taxonID	type	description	language	source
03968781AD2BFFE23412F8F0FE8A9B49.taxon	description	Specimens of Dolabrifera contain secondary metabolites with potential medical applications. For example, the propionate-derived compound Dolabriferol with antibiotic properties (Kigoshi & Kita, 2015) was originally characterized from specimens collected in the Caribbean region (Ciavatta et al., 1996). Subsequent studies on other Caribbean animals revealed the presence of chemically related, but distinct compounds (Jiménez-Romero, González & Rodríguez, 2012). Further research on specimens from the Eastern Pacific resulted in the isolation of 5 α, 8 α- epidioxycholest- 6 - en- 3 β- ol, a compound with antileishmanial properties (Clark et al., 2013). This chemical diversity may be suggestive of a more complex taxonomy than previously recognized. The recent discovery of cryptic species complexes in other biomedically important groups of sea slugs (Krug et al., 2013; Lindsay & Valdés, 2016) highlights the need for modern approaches to investigate the biodiversity of such groups. In this article, we examine the systematics of Dolabrifera dolabrifera and D. brazieri using a combination of molecular and anatomical data to determine the taxonomic status of these two nominal species and unveil possible cryptic diversity in this group.	en	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: 31-65
03968781AD2DFFE434A6F9DAFA439ECC.taxon	diagnosis	Diagnosis: Body oval to elongate, wider posteriorly, narrowing gradually towards head. Oral tentacles, rhinophores slit, flattened, wider distally. Oral tentacles emerging anteriorly from the head, clearly differentiated. Rhinophores dorsal, typically smooth. Parapodial lobes reduced, located behind body midline, fused posteriorly, overlapping partially in middle. Dorsum typically covered with tubercles of different sizes. Shell solid, calcified, reduced, flat, elongate, subtriangular, wider posteriorly, tapering anteriorly into a conspicuous nucleus. Jaws reduced to small section of the labial cuticle with simple denticles. Radula with large, subtriangular rachidian teeth, with several denticles on each side of the cusp; lateral teeth with long, coarsely denticulate cusps. The posterior end of buccal bulb, at insertion of the oesophagus, with two elongate structures (post-radular armature), of numerous elongate denticles. Gizzard muscular, with numerous irregular plates. Male reproductive system with distinct penial canal and penial sheath, where penial papilla is typically located. Remarks: Since its original introduction by Gray (1847), several nominal species have been described in or transferred to Dolabrifera, many of them based on shell material or preserved specimens. Nearly all of those species were confirmed to be members of Dolabrifera, but later synonymized with D. dolabrifera (see Engel & Hummelinck, 1936; Eales, 1944) or D. brazieri (see Hedley, 1917). However, the taxonomic assignment of two of those species is questionable. The first one, Dolabrifera holboelli Bergh, 1872, was described from a single specimen collected in ‘ Mare grönland’ (= Greenland Sea). Bergh (1872: pl. 5, figs 1 – 24) provided illustrations of the preserved animal, shell, radula, jaw, gill and other anatomical details (some of them reproduced in Supporting Information, Fig. S 2 S – U). All descriptions are consistent with the characteristics of a member of Dolabrifera, and examination of the holotype (ZMUC GAS- 2163) confirms the generic placement of this species in Dolabrifera. However, there are no confirmed reports of Dolabrifera from Greenland, nor from other temperate or cold-water regions in the North Atlantic; thus, the most likely explanation for this record is that the specimen was mislabelled. Based on the radular and shell morphology, D. holboelli resembles the tropical Indo-Pacific species D. dolabrifera because the lateral teeth have bifurcated apices with two equally large denticles, and the shell widens abruptly near the apical end, but none of these traits are conclusive. In the absence of molecular data or information on the geographical origin of the holotype, D. holboelli is here treated as an uncertain species. The second problematic species, Dolabrifera pelsartensis O’Donoghue, 1924, was described from ‘ Pelsart Island’ (Pelsaert Island), Western Australia, based