Reproduction of material in this information product for resale or other commercial purposes is prohibited without written permission of the copyright holders. It is part of the regular program activities and partial fulfillment of the Organization's role in relation to the identification of marine fisheries resources and biological data (FAO Program Element 232A3). We hope that this additional flexibility will provide convenience and usability for catalog users.
This is the first volume of the completely rewritten, revised and updated version of the original FAO Catalog of Cephalopods of the World (1984). Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae, with 23 genera and the 201 species known up to the date of completion of the volume. In particular, we thank Michel Lamboeuff for his help with the use of the FAO FISHSTAT database and for his review of the graphic sections afterwards.
Color photographs are included in this new version of the catalog to enrich the quality and usability of the book. Roper, for her technical and editorial assistance and invaluable support during many phases of the preparation of this catalogue.
INTRODUCTION
Plan of the Catalogue
Therefore, each major group and family is introduced with general descriptive notes, illustrations of diagnostic characters, highlights of its biology and relevance to fisheries. Species identification should only be attempted after verifying the family using the illustrated key to families. Due to the elasticity of tentacles and arms, total length is not a very accurate measurement.
The full known geographic range of the species, including areas of seasonal occurrence, is given in the text and shown on the map. The known depth range of the species is given, along with information on the salinity and temperature of its habitat where available. Here again, the quality and quantity of available information varies considerably between species.
The current compilation is necessarily incomplete, as only a fraction of the local names applied to specific entities are actually published. The local species name is preceded by the name of the country concerned in capital letters and, where necessary, by geographical specifications in lower case letters.
General Remarks on Cephalopods
Most cephalopod behaviors involve rapid changes in overall color and color patterns, and changes in skin texture, from smooth to heavily papillate, tubercular, or with erect flaps. The captured cephalopod prey is brought to the mouth and killed with the bites of powerful chitinous beaks equipped with powerful muscles. The gonads form a single mass at the posterior end of the mantle cavity, and the female gonoducts may be paired (in egopsids and incirrate octopuses) or single, as in other colloids and nautiluses.
They lack the gelatinous outer matrix found in squid and cuttlefish eggs and the outer shell found in Cirrate octopus eggs: the protective function is "replaced" by the behavior of female parents. The principle results obtained from the research generally confirm a very high growth rate in cephalopods, comparable to that of the fastest growing fish. There is a general consensus that spawning is a final event, despite the high variability in the duration of individual spawning periods (5 to 50% of ontogeny; . Nigmatullin, 2002) as well as the type of spawning, e.g.
The squids have broad sac-like bodies with lateral fins that are narrow and extend the length of the mantle; Several human deaths have been recorded in the western Pacific and Australia due to venomous bites from the small blue-ringed octopus, Hapalochlaenaspp.; the deadly toxin is a neural blocker considered a form of tetrodotoxin.
Interest to Fishery and Role in the Ecosystem
Jigging is the most commonly used method, accounting for almost half of the global cephalopod catch: all squid, mainly ommastrephids, but also some loliginids. The amount of cephalopods caught as bycatch in bottom trawls for fish fisheries attracted increasing attention to these stocks in the 1980s; this led to the development of (mainly) inland trawl fisheries specifically targeting squid, particularly in the South Atlantic Ocean. Bottom trawling can be very dangerous to benthic habitats due to the physical damage it causes to the seabed and associated fauna and due to the lack of selectivity.
5 Distribution of world ommastrephid light fisheries (illustrations based on nighttime satellite imagery) (from Rodhouseet al., 2001). Commercial exploitation of the cosmopolitan family Histioteuthidae could also be considered, as at least one large commercial catch has been made in the North Atlantic (see Okutani, personal communication, in Clarke, 1996). Almost all of our knowledge of the general biology of cephalopods is actually limited to shelf-dwelling species.
Therefore there is an important challenge to deepen our knowledge and to learn the details of the distribution, life history and biology of the exploited species in order to allow the rational use of the reserves. A thoroughly studied case is that of the snapper fishery at Endeavor Bank (Balguerias et al., 2000), which was replaced by a cephalopod fishery between the 1960s and 1970s.
Illustrated Glossary of Technical Terms and Measurements
Armature – The grasping structures of the arms and tentacular bats, including suction cups and/or hooks. Brachial lobe (of the brain) – The most anterior part of the brain located ventral to the esophagus. Cuttlebone (= sepion) – The calcareous (chalky) elongated, supporting protective and buoyant shield in the dorsal part of the mantle of squids (Fig. 18).
