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Squids of the family Onychoteuthidae Gray, 1847 in the southeastern Pacific Ocean.

Calamares de la familia Onychoteuthidae Gray, 1847 en el Oceano Pacifico suroriental

Hooked squids (family Onychoteuthidae Gray, 1847) inhabit all oceans of the world except the Arctic (Bolstad, 2010). This family is composed of 25 species belonging to seven genera (Bolstad, 2010). In the southeastern Pacific Ocean (SEP), various studies have reported a total of seven onychoteuthid squid species and four genera (Thore, 1959; Roper et al., 1984; Nesis, 1987; Fisher & Hureau, 1988; Rocha et al., 1991; Cardoso, 1991; Alexeyev, 1994; Rocha, 1997; Paredes et al., 1999). More recently, following reviews of specimens from Chilean museums and newer literature, the list has been reduced to five species and three genera (Vega et al., 2001; Vega, 2009; Ibanez et al., 2009). A complete review of the family has also revealed new species, new records, and new combinations (Bolstad, 2007, 2008, 2010). However, few specimens from the SEP were available for these studies. Herein, we have reviewed the literature and specimens from museum collections to elucidate the true onychoteuthid fauna of the SEP. Historically, the most commonly reported species from the SEP has been Onychoteuthis banksii. However, Bolstad (2008, 2010) recognized that many specimens attributable to this species are incorrectly identified, and O. banksii (sensu stricto) appears to only inhabit the Atlantic Ocean. Therefore, it is vital to reassess the Onycho-teuthis specimens collected from the SEP.

The present work was based on a review of the museum collections from the Museo Nacional de Historia Natural (MNHNCL), Santiago, Chile; Museo Zoologico de la Universidad de Concepcion (MZUC), Concepcion, Chile; Coleccion Flora y Fauna, Profesor Patricio Sanchez, Pontificia Universidad Catolica de Chile, Santiago, Chile (SSUC); Santa Barbara Museum of Natural History (SBMNH), Santa Barbara, USA; National Institute of Water and Atmospheric Research, (NIWA), Wellington, New Zealand, and the Museum of New Zealand Te Papa Tongarewa (NMNZ), Wellington, New Zealand. Description, measurements, and counts followed Roper & Voss (1983). The following abbreviations are used: TL: total length; ML: mantle length; MW: mantle width; HL: head length; HW: head width; AL: arm length 1 to 4R/L; FL: fin length; FW: fin width; TnL: tentacle length; CL: tentacle club length; CS: count of carpal suckers on the tentacle club; MH: count of manus hooks on the tentacle club; TPS: count of terminal pad suckers on the tentacle club.

Systematics

Family: Onychoteuthidae Gray, 1847 Genus: Onychoteuthis Lichtenstein, 1818 Type species: Onychoteuthis aequimanus Gabb, 1868 Material examined: Onychoteuthis "banksif" (2 specimens) 70, 98 mm ML (off Valparaiso, southeastern Pacific, 300-400 m depth) MNHNCL 300016. Collected by N. Bahamonde, August 10, 1960. Onychoteuthis "banksii" 70 mm ML (near Eastern Island, caught at surface) MNHNCL 300005. Collected by L. DiSalvo, February 1986.

Comparative material examined: Onychoteuthis aequimanus (3 specimens) 78, 87 and 100 mm ML (Tasman Sea, 37[degrees]S, 167[degrees]E) NMNZ 160477. Collected in 1960. Onychoteuthis aequimanus mantle damaged (Kermadec Islands, 27[degrees]50.8'S, 178[degrees]50.2'E, 20 m depth) NMNZ 074466. Collected June 18, 1976. Onychoteuthis aequimanus 150 mm ML (23[degrees]17.2'S, 173[degrees]1.5'E) NMNZ 181412. Collected in May 2005. Onychoteuthis cf. aequimanus 91 mm ML (40.51[degrees]S, 178.2[degrees]E, 100 m depth) NIWA 32754. Collected in February 1998.

