Doryteuthis pleii (Blainville, 1823) (Figs. 1, 3, 5, 6)

Synonymy see Cohen (1976). Complement:

Loligo plei [sic]: Cohen, 1976: 341 (figs. 25-26); Whitaker,

1980: 39; Hanlon et al., 1983: 638 (fig. 4); Roper

et al., 1984: 98; Haimovici & Perez, 1991: 227; Costa & Fernandes, 1993: 753 (tab. 1); DiMarco & Hanlon, 1997: 93 (fig. 2); Anderson, 2000: 606 (tab. 2); Jackson & Forsythe, 2002: 677; Perez et al., 2002: 271 (tab. 2); Pineda et al., 2002:71 (tab.1); Cardoso et al., 2005:373 (tab. 4); Martins & Perez, 2006: 24 (fig. 1); Perez et al., 2006: 750 (fig. 1); Rodrigues & Gasalla, 2008: 634 (tab. 1).

Doryteuthis plei [sic]: Brakoniecki, 1984: 436; Haimovici & Andriguetto, 1986 _ENREF_13: 5 (fig. 1); Grassi et al., 1990: 260; Vecchione et al., 2005: 25 (tab. 2); Jereb & Roper, 2010: 55 (fig. 83); Marian, 2010 _ENREF_20: 2 (fig.1); de Pasquier & Carroz,2012:37 (fig.4); de Aguiar et al., 2012: 17 (tab. 1); Marian, 2012: 250 (figs. 1-21); Marian & Domaneschi, 2012: 93 (figs. 1-20); de Luna Sales et al., 2013: 294 (fig. 1); Barcellos & Gasalla, 2014: 3 (figs. 1, 2, 3 and 6).

Doryteuthis pleii: Allcock et al., 2015: 18 (tab. 2); Willems et al., 2015: 28 (tab. 2).

Type locality: Martinique, West Indies, western North Atlantic Ocean.

Holotype: Muséum National d’Histoire Naturelle, Paris [despite the label of the specimen thought to be the holotype is missing ( Cohen, 1976)].

Material examined: Brazil; Rio de Janeiro: off São João da Barra, MZSP 15294, 24 spm. (“Exp. Dep. Zool.” coll., July 1963). off Farol de São Thomé, 22°08′S, 41°00′W, MZSP 27095, 22 m, 1 spm. (R/V “ W. Besnard ” coll., 12 March 1971). off Cabo Frio, 22°51′S, 41°52′W, MZSP 27092, 48 m, 17 spm. (R / V “ W. Besnard ” coll., 01 December 1970). 22°59′S, 42°06′W, MZSP 102761, 25- 33 m, 2 spm. (1 June 1987). off Saquarema, 23°02′S, 42°37′W, MZSP 101893, 50- 62 m, 1 spm. (19 November 2007). off Niterói, 23°05′S, 43°06′W, MZSP 110029, 30- 50 m, 2 spm. (Mendonça Jr. coll., 05 December 2012). off Angra dos Reis, Vila Velha Beach, MZSP 27063, 1 spm. (“ Exp. Dep. Zool. ” coll., 21 July 1966). off Paraty, 23°11′S, 44°38′W, MZSP 58954, 2 spm. (Simone coll., 12 January 2006). MZSP 27061, 13 spm. (July 1983). MZSP 32965, 2 spm. (Fishermen coll., March 2001). São Paulo: off Ubatuba, MZSP 27076, 8 spm. (Boffi coll., 08 February 1970). MZSP 101892, 1 spm. (Caires coll., 2011). MZSP 27084, 2 spm. (Local fishermen coll., 12 March 1970). Lázaro Beach, MZSP 27087, 2 spm. (Sazima coll., 04 April 1970). off Maranduba, 23°39′S, 45°05′W, MZSP 27101, 28- 32 m, 4 spm. (R / V “ W. Besnard ” coll., 05 December 1970).off Santos, MZSP 90936,2spm.(Cunha coll., March 2009). MZSP 27097, 42 m, 4 spm. (R / V “ W. Besnard ” coll., 05 December 1970). MZSP 86427, 8 spm. (R / V “Miragem Boat” coll., March 1988). off Praia Grande, Boqueirão, 24°01′S, 46°25′W, MZSP 80342, 1 spm. (Simone coll., 1997). 24°05′S, 46°20′W, MZSP 110145, 2 spm. (10 April 2012). off São Sebastião, Alcatrazes archipelago. 24°07′S, 45°40′W, MZSP 102400, 46 m, 3 spm. (Amaral coll., 04 October 2011). 24°07′S, 45°40′W, MZSP 102401, 46 m, 2 spm. (Amaral coll., 04 October 2011). 24°24′S, 46°05′W, MZSP 27159, 52 m, 25 spm. (R / V “ W. Besnard ” coll., 21 January 1971). off Itanhaém, 24°24′S, 46°18′W, MZSP 27093, 45 m, 13 spm. (R / V “ W. Besnard ” coll., 05 March 1971). off Peruíbe, 24°20′S, 46°49′W, MZSP 27094, 79 m, 28 spm. (R / V “ W. Besnard ” coll., 06 March 1971). 24°38′S, 45°05′W, MZSP 27102, 102 m, 6 spm. (R / V “ W. Besnard ” coll., 06 March 1971). off Barra do Una, 24°40′S, 46°49′W, MZSP 27096, 38 m, 10 spm. (R / V “ W. Besnard ” coll., 06 December 1970). off Cananéia, Castilho Island – ESEC Tupiniquins, 25°00′S, 47°55′W, MZSP 83165, 5 spm.(Guimarães, Lima & Amaral coll., 24 January 2007). 25°26′S, 45°35′W, MZSP 27091, 125- 128 m, 10 spm. (R / V “ W. Besnard ” coll., 05 July 1970). 25°44′S, 46°37′W, MZSP 27089,111 m, 6 spm. (R / V “ W. Besnard ” coll., 04 March 1971). Paraná: off Pontal do Sul, 25°36′S, 47°30′W, MZSP 27104, 50 m, 2 spm. (R / V “ W.Besnard ” coll., 07 December 1970). 25°59′S, 47°15′W, MZSP 27105, 77- 82 m, 4 spm. (R / V “ W. Besnard ” coll., 07 December 1970). Santa Catarina: São Francisco do Sul Island, 26°17′S, 48°22′W, MZSP 27162, 10 spm. (R / V “ W. Besnard ” coll., February 1977). off Itajaí, MZSP 92949, 51 m, 2 spm. (R / V “NPq Soloncy Moura” coll., 09 November 2009). off Imbituba, 28°18′S, 48°34′W, MZSP 27090, 50 m, 12 spm. (R / V “ W. Besnard ” coll., 01 March 1971). Rio Grande do Sul: off Balneário Pinhal, 30°37′S, 49°25′W, MZSP 27058, 150 m, 4 spm. (GEDIP Project coll., 05 August 1972).

