Systematics of the Onychoteuthidae Gray, 1847 (Cephalopoda: Oegopsida)
Squids in the family Onychoteuthidae Gray, 1847 have been reported from every ocean but the Arctic, are taken frequently in deep-sea fisheries by catch, and are ecologically important in the diets of many marine predators including cetaceans, pinnipeds, sharks, and seabirds. However, the diversity and systematic of the family have remained poorly understood. Of the 60+ nominal species, 12–14 have generally been accepted in recent studies. Challenges to clarity include insufficient species descriptions, original descriptions published in eight languages and often based solely on early life stages, non-designation or subsequent loss of type material, and the existence of several unresolved species complexes. In light of the general systematic disarray of the Onychoteuthidae, a global revision of the family follows, based on ~1500 specimens examined from 19 repositories. Type material has been examined wherever possible; for some species, photographs of type specimens, original illustrations, and/or the original descriptions have provided the only information available. It has not been possible to fully disambiguate taxa in some cases (e.g. Gen. nov. 2), given the limited material and information available, but for all species treated in this revision (25 out of 26 species; no material was available for Kondakovia nigmatullini), descriptions and illustrations are provided to a consistent standard that will enable their reidentification. External and internal morphological characters and states are described for sub adult to adult stages of most species, with external characters reported through ontogeny as permitted by available material. Historically important characters are treated (general external morphology, body proportions, tentacle clubs, photophores, gladius, lower beak, radula), augmented by several more recently recognised characters (palatine teeth, detailed morphology of the tentacular hooks in adults, tentacular suckers in paralarvae, chromatophore patterns). The systematic value of both historical and new morphological characters at the generic and species levels are discussed; at all ontogenetic stages, tentacular club and hook morphology are considered the most valuable characters, although body proportions and gladius also prove useful. Partial disambiguation of the Onychoteuthis banksii complex has been possible in the Pacific and Atlantic Oceans, resulting in the resurrection of Onychoteuthis bergii Lichtenstein, 1818 and Onychoteuthis aequimanus Gabb, 1868, the description of two new species, Onychoteuthis lacrima and Onychoteuthis prolata (in press), and the expansion of one species’ recognised distribution (Onychoteuthis compacta) to include the Atlantic Ocean. The genus Moroteuthis Verrill, 1881 is considered a junior synonym of Onykia Lesueur, 1821, in accordance with the findings of several earlier authors. However, morphological differences in the species ‘Moroteuthis’ ingens necessitate the resurrection of the subgenus Moroteuthopsis Pfeffer, 1908b, with all other Onykia species placed into a new subgenus, Onykia (Onykia). Sexual dimorphism is reported in the beaks of Onykia (Moroteuthopsis) ingens (new comb.), and revised sex-specific equations are given for estimating this species’ biomass based on LRL. Morphological and historical genetic data suggest a more distant relationship between Onykia and the species ‘Moroteuthis’ knipovitchi Filippova, 1972 than was suggested by earlier classifications. This species is therefore considered to represent an undescribed genus, herein referred to as Gen. Nov. 1, which cannot be more fully diagnosed and described at present due to limited material. The generic position of ‘Onykia’ rancureli (Okutani, 1981) is also uncertain; it may be allied to Walvisteuthis virilis Nesis & Nikitina, 1986 (family Walvisteuthidae Nesis & Nikitina, 1986), but confirmation is impossible without examining type material of W. virilis. A second new genus, Gen. Nov. 2, is therefore described for ‘Onykia’ rancureli and several morphological variants reported from the Pacific and Atlantic Oceans. Given that the majority of available onychoteuthid material was collected after 1950, resulting in the descriptions of over half of the generally accepted genera and species since 1960, ongoing collection programmes are necessary to further resolve onychoteuthid systematic.