjaleg.; under rocks, 28°C; ICML-UNAM 18320•  3 specs; San Diego Norte, Gulf of California; 25°12´15.48˝ N, 110°41´41.64˝ W;  13 m;  29 Oct. 2016; R. Granjaleg.; under rocks, 29°C; ICML- UNAM 18321•  1 spec.; San Mateo Norte, Gulf of California; 25°22´43.54˝ N, 110°59´34.26˝ W;  12 m;  31 Oct. 2016; R. Granjaleg.; under rocks, 29°C; ICML-UNAM 18322•  10 specs; Isla San Diego, Gulf of California; 25°12´12.78˝ N, 110°41´40.14˝ W;  10–11 m;  31 Aug. 2008; F. Solísand Solís-Wolfowitzleg.; under rocks; ICML-UNAM 18323•  5 specs; El Peruano, Guaymas, Sonora, Gulf of California; 27°54´30.06˝ N, 110°58´10.68˝ W;  12 m;  12 Aug. 2011; C. Sánchezleg.; under rocks, ICML-UNAM 18324•  1 spec.; La Lobera, Bahía de La Paz, Gulf of California; 24°36´56.39˝ N, 110°24´00.57˝ W;  7 m;  2 Mar. 2010; F. Solísleg.; under rocks, ICML-UNAM 18325•  1 spec.; Isla Larga(estación Este), Marietas Islands; 20°41´08.00˝ N, 105°34´00.00˝ W;  9 m;  22 Mar. 1996; F. Solísand C. Nepoteleg.; under rocks; ICML-UNAM 18326•  1 spec.; Los Islotes, Bahía de La Paz, Gulf of California; 24°35´00.00˝ N, 110°25´00.00˝ W;  18.3 m;  28 Apr. 1997; H. Cortésleg.; sediment under rocks; ICML-UNAM 18327.   PANAMA• 2 specs; Panama;  24 Jun.–23 Jul. 1872; Hassler expedition leg.; MCZ OPH-116.  Other material See: Supplementary material.     Holotypedescription DD = 14.8 mm; 5 arms, AL = 69.1 mm. Disc flat, nearly circular. Dorsal disc densely covered by rounded, small granules, slightly separated from each other; granule density 113 mm-2. Granules at periphery of disc and on base of arms slightly larger than on central part of disc. Radial shields covered by granules ( Fig. 1A). Interradii covered with granulation, similar to periphery and base of arms ( Fig. 1B). Four genital slits in each interradius ( Fig. 1B); proximal genital slits oval, in contact with distal part of oral shield and with 1 stLAP; distal genital slits rounded, elongated, placed between 5 thand 6 tharm segment and close to margin of disc, surrounded by granulation and numerous, elongated and imbricated scales next to side of lateral arm plates ( Fig. 1B). Oral shields slightly broader than long, rounded triangular, with convex obtuse proximal angle, straight to convex distal edge, obtuse lateral angles. Madreporite with central, circular, shallow depression, located in distal part of oral shield. Adoral shields naked, as long as broad, triangular and completely separated from each other. Jaws bear 8–9 oral papillae on each side: LyOs small, 2 × as long as broad, angled upwards; AdShSp largest, rounded; 2°AdShSp elongated and narrow; LOPas 4–5 conical and pointed, slightly separated; IPa reduced, elongated and pointed; TPa pair at apex of jaw, elongated, robust and pointed, slightly separated. vT similar in shape and slightly larger than TPa. Four teeth: ventralmost broader than long, quadrangular but with rounded borders; dorsalmost pointed, slender and smallest. One OPRSp, large and pointed at each side, visible within buccal cavity. Granules covering oral plates and below adoral shields larger than those on dorsal disc and interradii ( Fig. 1C–D).   Fig. 1.  Ophioderma hendleri sp. nov., holotype (ICML-UNAM 18315, DD = 14.8 mm). A. Dorsal disc. B. Ventral disc. C. Mouth. D. Structures of a jaw (according to Hendler 2018). E. Basal dorsal arm. F. Dorsal arm. G. Ventral arm. H. Lateral arm plates and arm spines. I. Dorsal view. See Material and methods for abbreviations.   Fig. 2.SEM images of skeletal elements of  Ophioderma hendleri sp. nov., paratype (ICML-UNAM 18323, DD = 15 mm). A–B. Radial shield (external view), deepest pores marked with arrows. C. Center of the distal portion of the radial shield, open meshed pores. D. Dorsal arm plate, with one spur. E. Ventral arm plate, with one spur. F. Lateral arm plate, external face, with two spurs and one condyle. G. Lateral arm plate, lateral face, with spine articulations. H. Lateral arm plate, internal face. I. Outer surface of the LAP (F) with finely meshed and polygonal knobs. J. Spine articulations with a weak sigmoidal fold and tilted lobes. K. Arm vertebra, distal face. L. Arm vertebra, proximal face. M. Tentacle scale. N. Arm spine. O. Dental plate fragments. P. Oral plate, abradial face. Q. Oral plate, adradial face.   