Funnel groove – The depression on the posteroventral surface of the head into which the anterior part of the funnel lies (Fig. 17). Gladius length (GL) - Sometimes used as a measurement of body (= cloak) length when direct measurement of the cloak is unreliable (usually due to warping). Esophagus – The part of the digestive tract between the buccal mass and the stomach (Fig. 19).
Pelagic–(1) Free swimming in the open ocean; (2) the area of the ocean away from the ocean floor. Penis – The long, muscular end of the male gonoduct that serves to transfer spermatophores to the female (Fig. 29).
Key to Recent Cephalopod Groups and Families
Four longitudinal rows (series) of suction cups on manus of tentacular clubs; fins united at posterior end of mantle; medial posterior margin of fins concave (Fig. 52). 13a. Tentacles and clubs absent in adults, although present in larvae or occasionally in juveniles (Taningia), but, when present, always with rudimentary clubs armed with few teats (Fig. 56). The homogeneity of this classification becomes apparent when this type of locking cartilage is compared with the more highly specialized types (Fig. 54).
17a. Hooks present on tentacular rods (Fig. 61a and b); tentacles and clubs are lost in mature females. Family Pholidoteuthidae (Fig. 62) 19b. Tentacles lost in adults; tentacles in juveniles are small and weak, with some (about 6) weak. 30a. Cartilage oval enclosing the funnel with 1 or 2 knobs directed towards the center of the concavity (Fig. 54d).
33a. Six rows of longitudinal suckers on wings I to III, 4 longitudinal rows of suckers on wings IV; extremely long tail (greater than the length of the mantle), like a rubber-like extension of the gladius; no tail feathers (Fig. 78). Family Opisthoteuthidae (Fig. 84) 37b. Intermediate contractile membrane ('secondary web') present between each arm and the primary.
CHAMBERED NAUTILUSES
Decapod crustaceans appear to be a standard food for adults, but echinoid fragments, fish bones, coleoid beaks and nautilus tentacles have also been observed in the tumors of examined animals. Interest to fisheries: At least 2 of these species are of commercial value as food, largely at the artisanal and subsistence level, and nautilus shells are sold commercially in the shell trade (e.g. in Indonesia, Fiji, New Caledonia and the Philippines) . Recently, Ward and Saunders (1997) erected a new genus, Allonautilus, for the most unusual member of the group, A.
Diagnostic features: Navel open, a deep, rounded shoulder concavity, approximately 15 to 16% of the shell diameter at its widest point; inner coils of the shell visible. Habitat and Biology: Nautilus macromphalus inhabits continental shelf and slope waters associated with coral reefs, from the surface to a depth of approximately 500 meters. Collected alive at a depth of approximately 65 m on the outer slope of the barrier reef in New Caledonia; in the Coral Sea it is caught by traps at depths between 300 and 400 meters.
Shell color patterns are variable: irregular brown to reddish brown stripes radiate from the umbilicus to the vent in the usual color, but this stripe may be reduced to varying degrees, leaving the umbilicus and even most of the white wings. Size: Shell diameters are typically between 170 and 180 mm around Fiji and the Philippines, larger in Western Australian populations (ie mean diameter up to 222 mm). The outer layers of the shells are sometimes removed, leaving the outer surface a silver layer of mother of pearl.
Remarks: The Western Australian population includes the largest known specimens with an average shell diameter of 222 mm and an average weight of 1 675 g. Nautilus repertus Iredale, 1944, Australian pearl-coloured Nautilus. Australian zoological type locality: Rottnest Island and Pelsart Island, Western Australia]. Remarks: Very similar in size and weight to Nautilus pompilius, from which it can be distinguished by the absence of callus and the reduced coloration lacking in the umbilical region.
Nautilus scrobiculatusLightfoot, 1786, a catalog of the Portland Museum, latterly owned by the Dowager Duchess of Portland, deceased. Apart from the larger navel, Allonautilus scrobiculatus differs from the other nautiluses in the very special features of the shell, including folds and crusting of the periostracum (see Figure 97). Nautilus perforatus Conrad, 1847, Notes on shells, with description of new genera and species. Journal of the Academy of Natural Sciences, Philadelphia, Second Series type locality: unknown].
CUTTLEFISHES
LIST OF NOMINAL SPECIES
LIST OF SPECIES BY MAJOR FISHING AREAS
INDEX OF SCIENTIFIC AND VERNACULAR NAMES
LIST OF COLOUR PLATES