on two preserved specimens. O’Donoghue (1924: pl. 27, fig. 3, pl. 29, figs 26 – 28) illustrated a preserved animal (Supporting Information, Fig. S 2 R) and a radula, which shows mid-lateral teeth with two to three denticles and simple cusps. No illustrations of the shell or other anatomical features were provided. With the available information, it is difficult to compare D. pelsartensis with the species here described. The radular morphology is most similar to that of D. brazieri, which appears to be endemic to temperate Eastern Australia and New Zealand. Examination of material from Western Australia is necessary to clarify the status of this species. Another species, Dolabrifera triangularis Watson, 1884, has almost certainly been erroneously assigned to Dolabrifera. Watson (1884) described D. triangularis based on a shell collected from Simon’s Bay, South Africa. Although Watson (1884) described the shell in detail, provided no illustrations and the live animal is unknown. Years later, Watson (1886: 673 – 674, pl. 50, fig. 7; Supporting Information, Fig. S 2 V, W) redescribed and illustrated this shell, which is nearly triangular, short, with a large nucleus. The only remaining syntype (NHMUK 1887.2.9.2296) here examined, matches this description, but is much shorter and thicker that other shells of Dolabrifera spp. In addition, no specimens of Dolabrifera have been documented in False Bay and surrounding areas despite the comprehensive studies (Gosliner, 1987; Zsilavecz, 2007). Based on the available evidence, we cannot identify D. triangularis and therefore is here regarded as a nomen dubium.	en	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: 31-65
03968781AD2CFFEA3446FBC5FEEA9A66.taxon	description	(FIGS 3 A – G, 4 – 6)	en	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: 31-65
03968781AD2CFFEA3446FBC5FEEA9A66.taxon	description	Dolabrifera pacifica Pease in Reeve, 1868: pl. 1, fig. 3. Type locality: Islands of the Pacific. Dolabrifera marmorea Pease in Reeve, 1868: pl. 1, fig. 5 a, b. Type locality: Sandwich Islands (= Hawaiian Islands). Type material	en	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: 31-65
03968781AD2CFFEA3446FBC5FEEA9A66.taxon	description	Dolabrifera marmorea Pease in Reeve, 1868, one syntype at NHMUK 20160139. Dolabrifera pacifica Pease in Reeve, 1868, two syntypes at NHMUK 20160140. Material examined Hekili Point, Maui, Hawaiian Islands, 1 m depth, 21 June 2016, one specimen 19 mm preserved length (CPIC 01654). Maliko Bay, Maui, Hawaiian Islands, 1 m depth, 22 June 2016, five specimens 5 – 14 mm preserved length (CPIC 01657). Mama’s Fish House Reef, Maui, Hawaiian Islands, 1 m depth, 25 June 2016, five specimens 9 – 12 mm preserved length (CPIC 01696); 27 June 2016, one specimen 4 mm preserved length (CPIC 01718); 28 June 2016, two specimens 2 – 3 mm preserved length (CPIC 01752). Sharks Cove, O‘ahu, Hawaiian Islands, 1 m depth, 5 July 2016, one specimen 10 mm preserved length (CPIC 01770). Kaloko, Kona, Hawai‘i, Hawaiian Islands, 26 July 2014, two specimens 16 – 19 mm preserved length, leg. S. Medrano (CPIC 01130). Onekahakaha, Hilo, Hawai‘i, Hawaiian Islands, 26 July 2014, two specimens 12 – 20 mm preserved length, leg. S. Medrano (CPIC 01132). Merizo, Guam, intertidal, 16 June 2009, one specimen 12 mm preserved length, leg. E. Ornelas-Gatdula (CPIC 00413). In front of Bali Sol Hotel, Nusa Dua, Bali, Indonesia (8 ° 48.0 ′ S, 115 ° 14.0 ′ E), intertidal, 22 May 1986, two specimens 35 – 40 mm preserved length, leg. J. McLean (LACM 1986 - 164.16). Cement Mixer Reef, Madang, Papua New Guinea, 4 December 2012, one specimen 14 mm preserved length, leg. J. Goodheart (CPIC 00820). Rempi, Madang, Papua New Guinea, 21 November 2012, one specimen 10 mm preserved length, leg. Papua New Guinea Biodiversity Expedition 2012 (CASIZ 191380). Kranket Island, Madang, Papua New Guinea (5 ° 11 ′ 30.9 ″ S, 145 ° 49 ′ 6.7 ″ E), 6 November 2012, one specimen 6 mm preserved length, leg. Papua New Guinea Biodiversity Expedition 2012 (CASIZ 191000). Platier Flacourt, Taolagnaro, Madagascar (25 ° 1.7 ′ S, 47 ° 0.1 ′ E), 13 May 2010, two specimens 24 – 25 mm preserved length, leg. Atimo Vatae Expedition (CASIZ 194443). Plage Monseigneur, Taolagnaro, Madagascar (25 ° 2.1 ′ S, 46 ° 59.9 ′ E), 15 May 2010, two specimens 18 – 26 preserved length, leg. Atimo Vatae Expedition (CASIZ 1 9 4 4 2 8). Shib Ammar, Saudi Arabia (19 ° 34.232 ′ N, 40 ° 0.519 ′ E), 20 m depth, 3 March 2013, tissue sample, leg. A. Anker (CASIZ 192302). Description External morphology: Body up to 100 mm long, oval to elongate, flattened, broader posteriorly, narrowing gradually towards head. Dorsum covered with small, rounded, scattered tubercles, each with retractile, highly ramified, papilla on apex; tubercles typically tightly arranged, giving animal bumpy (Fig. 3 A, G) or rough appearance (Fig. 3 C – F), in some specimens surrounded by large areas of smooth tissue (Fig. 3 B). 