Fig. 3.  Ophioderma hendleri sp. nov., paratype (ICML-UNAM 10582, DD = 4.8 mm). A. Dorsal view. B. Basal dorsal arm. C. Dorsal arm. D. Ventral arm. E. Dorsal disc. F. Ventral disc. G. Interradii. H. Mouth.   Fig. 4.  Ophioderma hendleri sp. nov., paratype (ICML-UNAM 18319, DD = 2.2 mm). A. Dorsal view. B. Basal dorsal arm. C. Dorsal arm. D. Ventral arm. E. Dorsal disc. F. Ventral disc. G. Interradii. H. Mouth.   Fig. 5. A–D.  Ophioderma hendleri sp. nov. A–B. In vivo coloration in natural habitat. C–D. In vivo coloration of collected specimens. – E–J.  Ophioderma panamense Lütken, 1859, holotype (NHMD- 107679, DD = 21.9 mm). E. Dorsal disc. F. Ventral disc. G. Mouth. H. Dorsal arm. I. Ventral arm. J. Dorsal view.   Table 1.Comparison of morphological characters between some species of  Ophiodermafrom the Eastern Pacific.       O. hendleri sp. nov.   O. panamense   O. pentacanthum   O. variegatum   Radial shields covered by granules naked covered by granules covered by granules   Base of the arms covered by granules and scales covered by granules and scales covered by granules and scales covered by granules   Arm-spines up to 11; short and more or less robust up to 12; short and robust up to 6; large and thin, the ventral-most being much larger and robust up to 9; very large and thin   Tentacle scales adradial slightly larger than abradial adradial slightly larger than abradial adradial much larger than abradial adradial slightly larger than abradial   Oral shields broader than long, rounded triangular, with rounded obtuse proximal angle, straight distal edge, obtuse lateral angles broader than long, rounded triangular, with rounded obtuse proximal angle, straight distal edge, obtuse lateral angles broader than long, rounded triangular, with rounded obtuse proximal angle, straight distal edge, obtuse lateral angles longer than broad, oval, proximal margin rounded obtuse angled, distal edge convex, lateral margins straight   Adoral shields naked; equally long as broad covered by granules naked; longer than broad naked; longer than broad   Genital slits short and rounded; distally surrounded by granules and scales short and rounded; distally surrounded by granules and scales large and slender; distally surrounded by granules and scales short and rounded; distally surrounded only by scales   Coloration in vivo disc dark brown mottled with small and white spots, ventral margin beige; arms with transverse white bands disc green, brown or grey mottled with red, orange or yellow, sometimes a central white spot; arms with transverse bands lighter and darker disc orange with dark brown blotches; arms with transverse brown bands disc olive-green with dark pink or red; arms with transverse olivegreen and dark pinkred bands   Depth intertidal – 37 m intertidal – 20 m 102 – 183 m intertidal – 110 m Dorsal base of arms covered by granules and with approximately 11 oval scales, located laterally at first 3 dorsal arm plates ( Fig. 1E). Dorsal arm plates 3 × broader than long, rectangular, overlapping; proximal edge straight, distal edge slightly concave with rounded lateral edges ( Fig. 1F). Distalmost dorsal arm plates smaller, longer than broad, triangular. First ventral arm plate small, broader than long, with rounded edges; in contact with adoral shields. Second ventral arm plate quadrangular, as broad as long ( Fig. 1C). Next ventral arm plates slightly longer than broad, quadrangular; proximal margin truncate and distal edge convex ( Fig. 1G). Distalmost ventral arm plates wider distally, with a tapering proximal