Description. Small squids (ML 70-106 mm), arms medium length (29-57% ML) (Fig. 1). Head with 9-10 nucal folds (Fig. 2). Fins rhombic, length about half of mantle length (52-59% ML). Gladius visible along anterior mantle zone. Arms with two sucker rows; tentacle club length 22-34% ML, with 21-22 large hooks, 8 carpal suckers, and 14-16 terminal pad suckers (Fig. 1, Table 1).

Genus: Onykia Lesueur, 1821

Subgenus: Onykia (Moroteuthopsis) Pfeffer, 1908

Type species: Onykia (Moroteuthopsis) ingens (Smith, 1881)

Material examined: "Moroteuthis" ingens (3 specimens) 93, 87, 90 mm ML (southern Chile, 55[degrees]29'S, 70[degrees]03W, 350-357 m depth) MNHNCL 5747. Collected by S. Aviles & P. Ojeda, December 3, 1977. "Moroteuthis" ingens 380 mm ML (southern Chile, 52[degrees]06'S, 75[degrees]47W) MNHNCL 5805. Collected November 22, 1977.

Comparative material examined: Onykia ingens 178 mm ML (Campbell Plateau, 52[degrees]26'S, 170[degrees]31'E, 480-482 m depth) NMNZ 074116. Collected January 18, 1977. Onykia ingens 115 mm ML (Bounty Plateau, 47[degrees]40.68'S, 178[degrees]0.72'E, 840 m depth) NMNZ 118344. Collected December 9, 1990. Onykia ingens 170 mm ML (43.06[degrees]S, 175.64[degrees]E, 460 m depth) NIWA32754. Collected October 7, 1996.

Description. Large squids (ML 87-380 mm) (Fig. 3). Mantle muscular, rugose, with small, fleshy, irregular 'warts' over all surfaces (Fig. 4). Arms long with two sucker rows (61-97% ML). Fins broad, smooth, rhombic, length about half of mantle length (46-57% ML); tentacle club narrow (Fig. 5), length 29-35% ML, with 25-28 large hooks, 12 carpal suckers, and 13-17 terminal pad suckers (Fig. 2, Table 1).

Genus: Onykia Lesueur, 1821

Subgenus: Onykia (Onykia) Lesueur, 1821

Type species: Onykia (Onykia) robsoni (Adam, 1962) Material examined: "Moroteuthis" robsoni 380 mm ML (off Los Vilos, central Chile, 32[degrees]S, 450 m depth) MNHNCL 1950.

Comparative material examined: Onykia robsoni 265 mm ML (42.83[degrees]S, 177.2[degrees]E, 809 m depth) NIWA 32750. Collected July 21, 1995.

Description. Large squid (380 mm ML) (Fig. 6). Mantle fleshy, spindle-shaped, narrow cone over posterior (Fig. 6). Dorsal and ventral surfaces of mantle studded with, well-separated, blister-like warts (Fig. 7). Fin length about half of mantle length (55% ML). Arms longs with two sucker rows (55-75% ML); tentacle club length 38% ML, with 25-27 large hooks, 9-19 carpal suckers, and 12-13 terminal pad suckers (Fig. 6, Table 1).

Genus: Kondakovia Filippova, 1972 Type species: Kondakovia longimana Kondakovia nigmatullini Laptikhovsky, Arkhipkin & Bolstad, 2008

Material examined: Kondakovia "longimana" 62 mm ML (44[degrees]44'S, 68[degrees]20'W) MNHNCL 841. Collected April 24, 1984.

Description. Small squid (62 mm ML), arms long with two sucker rows (64-87% ML). Fin broadly rhombic and short (29% ML); tentacle club length 30% ML, with 27 large hooks, 12 carpal suckers, two complete series of marginal suckers and 12-21 terminal pad suckers (Fig. 8, Table 1). Mantle head and arms with reticulated skin sculpture (Fig. 8).