Distribution: Cape Hatteras (36°N) to northern Argentina (35°S) usually associated with Brazilian warm current. Also including Gulf of Mexico, Caribbean Sea, Bermuda and Bahamian and Caribbean islands (Jereb & Roper, 2010).

Description

Mantle: Cylindrical; muscular (~ 91 mm; Table 1); usually long and slender (Fig. 1A), ~ 4-times longer than wide.

Head: Eye diameter approximately 13% of mantle length (Table 2). Width similar to length values.

Chromatophores: Widely distributed, with higher densities on dorsal region including head (around eyes), dorsal midline of mantle (highly concentrated beginning of fins), fins and arms/tentacles. Mature males presenting striped pattern along ventral surface of mantle (Fig. 1A).

Fins: Rhomboid, slightly ovate comprising ~ 48% ML (Table 2), connected posteriorly to posterior end of mantle (Fig. 1A).

Arms: Muscular, broad at base and gradually tapering into thin tips; formula III>IV>II>I (~ lengths: 33 mm, 30 mm, 29 mm, 23 mm);biserial suckers.Hectocotylized portion ~ 39% of left ventral arm length reaching arm’s tip (Table 2), only left row of suckers modified into small peduncles. Bar between row of peduncles and row of suckers connecting them in ladder pattern (Fig. 1E).

Tentacle: Club long and thin, ~ 28% ML (Table 2), carpus, manus and dactylus easily defined; 4 rows of suckers diagonally distributed along club, more visible at manus and dactylus regions; 2 central rows at manus region much bigger than other suckers. Each sucker with small chitin ring of teeth inside. Longitudinal muscle around sucker visible (Figs. 1B, 1D).

Circulatory system: Normal Loliginidae form (Fig. 6A). One central systemic croissant-shaped heart; two small- er peripheral branchial round hearts associated to gills. Two aortae: (i) anterior aorta delivering blood to head, arms and tentacles; (ii) posterior aortic trunk, divided in: (a) visceral aorta, sending blood to posterior part of mantle and visceral organs; (b) mantle aorta, distributing blood to anterior region of mantle. Blood return to heart (i) from posterior vena cava and (ii) from lateral nephridial vessel, both ending in a branchial heart. Branchial artery leaving branchial heart, and ending in gills complex; after this, returning to heart through branchial vein (Fig. 3A).

Digestive system (Figs. 5, 6): Pigmented tip of upper beak short and robust (Fig. 1C). Radula usual for the family; rachidian and first lateral teeth bearing primary large projection and secondary smaller one(s) (Fig. 4A). Usual digestive system for Loliginidae: buccal bulb occupying ~ half of inner head volume. Esophagus narrow, running straight posteriorly along ~ ⅕ of mantle length. Stomach spherical, wall weakly muscular, occupying ~ 20% of visceral volume, located in middle level of visceral sac; posterior caecum wide, extending up to posterior mantle space. Intestine narrow, originating in YY region of stomach, performing simple loop in anterior region of visceral sac. Anus with two papillae, located in middle level of pallial cavity, preceded by short flexible stalk. Digestive gland wide and located between four retractile muscles of head and funnel. Esophagus passing through digestive gland ventro-dorsally.

Male genital system: Normal genital system for Loliginidae, including characters of penis, Needham sac, spermatophoric gland and a single testicle (Figs. 6A, 6B). Hectocotylus described above.

Female genital system: Normal genital system for Loliginidae, including characters of vagina, oviduct,oviducal gland, ovary, a pair of nidamental glands and pair of accessory nidamental glands (Fig. 5). Nidamental glands more uniform, with no slender tip (Fig. 5B).

Central nervous system (Fig. 3B): Central mass slightly divided into anterior and posterior regions. Giant fibers running from posterior end, along dorsal portion of mantle; stellate ganglion visible beneath gills. One pair of optic lobes mushroom-shaped, flanking central mass on each side. Several nerves running anteriorly towards arms and tentacles, forming ring around buccal bulb (Fig. 3B: ib).