angle. Paired rounded depressions between most proximal ventral arm plates ( Fig. 1B). LAPs conspicuous, semicircular, broader than long ( Fig. 1H). LAP with up to 11 arm spines. First and 2 ndLAP with 3 arm spines; 3 rdwith 4 arm spines; 4 thwith 5; 5 thwith 6; 6 thwith 7; 7 thwith 8; 8 thwith 9; 9 th–30 tharm plates with 10–11 arm spines; 35 thwith 9 spines; 40 th–50 thwith 8; 60 thwith 7; 70 thwith 6; 75 thwith 5; 80 thwith 4;>80 thwith 3 arm spines. Arm spines with blunt tip; length approximately ½ of LAP. Dorsalmost arm spine shortest, blunt; ventralmost arm spine longest and more robust, as long as 1 arm segment, almost in contact with tentacle scale of succeeding joint ( Fig. 1H). Two tentacle scales; adradial tentacle scale ovoid, approximately ½× length of ventral arm plate; abradial tentacle scale slightly shorter, subtriangular ( Fig. 1G). General coloration dark brown (dry specimen) ( Fig. 1I). Dorsal side: disc dark brown and mottled with small white spots; middle of margin on each interradius of disc bearing white spot ( Fig. 1A); arms dark brown with small black spots only observed microscopically and with every 5–6 arm plates with transverse white bands, located between distal part of ventral plate and proximal part of next arm plate ( Fig. 1F, I). Ventral side: interradii, center and proximal part beige, margin dark brown with some beige marks ( Fig. 1B). Jaw beige but oral shields can display brown color ( Fig. 1C). Ventral arm plates beige, but that on tip of arms dark brown with some beige transversal lines ( Fig. 1G). LAPs dark brown with some beige marks; arm spines dark brown and generally beige color on base and tip ( Fig. 1H).  Disarticulated ossicles Specimen analyzed: 1 spec., paratypeICML-UNAM 18323 (DD = 15 mm, AL = 80.8 mm). Radial shield (external view) irregularly triangular, with convex proximal margin, incised abradial edge with projection, convex to straight distal edge and irregular adradial edge, series of 8 pores on median to proximal margin (which are covered/overlapped by disc scales and granules in intact animal) ( Fig. 2A– B), most proximal ones deepest. Centerof distal portion of radial shield covered in intact animal. Outer surface of stereom is open mesh of pores and small knobs ( Fig. 2C). Dorsal arm plate slightly arched and somewhat rectangular, 3× as wide as long; proximal margin convex and distal margin straight, 1 spur on proximal portion of external surface ( Fig. 2D). Ventral arm plate as long as wide, quadrangular with proximal side truncated and pointed spur, lateral sides concave forming border of tentacle pore ( Fig. 2E). LAP D-shaped, 2× as high as wide, with 8 spine articulations sunken in notches of distal edge ( Fig. 2F–G). Ventral portion of LAP projecting ventro-proximalwards and ventro-distal tip projecting ventralwards ( Fig 2F). Ventro-proximal margin with condyle ( Fig. 2F). Proximal edge of LAP with 2 prominent and elongated spurs, protruding and modifying central-proximal edge ( Fig. 2F), between spurs and across remaining proximal margin discernible band of different stereom structure ( Fig. 2F). Outer surface finely meshed with relatively large polygonal knobs ( Fig. 2I). Inner surface of LAP with continuous ridge on proximal edge, and at ventro-distal margin 2 spurs matching those on external surface ( Fig. 2H), and 3 pores just below center. Spine articulations ventralwards increasing in size ( Fig. 2G). Lobes with weak sigmoidal fold, tilted, curved, slightly joined on proximal margin by 3 or 4 knobs; ventral lobe smaller than dorsal lobe ( Fig. 2J). Proximal vertebrae almost round, as wide as long, with large aboral muscle flange and smaller oral muscle flange ( Fig. 2L). Distal face of vertebrae with large muscle flanges, with typical zygospondylous articulation ( Fig. 2K). Tentacle scale longer than wide, with scale-like surface ( Fig. 2M). Spines with scale-like surface and blunt tip ( Fig. 2N). Dental plate consists of several pieces, 1 stpiece bears 2 TPas on small round socket and 1 wide tooth socket, while rest of pieces with single tooth socket per piece ( 4 intotal), none of sockets penetrate plate ( Fig. 2O). Oral plates as long as high, abradial face with muscle fossa highly triangular, almost covering whole surface ( Fig. 2P), adradial face in middle part with foot basin ( Fig. 2Q).    