In the southeastern Pacific Ocean (SEP), onychoteuthid squids are poorly represented in regional and international collections, and holdings represent only four species. We reported the presence of four species based on museum collections and an examination of the literature. The genera Onykia and Kondakovia in the SEP are distributed in cold waters of the southern Ocean down to 30[degrees]S, while Onychoteuthis is common in warmer oceanic waters from 30[degrees]N to 30[degrees]S (Ibanez et al, 2009; Bolstad, 2010).

Previous reports of O. "banksii " in Peru and Chile (Thore, 1959; Cardoso, 1991; Rocha, 1997; Paredes et al., 1999; Vega et al., 2001; Ibanez et al., 2009; Vega, 2009) probably correspond to O. aequimanus. Specimens of this species from ML 100-350 mm are common in stomach contents of swordfish (Xiphias gladius) in oceanic waters off Chile between 32[degrees]S-8[degrees]S and 80[degrees]W-5[degrees]W (Ibanez et al., 2004; Castillo et al., 2007). The three specimens of O. aequimanus from MNHNCL represents a new distribution record for this species.

Squids identified as "Moroteuthis'" from MNHNCL belong to the genus Onykia. Onykia ingens inhabits the continental shelf of Chile, from 42[degrees]S southward in cold waters of Antarctic origin deeper than 300 m (Ibanez et al., 2009). Alexeyev (1994) found one juvenile specimen of O. ingens at 39[degrees]04'S, 93[degrees]50'W. This species has been reported in stomach contents of Patagonian toothfish (Dissostichus eleginoides) from the continental slope (>500 m) of southern Chile (43[degrees]S) (Murillo et al., 2008). Onykia robsoni, in Chile, is distributed in more northern waters than O. ingens, from 32[degrees]S southward. In addition, Alexeyev (1994) found several juvenile specimens of O. robsoni between 35[degrees]and 0[degrees]S.

Kondakovia longimana has been reported from stomach contents of hoki (Macruronus novaezelandiae) and southern hake (Merluccius australis) in southern Chile (52[degrees]S, 76[degrees]W) at 300 m depth (Rocha et al., 1991). This squid has also been reported from stomach contents of Patagonian toothfish (D. eleginoides) from the continental slope (>500 m) of southern Chile (43[degrees]S) (Murillo et al., 2008). The only museum specimen (MNHNCL 841) of K. "longimana" examined herein corresponds to a juvenile of K. nigmatullini. It is likely that all records in Chile of K. longimana from stomach contents, for example, correspond to a K. nigmatullini. This specimen represents a new distribution record for this species in addition to type material (holotype and paratype) collected near to Falkland Islands (Laptikhovsky et al., 2008).

Rocha (1997) included Filippovia knipovitchi (Filippova, 1972) in Chilean waters based on the Southern Ocean distribution reported by Fisher & Hureau (1988), but we did not find this species represented in the present material.

We don't found any Onychoteuthid squid from Chile or Peru in the collections of NIWA, NMNZ, SSUC, MZUC and SBNH, only in MNHNCL. Onychoteuthid squids from the southeastern Pacific Ocean have been poorly studied and it is likely that more species from another genera such as Notonykia and Filippovia are also present in these waters. During the development of this work we found three specimens of Notonykia sp. (identified by K. Bolstad) at 280 km off central Chile (34.09[degrees]S, 75.08[degrees]W).

DOI:10.3856/vol44-issue2-fulltext-23

Received: 21 January 2015; Accepted: 27 March 2016

ACKNOWLEDGEMENTS

We thank Sergio Letelier, Bruce Marshall and Sadie Mills for their help in examining specimens from the MNHNCL (Santiago, Chile), NMNZ (Wellington, New Zealand) and NIWA (Wellington, New Zealand). We thank Kathrin Bolstad and Heather Braid (Auckland University of Technology, Auckland, New Zealand) for the comments in the manuscript. This work was partially funded by FONDECYT 3110152.

REFERENCES

Alexeyev, D.O. 1994. New data on the distribution and biology of squids from the Southern Pacific. Ruthenica, 4(2): 151-166.