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. External sperm groove clearly visible on right side of animal, between mantle cavity and base of right oral tentacle. Colour: Background colour variable, pink, purple, dark brown, or red, with spots and / or irregular patches of different colours. Mantle margin with same general colour as rest of body. Ventral side typically brown with numerous white, green or brown spots. Digestive system: Radular formulae: 38 × 78.1.41 (CPIC 00336), 37 × 62.1.56 (CPIC 00336), 25 × 58.1.70 (LACM 1986 - 164.1), 28 × 56.1.46 (CPIC 00820); radulae typically asymmetrical, with more teeth in one half-row. Rachidian teeth with robust central cusp, two large denticles on each side (Fig. 4 A, D); teeth wider at base, both upper and lower ends of teeth convex. Lateral teeth elongate (Fig. 4 B, E), with long bases; each cusp with basal, blunt, inner denticle, apex bifurcated into two blunt denticles equal in size. Outer teeth very elongate (Fig. 4 C, F), with or without a secondary denticle, apex bifurcate or simple. Jaws with simple, irregular denticles (Fig. 4 J). Two elongate structures (post-radular armature) located posterior to radula, at oesophagus insertion point, with numerous spines with elongate bases, triangular cusps (Fig. 4 K). Gizzard plates variable in shape, typically irregular, with some striations more visible near centre (Fig. 4 G – I). Shell: Shell flat, well calcified, oval (Fig. 5); nucleus conspicuous, formed by single lobe, occasionally two (Fig. 4 L – O). 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 abruptly near apical end, but more gradually in some specimens. Widest portion of shell mid-length forward, generally close to nucleus. Sculpture of conspicuous growth lines. Male reproductive system: Penial canal large, muscular, lacking tubercles, with two proximal retractor muscles, one on each side of opening into body wall (Fig. 6 C, D). Penial canal narrows and expands abruptly into oval penial sheath containing penial papilla. Oval section connects to large, strong retractor muscle connecting to ventral side of body wall internally. Penial papilla elongate, with curved cusp and longitudinal groove (Fig. 6 A, B).	en	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: 31-65
03968781AD26FFED37FFFA38FDCF9BDD.taxon	description	(FIGS 3 H, 7 – 9)	en	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: 31-65
03968781AD26FFED37FFFA38FDCF9BDD.taxon	materials_examined	Type material	en	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: 31-65
03968781AD26FFED37FFFA38FDCF9BDD.taxon	materials_examined	Material examined Lord Howe Island, New South Wales, Australia (31 ° 32.94 ′ S 159 ° 03.72 ′ E), 1887, six specimens, 32 – 45 mm preserved length (AM C. 54298). Boat Cove, Raoul Island, Kermadec Islands, New Zealand (29 ° 16.79898 ′ S, 177 ° 53.66268 ′ W), 14 May 2011, 0.25 m depth, one specimen, 39 mm preserved length (AM C. 475882). NW corner of North Meyer Island, Kermadec Islands, New Zealand (29 ° 14.4984 ′ S, 177 ° 52.6734 ′ W), 5 – 15 m depth, 13 May 2011, one specimen 45 mm preserved length (AM C. 475783). Description External morphology: Body up to 150 mm long, oval to elongate, flattened. Dorsum covered with large, blunt, rounded to semi-conical tubercles, each with retractile, ramified, papilla on apex; large tubercles typically surrounded by smaller rounded tubercles (Fig. 3 H). Rhinophores enrolled, elongated. Oral tentacles wide, flattened, 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. 