Dolabrifera nicaraguana, PILSBRY, 1896

Valdés, Ángel, Breslau, Eric, Padula, Vinicius, Schrödl, Michael, Camacho, Yolanda, Malaquias, Manuel António E., Alexander, Jennifer, Bottomley, Morgan, Vital, Xochitl G., Hooker, Yuri & Gosliner, Terrence M., 2018, Molecular and morphological systematics of Dolabrifera Gray, 1847 (Mollusca: Gastropoda: Heterobranchia: Aplysiomorpha), Zoological Journal of the Linnean Society 184, pp. 31-65 : 51-52

DOLABRIFERA NICARAGUANA PILSBRY, 1896

( FIGS 3I–J, 14–16)

Dolabrifera nicaraguana Pilsbry, 1896: 124 , pl. 63, figs 12–16. Type locality: San Juan del Sur , Nicaragua (Pacific coast).

Type material

Three syntypes at ANSP 67517 (dry) and ANSP A7048 (wet).

Material examined

Puerto Vallarta , Mexico, intertidal, 2 May 2002, one specimen 15 mm preserved length ( CPIC 00547 ) . El Salvador, no locality data or date, two shells, leg. A.J. Ferreira ( LACM 10383–10384 ) . Costa Rica, no locality data or date, two shells, leg. A.J. Ferreira ( LACM 10381–10382 ) . North side of San Juan del Sur Bay, Nicaragua (11°15′N, 85°52′W), 25 Jan 1974, six specimens 10–13 mm preserved length, one shell removed ( CASIZ 001449 ) . Bahía Salinas , Guanacaste, Costa Rica (11°02′N, 83°42′W), intertidal, 10 February 1935, two specimens 30–35 mm preserved length ( LACM 1935 - 107.25 ) ; one specimen 35 mm preserved length ( LACM 1935 - 107.25 ) . Puerto Parker , Guanacaste, Costa Rica (10°56′N, 85°48′47″W), intertidal, 9 February 1935, one specimen 23 mm preserved length ( LACM 1935 - 115.17 ) . South of Punta Mala , Puerto Culebra, Guanacaste, Costa Rica (10°36′30″N, 85°42′15″W), intertidal, 24 February 1934, three specimens 25–35 mm preserved length ( LACM 1934 - 130.21 ) ; eight specimens 20–30 mm preserved length ( LACM 1934 - 130.21 ) . Bahía San Josecito , Provincia Puntarenas, Costa Rica, 14 January 2003, two specimens 24–28 mm preserved length ( CPIC 00551 ) . North side of Isla del Caño, Costa Rica (8°43′15″N, 83°53′07″W), intertidal, 18–19 March 1972, four specimens 10–15 mm preserved length, leg. J.H. McLean ( LACM 1972 - 68.45 ) . Bahía Honda , Panama (7°43′30″N, 81°32′40″W), intertidal, 28 March 1939, seven specimens 22–30 mm preserved length ( LACM 1939 – 52.1 ) . South of Punta Marzo , Bahía Octavia , Choco, Colombia (6°49.8′N, 77°49.6′W, intertidal, 27 January 1935, one specimen 21 mm preserved length ( LACM 1935 – 73.9 ). Bahía Cupica , Choco , Colombia (6°39.2′N, 77°30.7′W, intertidal, 26 January 1935, one specimen 22 mm preserved length ( LACM 1935 – 73.9 ). Puerto Utria , Choco , Colombia (5°59.2′N, 77°21.3′W, intertidal, 23 January 1935, one specimen 30 mm preserved length ( LACM 1935 – 52.25 ); 25 February 1938, three specimens 23–31 mm preserved length ( LACM 1938 - 126.18 ). Isla Gorgona , Cauca, Colombia (3°0.5′N, 78°11.8′W), intertidal, 12 February 1934, one specimen 24 mm preserved length ( LACM 1934 – 92.17 ). Cabo de San Francisco , Ecuador (0°39′30″N, 80°06′30″W), intertidal, two specimens 32–37 mm ( LACM 1938 - 116.11 ). Sombrero Chino , Isla Santiago , Islas Galápagos, Ecuador (0°22′20″S, 90°34′30″W), intertidal, 22 March 1971, six specimens 15–22 mm preserved length ( LACM 1971 - 64.6 ). Punta Alfaro , Isla Isabela , Islas Galápagos, Ecuador (0°25′20″S, 90°57′10″W), intertidal, 25 March 1971, three specimens 8–12 mm preserved length ( LACM 1971 - 70.10 ). Isla Pinzón , Islas Galápagos, Ecuador (0°35′50″S, 90°39′15″W), intertidal, 25–26 March 1971, five specimens 16–22 mm preserved length ( LACM 1971 - 72.16 ). Darwin Research Station , Academy Bay , Isla Santa Cruz , Islas Galápagos, Ecuador (0°45′06″S, 90°15′38″W), 1 m, March 1971, one specimen 23 mm preserved length ( LACM 1971 - 45.20 ); intertidal, March 1971, one specimen 21 mm preserved length ( LACM 1971 - 44.17 ). Flamingo Cove, near Post Office Bay, Isla Floreana, Islas Galápagos, Ecuador (1°14′S, 90°27′30″W), intertidal, 15 March 1971, two specimens 13–24 mm preserved length ( LACM 1971 - 52.13 ). Isla de la Plata, Ecuador (1°16′S, 81°05′10″W), intertidal, 22 January 1933, one specimen 45 mm preserved length ( LACM 1933 - 22.21 ). Salinas, Santa Elena, Ecuador (2°12′S, 81°58′W), 5–6 March 1970, one specimens 28 mm preserved length, leg. J.H. McLean ( LACM 1970 – 9.29 ). South Point of Santa Elena, Ecuador (2°12′23″S, 81°00’05″W), 8 February 1934, three specimens 40 mm preserved length ( LACM 1934 – 81.15 ). Balneario Punta Sal, Tumbes, Peru, intertidal, 9 August 2005, one specimen 16 mm preserved length, leg. K. Nakamura ( CPIC 00194 ).