Bolstad, K.S. 2007. Systematics and distribution of the New Zealand onychoteuthid fauna (Cephalopoda: Oegopsida), including a new species, Notonykia nesisi sp. nov. Rev. Fish Biol. Fish., 17: 305-335.

Bolstad, K.S. 2008. Two new species and a review of the squid genus Onychoteuthis Lichtenstein, 1818 (Oegopsida: Onychoteuthidae) from the Pacific Ocean. Bull. Mar. Sci., 83: 481-529.

Bolstad, K.S. 2010. Systematics of the Onychoteuthidae Gray, 1847 (Cephalopoda: Oegopsida). Zootaxa, 2696: 1-186.

Cardoso, F. 1991. Los calamares y potas (Cephalopoda: Teuthoidea) del mar peruano. Biota, 15(97): 2-13.

Castillo, K., C.M. Ibanez, C. Gonzalez & J. Chong. 2007. Dieta del pez espada Xiphias gladius Linnaeus, 1758 en distintas zonas de pesca frente a Chile central durante el otono de 2004. Rev. Biol. Mar. Oceanogr., 42(2): 149-156.

Fisher, W. & J.C. Hureau. 1988. Fichas FAO de identificacion de especies para los fines de la pesca. Oceano Austral (Areas de pesca 48, 58 y 88, area de la Convencion CCAMLR). Publicacion preparada y publicada con el apoyo de la Comision para la Conservacion de los Recursos Vivos Marinos Antarticos. Roma, I(1): 232 pp.

Ibanez, C.M., P.A. Camus & F.J. Rocha. 2009. Diversity and distribution of cephalopod species of the coast off Chile. Mar. Biol. Res., 5: 374-384.

Ibanez, C.M., C. Gonzalez & L. Cubillos. 2004. Dieta del pez espada Xiphias gladius Linnaeus, 1758, en aguas oceanicas de Chile central en invierno de 2003. Invest. Mar., Valparaiso, 32(2): 113-120.

Laptikhovsky, V., A. Arkhipkin & K.S. Bolstad. 2008. A second species of the squid genus Kondakovia (Cephalopoda: Onychoteuthidae) from the sub-Antarctic. Polar Biol., 32: 21-26.

Murillo, C., C. Oyarzun & I. Fernandez. 2008. Variacion latitudinal y estacional en la dieta de Dissostichus eleginoides Smitt, 1898 (Perciformes: Nototheniidae) en ambientes profundos de la costa centro-sur de Chile. Gayana, 72: 94-101.

Nesis, K.N. 1987. Cephalopods of the world. T.F.H. Publications, New Jersey, 351 pp.

Paredes, C., P. Huaman, F. Cardoso, R. Vivar & V. Vera. 1999. Estado actual del conocimiento de los moluscos acuaticos en el Peru. Rev. Peru. Biol., 6(1): 5-47.

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Rocha, F.J., O. Poblete & N. Bahamonde. 1991. Cefalopodos en contenidos gastricos de Merluccius australis polylepis Ginsburg y Macruronus magellanicus Lonnberg (Chile), Invest. Pesq., 36: 51-65.

Roper, C.F.E. & G.L. Voss. 1983. Guidelines for taxonomic descriptions of cephalopod species. Mem. Nat. Mus. Victoria, 44: 49-63.

Roper, C.F.E., M.J. Sweeney & C.E. Nauen. 1984. Cephalopods of the world. An annotated and illustrated catalogue of species of interest to fisheries. FAO Fish. Syn., 125(3): 1-277.

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Vega, M.A., S. Letelier & E. Carreno. 2001. Coleccion de cefalopodos del Museo Nacional de Historia Natural: catalogo especies de aguas chilenas. Museo Nacional de Historia Natural Santiago, Chile. Publ. Ocas., 57: 1-88.

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Christian M. Ibanez (1) & Alina F. Cifuentes-Bustamante (1)

(1) Departamento de Ecologia y Biodiversidad, Facultad de Ecologia y Recursos Naturales Universidad Andres Bello, Santiago, Chile

Corresponding author: Christian M. Ibanez (ibanez.christian@gmail.com)

Corresponding editor: Cristian Aldea

Caption: Figure 1. Onychoteuthis aequimanus MNHNCL 1954-2. a) Complete specimen, ML 106 mm, b) Club, 29 mm.