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 or greenish-brown, sometimes pale grey or nearly black, with irregular darker, lighter areas. Dorsum often covered with dark brown network of anastomosing lines surrounding tubercles, which do not penetrate mantle margin, rhinophores. Mantle margin typically green, with whitish spots. Digestive system: Radular formulae: 33 × 92.1.80 (AM C. 54298); radulae asymmetrical, with more teeth in one half-row. Rachidian teeth with robust central cusp, two large denticles on each side (Fig. 7 A); teeth wider at base, both upper and lower ends of teeth convex. Lateral teeth elongate (Fig. 7 B), with long bases; each cusp with proximal and distal elongate denticle on inner side, apex blunt, simple. Outer teeth very elongate (Fig. 7 C), with bifurcate apices. Jaws with simple, irregular denticles (Fig. 7 F). Post-radular armature with numerous spines with elongate bases, triangular cusps (Fig. 7 G). Gizzard plates variable in shape, typically irregular, with some striations more visible near the centre (Fig. 7 D). Shell: Shell flat, well calcified, oval (Fig. 8); nucleus conspicuous, formed by single lobe connected to rest of shell by narrow area (Fig. 7 H, I). On apertural view with nucleus on upper end, left side of shell straight, sometimes slightly concave or convex. Right side of shell convex, narrower near nucleus, widening gradually near mid-length. Widest portion of shell mid-length or slightly forward. Sculpture of conspicuous growth lines. Male reproductive system: Penial canal large, muscular, lacking tubercles, with two proximal retractor muscles, one on each side of opening into body wall (Fig. 9 B, C). Penial canal narrows and expands abruptly into elongate penial sheath section containing penial papilla. This section ends in large, strong retractor muscle connecting to ventral side of body wall internally. Penial papilla elongate, with curved cusp and longitudinal groove (Fig. 9 A).	en	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: 31-65
03968781AD3BFFF237D4FC01FC359A70.taxon	description	(FIGS 10 A – D, 11 – 13)	en	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: 31-65
03968781AD3BFFF237D4FC01FC359A70.taxon	materials_examined	Type material	en	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: 31-65
03968781AD3BFFF237D4FC01FC359A70.taxon	materials_examined	Material examined Lower Matecumbe Key, Florida, USA, intertidal, 22 September 1950, four specimens 21 – 34 mm preserved length (LACM 1955 - 59.4). Arrecife Gallega, Veracruz, Veracruz, Mexico, 27 June 2014, one specimen 47 mm preserved length (CNMO 6611). Veracruz, Mexico, 16 June 1973, six specimens 30 – 45 mm preserved length, leg. J. W. Tunnell (LACM 179119). Arrecife Isla Lobos, Veracruz, Mexico, intertidal, 3 June 1973, three specimens 18 – 44 mm preserved length, leg. J. W. Tunnell (LACM 179120). Urchin Cove, Jamaica, 24 May 2006, one specimen 27 mm preserved length, leg. Ann DuPont (LACM 173238). East side of Playa El Yaque’s Lagoon, Isla Tortuga, Venezuela, 18 March 2010, 0.5 m depth, one specimen 55 mm preserved length (ZMUB 84907.1); one specimen 54 mm preserved length (ZMUB 84907.2). Bocas del Drago, Bocas del Toro, Panama, 28 July 2015, one specimen 22 mm preserved length (CPIC 01563). Description External morphology: Body up to 90 mm long, oval to elongate, flattened, broader posteriorly, narrowing gradually towards head. Dorsum covered with numerous tubercles of different sizes, each with retractile, simple papilla on apex; larger tubercles surrounded by smaller ones (Fig. 10 A – D), tightly arranged, giving animal bumpy appearance. 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, brown or green, with spots and / or irregular patches of different colours. Mantle margin semi-translucent, with same general colour as rest of the body and light spots. Ventral side typically same colour as rest of the body with numerous white spots. Digestive system: Radular formulae: 34 × 90.1.77 (CPIC 01563); 35 × 90.1.95 (LACM 173238); radulae typically asymmetrical, with more teeth in one half-row. Rachidian teeth with robust central cusp, two large denticles on each side (Fig. 11 A, D); teeth wider at base, both upper, lower ends of teeth convex. Some rachidian teeth with sharp conical extensions on one of the apical ends (typically right one). Lateral teeth elongate, with long bases (Fig. 11 B, E); each cusp with basal, blunt, inner denticle, apex bifurcated into two blunt denticles, one clearly larger. Outer teeth very elongate, with or without secondary denticle, apex bifurcate or simple (Fig. 11 C, F). Jaws with simple, irregular denticles (Fig. 11 J). Post-radular armature with numerous spines with elongate bases, triangular cusps (Fig. 11 K). Gizzard plates variable in shape, typically irregular, with some striations more visible near centre (Fig. 11 G – I). Shell: Shell flat, well calcified, oval (Fig. 12); nucleus conspicuous, formed by single lobe (Fig. 12 A – D, I, J), occasionally two (Fig. 12 E – H, K, L). 