Paratypevariations Large specimens showed some differences in morphology compared to small ones ( Figs 1, 3–4). Large individuals have granules and scales on the base of the arms ( Fig. 1E), but in the smallest specimens (DD = ~ 2–4 mm) the scales are covered by granules ( Fig. 4B). The scales begin to appear as the animal grows; therefore, in specimens with DD = 4–6 mmthere are up to 3 elongated scales to either side of the base of the arms ( Fig. 3B), and with a DD = 7 mmthere are up to six scales; the number of scales in larger specimens reaches 11 ( Fig. 1E). Additionally, large individuals have numerous scales located on the distalmost part of the interradii ( Fig. 1B) but in the smallest specimens (DD = 3–6 mm) these scales are not evident, and instead, only granules are present ( Figs 3F–G, 4F–G). In specimens larger than DD = 7 mm, a row of elongated scales in the distal genital slits is evident. Finally, in the smallest specimens the adoral shields are completely covered by granules, which disappear as the animal grows, and individuals with DD = 5 mmtend to have totally or partially covered adoral shields. Some specimens can have beige specks on the LAPs, the arm spines and the tentacle scales. Moreover, some specimens from the Gulf of California ( Mexico) have a large white spot in the center of the dorsal disc. It is important to note that in vivo coloration ( Fig. 5A–D), generally conserved in all preserved specimens ( Figs 1, 3–4), can be used as a distinctive character in the field.    Habitat and distribution Widely distributed in the Eastern Tropical Pacific.  Ophioderma hendleri sp. nov.has been collected in the Mexican Pacific from the Gulf of California (in the States of Sonora and Baja California Sur), Jalisco, Michoacán, Guerrero and Oaxaca as well as in Revillagigedo, Marías and Marietas Islands. In Costa Ricait has been collected from Murciélago, Caño, and Cocos Islands; in Panamafrom Pearl Islands; and finally, in Colombiafrom the localities La Parguera and La Roñosa Rock (see: Supplementary material). The northernmost locality of its distribution is off San Francisquito Bay (Gulf of California, Mexico; 28° N) and the southernmost point is La Roñosa Rock, Colombia( 3° N). Due to its distribution,  O. hendleri sp. nov.can be considered as a species with a Panamic affinity, suggesting that, similar to other ophiuroids (e.g.,  Ophiocoma aethiops Lütken, 1859and  Ophiocomella alexandri( Lyman, 1860)), its distribution may extend to northern Peru, where a transitional zone between the Panamic and the Peruvian Provinces is located ( Granja-Fernández & Hooker 2020), but more sampling effort is needed to confirm its presence or absence in this area. The species inhabits depths between 0.40– 37 m.  Ophioderma hendleri sp. nov.is mostly collected under rocks, burrowed in the sediment and it is common to find it associated with coral and rocky reefs ( Granja-Fernández et al. 2014).    Remarks   Ophioderma hendleri sp. nov.is one the most widely distributed ophiodermatids in the Eastern Tropical Pacific and it is of interest to note that although it has been collected since 1936, it has been misidentified in all the reviewed collections as  O. variegatumor  O. panamense(see Discussion). In previous works, Granja-Fernández et al. (2014, 2015a, 2017) mentioned the presence of  Ophiodermasp. 