Caption: Figure 2. Detail of head, eye and nucal folds of Onychoteuthis aequimanus MNHNCL 1954-2. Complete specimen, ML 106 mm.

Caption: Figure 3. Onykia ingens MNHNCL 5747. a) Complete specimen, ML 93 mm, b) Club, 27 mm, c) carpal suckers, d) manus suckers.

Caption: Figure 4. Detail of mantle warts of Onykia ingens MNHNCL 5747. Complete specimen, ML 93 mm.

Caption: Figure 5. Detail of tentacle club of Onykia ingens MNHNCL 5747. Complete specimen, ML 93 mm.

Caption: Figure 6. Onykia robsoni MNHNCL 1950. a) Complete specimen, ML 380 mm, b) Club 118 mm.

Caption: Figure 7. Detail of mantle warts of Onykia robsoni MNHNCL 1950. Complete specimen, ML 380 mm.

Caption: Figure 8. Kondakovia nigmatullini MNHNCL 841. a) Dorsal view, ML 62 mm, b) ventral view.
Table 1. Measurements (mm) and counts of onychoteuthid squids. For
abbreviations see text, incomplete or damaged.

               MNHNCL           MNHNCL           MNHNCL        MNHNCL
               1954-1           1954-2            1947         5747-1

           Onychoteuthis    Onychoteuthis    Onychoteuthis     Onykia
             aequimanus       aequimanus       aequimanus      ingens

ML               98              106               70            93
MW               20               23               23            24
HL               15               21               23            19
HW               19               21               17            21
FL               53               63               37            43
FW               62               71               47            65
Arms I        29L/30R          31L/33R          25L/30R        60R/60L
Arms II       38L/37R          39L/42R          40L/39R        80R/75L
Arms III      35L/38R         47L/38R *         40L/42R        75R/65L
Arms IV       35L/35R         38L/35R *         35L/36R        80R/80L
TnL              55              105            80R/75L        115R/--
CL              22R              29R            24R/23L        27RJ--
CS               8R               8R             8R/8L         12R1--
MH              22R              21R            22R722L        28R/--
TPS             14R              16R             4R/4L         17R/--

             MNHNCL        MNHNCL         MNHNCL         MNHNCL
             5747-2        5747-3          5805           1950

             Onykia        Onykia         Onykia         Onykia
             ingens        ingens         ingens        robsoni

ML             87            90            380            380
MW             24            24            130             95
HL             25            23             75             56
HW             21            21             80             45
FL             50            51            190            210
FW             68            67            240            185
Arms I       65R/60L       55R/61L      155R*/204L    212R/125L *
Arms II      65R/85L       81R/81L      314R/305L      234R/246L
Arms III     85R/75L       72R/75L      276R/280L      240R/224L
Arms IV      70R/70L       75R/79L      316R/311L      285R/281L
TnL         130R/125L     156R/145L     582R/520L      830R/826L
CL           27R/27L       32R/30L      120R/105L      118R/145L
CS           12R/12L       12R/12L       12R/12L         10R/9L
MH           26R/28L       25R/26L       27R/26L        25R/27L
TPS          16R/L3L       14R/16L       16R/16L        12R/13L

              MNHNCL
                841

            Kondakovia
           nigmatullini

ML              62
MW              15
HL              20
HW              10
FL              18
FW              35
Arms I       40R/35L *
Arms II       54R/45L
Arms III      40R/42L
Arms IV       43R/44L
TnL           81R/70L
CL            19R/--
CS            12R/12L
MH            26R/27L
TPS           12R/21L
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Title Annotation:Short Communication
Author:Ibanez, Christian M.; Cifuentes-Bustamante, Alina F.
Publication:Latin American Journal of Aquatic Research
Date:May 1, 2016
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