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 abruptly near mid-length, more gradually in some specimens. Widest portion of shell mid-length or slightly forward. Sculpture of conspicuous growth lines. Male reproductive system: Penial canal long and muscular, lacking tubercles, with two proximal retractor muscles, one on each side of opening into body wall (Fig. 13 B – D). Penial canal ends in long retractor muscle connecting to ventral side of body wall internally. No distinguishable penial sheath. Penial papilla very long, occupies almost entire penial canal (Fig. 13 A).	en	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: 31-65
03968781AD3FFFF537F5F921FA529A46.taxon	description	(FIGS 3 I – J, 14 – 16)	en	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: 31-65
03968781AD3FFFF537F5F921FA529A46.taxon	materials_examined	Type material Three syntypes at ANSP 67517 (dry) and ANSP A 7048 (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. 3 I, 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. 14 A, 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. 14 B, 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. 14 C, F). Jaws with simple, irregular denticles (Fig. 14 J). Post-radular armature with numerous spines with elongate bases and triangular cusps (Fig. 14 K). Gizzard plates variable in shape, typically irregular, with numerous striations (Fig. 14 G – I). Shell: Shell flat, well calcified, oval to elongate (Fig. 15); nucleus conspicuous, formed by single lobe (Fig. 14 L – 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. 16 B, 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. 16 A).	en	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: 31-65
03968781AD3CFFF9363DF9CFFE249CE5.taxon	description	(FIGS 10 G – H, 17 – 19)	en	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: 31-65
03968781AD3CFFF9363DF9CFFE249CE5.taxon	materials_examined	Type material	en	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: 31-65
03968781AD3CFFF9363DF9CFFE249CE5.taxon	materials_examined	Material examined Urchin Cove, St Ann’s Bay, Jamaica (18 ° 27.20 ′ N, 77 ° 13.55 ′ W), 0.3 – 0.6 m depth, 25 May 2006, two specimens 10 – 15 mm preserved length, leg. A. DuPont (LACM 173265). Lauderdale-by-the-Sea, Florida, USA, 8 m depth, 18 June 2014, one specimen 21 mm preserved length, leg. A. Dimitris (CPIC 01134). Vista Alegre, Curaçao (12 ° 0.50 ′ N, 68 ° 53.30 ′ W), intertidal, 23 April 1939, nine specimens 14 – 40 mm preserved length (LACM 1939 - 215.1). Description External morphology: Body up to 80 mm long, oval, flattened, broader posteriorly, narrowing slightly towards head. Dorsum covered with densely packed tubercles of different sizes, each with elongate papilla on apex, some ramified at tips; larger tubercles surrounded by smaller ones, each bearing papilla, giving animal appearance of being densely covered with papillae (Fig. 10 G – H). 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, red, brown or green, sometimes white, with spots and / or irregular patches of different colours. Mantle margin semi-translucent, with same general colour as rest of body and few small white dots. Ventral side typically same colour as rest of the body with numerous white spots. Digestive system: Radular formulae: 32 × 61.1.60 (LACM 173265 A); 27 × 50.1.57 (LACM 173265 B); radulae nearly symmetrical, with about same number of teeth on each row. Rachidian teeth with robust central cusp, two large denticles on each side (Fig. 17 A, D); teeth wider at base, both upper and lower ends of teeth convex. Lateral teeth elongate, with long bases (Fig. 17 B, E); each cusp with two to three basal, blunt, inner denticles, apex bifurcated into two blunt denticles equal in size. Outer teeth very elongate, with secondary denticle, apex bifurcate (Fig. 17 C, F). Jaws with simple, irregular denticles (Fig. 17 J). Post-radular armature with numerous spines with elongate bases and low cusps (Fig. 17 K). Gizzard