1, which actually corresponds to  O. hendleri sp. nov.; therefore, these records are considered to be this species. In our revision of the specimens, we did not observe any differences in the morphological characters from different geographical areas, suggesting that intraspecific variation in  O. hendleri sp. nov.is not related to its distribution. On the other hand, it is very notable that this species in most cases presents the same color pattern (dorsal side dark brown, ventral side beige and arms with transversal lines; Figs 1, 3–5), with some exceptions (see Paratypevariations). This is in contrast to other  Ophiodermaspecies such as  O. panamenseor  O. variegatum, which can display a great variety of colors (green, yellow, pink, etc.; Nielsen 1932; Ziesenhenne 1955). This species is commonly found at the same localities as  O. panamenseand other ophiuroids such as  Ophiactis simplex(Le Conte, 1851),  Ophiactis savignyi(Müller & Troshel, 1842),  Ophiocoma aethiops,  Ophiocomella alexandri,  Ophionereis annulata(Le Conte, 1851),  Ophiothela mirabilis Verrill, 1867and  Ophiothrix( Ophiothrix) spiculataLe Conte, 1851. It is one of the most conspicuous  Ophiodermaspecies of the Eastern Tropical Pacific and it is always hiding under small and medium-sized rocks and in sediment of median size of rocky and coral reefs ( Fig. 5A–B). Its abundance has not been quantified, but using scuba diving, in an hour of sampling effort of all the species of ophiuroids at one locality, a total of 5– 6 specimenswere found, a smaller number compared to  O. panamense(8– 12 specimens). It tends to be gregarious, and it is very common to find at least two specimens occupying the same space. Field notes denote that  O. hendleri sp. nov.has been found in waters with a bottom temperature of 27–29°C. 2995182307 under rocks 2011-11-05 R R. Granja Mexico 21 16.030527 Chipehua 1 -95.35936 3 14 ICML-UNAM 18315 1 Oaxaca holotype 2995182309 [189,1337,300,327] 1994-02-19 1994-02-28 1994-02-19 MZUCR-ECH Gordon Hendler Costa Rica Coco Island 4 15 MZUCR-ECH 321 6 paratype 2995182315 under rocks 2012-06-01 R R. Granja Mexico 9 17.654167 Zacatoso 1 -101.62195 Zacatoso 4 15 ICML-UNAM 10582 2 Guerrero 2995182304 under rocks 2017-01-14 R R. Granja Mexico 7 15.750806 Chahue 1 -96.12318 4 15 ICML-UNAM 18316 1 Oaxaca 2995182316 under rocks 2016-01-29 R D. Rangel & R. Granja Mexico 9 15.687533 Tijera 1 -96.44208 4 15 ICML-UNAM 18317 2 Oaxaca 2995182311 sediment under rocks 2016-02-01 ICML-UNAM Rebeca Granja Mexico 20 16.24174 Frente a Llano Grande 1 -98.45129 Frente 4 15 ICML-UNAM 18318 1 Oaxaca 2995182317 sediment under rocks 2016-01-26 R D. Rangel & R. Granja Mexico 12 15.7197 La Jeringa 1 -96.16109 La Jeringa 4 15 ICML-UNAM 18319 1 Oaxaca 2995182306 under rocks 2016-10-28 R R. Granja Mexico Punta Lobos 16 24.47474 Espiritu Santo Island 1 -110.28907 Espiritu Santo Island 4 15 ICML-UNAM 18320 2 2995182302 under rocks 2016-10-29 R, ICML- R. Granja Mexico 13 25.2043 Gulf of California 1 -110.6949 San Diego Norte 4 15 ICML- UNAM 18321 3 2995182314 under rocks 2016-10-31 R R. Granja Mexico 12 25.378761 Gulf of California 1 -110.99284 San Mateo Norte 4 15 ICML-UNAM 18322 1 2995182310 under rocks 2008-08-31 ICML-UNAM F. Solis & Solis-Wolfowitz Mexico 11 25.20355 Gulf of California 1 -110.69448 Isla San Diego 4 15 ICML-UNAM 18323 10 2995182318 under rocks 2011-08-12 ICML-UNAM C. Sanchez Mexico El Peruano 12 27.90835 Gulf of California 1 -110.969635 Guaymas 4 15 ICML-UNAM 18324 5 Sonora 2995182301 under rocks 2010-03-02 ICML-UNAM F. Solis Mexico La Lobera 7 24.615664 La Lobera 1 -110.40016 Bahia de La Paz 4 15 ICML-UNAM 18325 1 2995182320 under rocks 1996-03-22 ICML-UNAM F. Solis & C. Nepote Mexico Isla Larga 9 20.685555 Isla Larga 1 -105.566666 Este 4 15 ICML-UNAM 18326 1 2995182324 sediment under rocks 1997-04-28 ICML-UNAM H. Cortes Mexico Los Islotes 18 24.583334 Los Islotes 1 -110.416664 Bahia de La Paz 4 15 ICML-UNAM 18327 1 2995182303 [189,1276,1146,1173] 1872-06-24 1872-07-23 1872-06-24 MCZ Panama Panama 4 15 MCZ OPH-116 2 Panama