Description

External morphology: Body up to 50 mm long, oval to elongate, flattened, broader posteriorly, narrowing gradually towards head. Dorsum covered with densely packed tubercles of different sizes, each with retractile, simple, short papilla on apex; larger tubercles surrounded by smaller ones ( Fig. 3I, J). Rhinophores enrolled, elongated. Oral tentacles wide, short, enrolled, with narrower bases. Parapodia fused together except for short region in posterior midline, forming two small flaps. Flaps partially overlapping in middle (right parapodium partially covers left one), allowing enough space anteriorly for inhalant opening and posteriorly for exhalant siphon to protrude; edge of parapodia tuberculate. External sperm groove clearly visible on right side of animal, between mantle cavity and base of right oral tentacle.

External coloration: Background colour variable, typically brown, greyish-green, or red; tubercles often lighter than background coloration, sometimes with papillae of different colours (pink). Mantle margin with same general colour as rest of body with light spots. Ventral side typically same colour as rest of body with numerous white spots.

Digestive system: Radular formulae: 25 × 86.1.84 ( CPIC 00547); 43 × 111.1.95 ( CPIC 00551); radulae slightly asymmetrical, with few more teeth in one half-row. Rachidian teeth with robust central cusp, two large denticles on each side ( Fig. 14A, D), two innermost denticles much larger than two outermost; teeth wider at base, both upper and lower ends of teeth convex. Some rachidian teeth with small sharp conical extensions on one of apical ends (typically right one). Lateral teeth elongate, with long bases ( Fig. 14B, E); each cusp with basal, narrow, inner denticle, apex bifurcated into two blunt denticles, one slightly larger. Outer teeth very elongate, with or without secondary denticle, apex bifurcate or simple ( Fig. 14C, F). Jaws with simple, irregular denticles ( Fig. 14J). Post-radular armature with numerous spines with elongate bases and triangular cusps ( Fig. 14K). Gizzard plates variable in shape, typically irregular, with numerous striations ( Fig. 14G–I).

Shell: Shell flat, well calcified, oval to elongate ( Fig. 15); nucleus conspicuous, formed by single lobe ( Fig. 14L–P). On apertural view with nucleus on upper end, left side of shell either concave or straight. Right side of shell convex, narrower near nucleus, typically widening slightly towards mid-length, abruptly in some specimens; some shells very narrow across entire length. Widest portion of shell mid-length backward. Sculpture of conspicuous growth lines.

Male reproductive system: Penial canal large and muscular, with a series of tubercles and two proximal retractor muscles, on each side of the opening into the body wall ( Fig. 16B, C). The penial canal widens gradually into the distal end, right before narrowing abruptly and widening again into a small, vesicular-like penial sheath, which contains the penial papilla. The distal end of the penial sheath connects to a long and strong retractor muscle attaching to the ventral side of the body wall internally. The penial papilla is small, triangular, with a longitudinal groove ( Fig. 16A).