plates variable in shape, typically short, wide, with some striations more visible near one end (Fig. 17 G – I). Shell: Shell flat, well calcified, elongate, width / length ratio greater than 1: 2 (Fig. 18); nucleus inconspicuous, short, formed by single lobe. On apertural view with nucleus on upper end, left side of shell either concave or straight. Right side of shell convex, narrower near nucleus, widening gradually towards mid-length. Widest portion of shell mid-length or slightly backward. Sculpture of conspicuous growth lines. Male reproductive system: Penial canal long, muscular, lacking tubercles, with two proximal retractor muscles, one on each side of opening into body wall (Fig. 19 B, C). Penial canal ends in long retractor muscle connecting to ventral side of body wall internally. No distinguishable penial sheath. Penial papilla very long, occupies almost entire penial canal (Fig. 19 A).	en	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: 31-65
03968781AD30FFFF3632FB14FAF89A4A.taxon	description	(FIGS 10 E – F, 20 – 22) Type material Holotype at ZSM Mol 20160193, Porto Moniz, Madeira, September 2014, 18 mm preserved length. Material examined Porto Moniz, Madeira, September 2014, one specimen 17 mm preserved length, leg. P. Wirtz (ZSM Mol 20160194), one specimen 19 mm preserved length, leg. P. Wirtz (ZSM Mol 20160195), one specimen 20 mm preserved length, leg. P. Wirtz (ZSM Mol 20160196), one specimen 20 mm preserved length, leg. P. Wirtz (ZSM Mol 20160197). Ilhéu Bombom, Ilha do Príncipe, São Tomé and Príncipe, 20 January 2009, one specimen 11 mm preserved length (CASIZ 179395). South of Terrebonne Bay, LA, USA (28 ° 5.57 ′ N, 91 ° 1.34 ′ W), 56 – 58 m depth, 4 June 2005, four specimens 12 – 14 mm preserved length, leg. E. Garcia (LACM 179615). Marsh Island, LA, USA (27 ° 98.34 ′ N, 92 ° 22.42 ′ W), 68 – 86 m depth, 22 June 2005, two specimens 11 – 16 mm preserved length, leg. E. Garcia (LACM 179619). Description External morphology: Body up to 80 mm long, oval to elongate, flattened, broader posteriorly, narrowing gradually towards head. Dorsum covered with small, rounded, scattered tubercles, each with retractile, highly ramified, papilla on apex (Fig. 10 E, F); tubercles typically surrounded by large areas of smooth tissue. 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. 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 greenish-grey or brown, sometimes red, with spots and / or irregular patches of different colours. Mantle margin with same general colour as rest of body. Digestive system: Radular formula: 40 × 70.1.71 (ZSM Mol 20160195); 38 × 71.1.72 (ZSM Mol 20160196); radulae symmetrical, with just about same number teeth in both half-rows. Rachidian teeth with robust central cusp, two large denticles on each side (Fig. 20 A, D), all similar in size; cusp with or without denticles on both sides; rachidian teeth wider at base, both upper and lower ends of teeth convex. Some rachidian teeth with small extensions on apical ends. Lateral teeth elongate, with long bases (Fig. 20 B, E); each cusp with basal, narrow, inner denticle, apex bifurcated into two blunt denticles equal in length. Outer teeth very elongate, with or without secondary denticle, apex bifurcated (Fig. 20 C, F). Jaws with simple denticles (Fig. 20 J). Post-radular armature with numerous spines with elongate bases and triangular cusps (Fig. 20 K). Gizzard plates variable in shape, typically irregular, with numerous striations (Fig. 20 G – I). Shell: Shell flat, well calcified, oval to elongate (Fig. 21); nucleus conspicuous, formed by single lobe (Fig. 20 L – O). On apertural view with nucleus on upper end, left side of shell either concave or straight. Right side of shell convex, narrower near nucleus, widening slightly towards mid-length. Widest portion of shell mid-length. Sculpture of conspicuous growth lines. Male reproductive system: Penial canal large, muscular, with two proximal retractor muscles, on each side of opening into body wall (Fig. 22 B – D). Penial canal widens gradually into oval penial sheath, containing penial papilla. Distal end of penial sheath connects to thick and strong retractor muscle attaching to ventral side of body wall internally. Penial papilla elongate, triangular, with longitudinal groove (Fig. 22 A).	en	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: 31-65
