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Teuthowenia pellucida Chun, Cycloteuthis Joubin, Cycloteuthis sirventi Joubin, Cycloteuthis akimushkini Filippova, Discoteuthis Young and Roper, Discoteuthis discus Young and Roper, Discoteuthis laciniosa Young and Roper, Enoploteuthis dOrbigny in Rppell, Enoploteuthis leptura leptura Leach, Enoploteuthis leptura magnoceani Nesis, Enoploteuthis anapsis Roper, Enoploteuthis chunii Ishikawa, Enoploteuthis galaxias Berry, Enoploteuthis higginsi Burgess, Enoploteuthis jonesi Burgess, Enoploteuthis obliqua Burgess, Enoploteuthis octolineata Burgess, Enoploteuthis reticulata Rancurel, Enoploteuthis semilineata Alexeyev, Abralia Gray, Abralia armata Quoy and Gaimard, Abralia andamanica Goodrich, Abralia astrolineata Berry, Abralia astrostricta Berry, Abralia dubia Adam, Abralia fasciolata Tsuchiya, Abralia grimpei Voss, Abralia heminuchalis Burgess, Abralia marisarabica Okutani, Abralia multihamata Sasaki, Abralia omiae Hidaka and Kubodera, Abralia redfieldi Voss, Abralia renschi Grimpe, Abralia robsoni Grimpe, Abralia siedleckyi Lipinski, Abralia similis Okutani and Tsuchiya, Abralia spaercki Grimpe, Abralia steindachneri Weindl, Abralia trigonura Berry, Abralia veranyi Rppel, Abraliopsis Joubin, Abraliopsis hoylei Pfeffer, Abraliopsis affinis Pfeffer, Abraliopsis atlantica Nesis, Abraliopsis chuni Nesis, Abraliopsis falco Young, Abraliopsis felis McGowan and Okutani, Abraliopsis gilchristi Robson, Abraliopsis lineata Goodrich, Abraliopsis morisii Verany, Abraliopsis pacificus Tsuchiya and Okutani, Abraliopsis tui Riddell, Watasenia Ishikawa, Watasenia scintillans Berry, Key to the genera of Gonatidae.

Gonatus Gray, Gonatus fabricii Lichtenstein, Gonatus kamtschaticus Middendorff, Gonatus madokai Kubodera and Okutani, Gonatus steenstrupi Kristensen, Berryteuthis Naef, Berryteuthis magister Berry, Berryteuthis anonychus Pearcy and Voss, Gonatopsis Sasaki, Gonatopsis octopedatus Sasaki, Gonatopsis borealis Sasaki, Gonatopsis japonicus Okiyama, Gonatopsis makko Okutani and Nemoto, Gonatus antarcticus Lnnberg, Gonatus berryi Naef, Gonatus californiensis Young, Gonatus onyx Young, Gonatus oregonensis Jefferts, Gonatus pyros Young, Gonatus ursabrunae Jefferts, Berryteuthis magister nipponensis Okutani and Kubodera, Berryteuthis magister shevtsovi Katugin, Eogonatus Nesis, Eogonatus tinro Nesis, Histioteuthis dOrbigny, Histioteuthis bonnellii Ferussac, Histioteuthis hoylei Goodrich, Histioteuthis miranda Berry, Histioteuthis reversa Verrill, Histioteuthis arcturi Robson, Histioteuthis atlantica Hoyle, Histioteuthis celetaria celetaria Voss, Histioteuthis celetaria pacifica Voss, Histioteuthis corona berryi Voss, Histioteuthis corona cerasina Nesis, Histioteuthis corona corona Voss and Voss, Histioteuthis corona inermis Taki, Histioteuthis eltaninae Voss, Histioteuthis heteropsis Berry, Histioteuthis macrohista Voss, Histioteuthis meleagroteuthis Chun, Histioteuthis oceani Robson, Lycoteuthis Pfeffer, Lycoteuthis lorigera Steenstrup, Lycoteuthis springeri Voss, Nematolampas Berry, Nematolampas regalis Berry, Nematolampas venezuelensis Arocha, Selenoteuthis Voss, Selenoteuthis scintillans Voss, Lampadioteuthis megaleia Berry, Magnapinna Vecchione and Young, Magnapinna pacifica Vecchione and Young, Magnapinna atlantica Vecchione and Young, Magnapinna talismani Fischer and Joubin, Mastigoteuthis Verrill, Mastigoteuthis agassizii Verrill, Mastigoteuthis atlantica Joubin, Mastigoteuthis cordiformis Chun, Mastigoteuthis danae Joubin, Mastigoteuthis dentata Hoyle, Mastigoteuthis famelica Berry, Mastigoteuthis glaukopis Chun, Mastigoteuthis hjorti Chun, Mastigoteuthis magna Joubin, Mastigoteuthis microlucens Young, Lindgren and Vecchione Mastigoteuthis psychrophila Nesis, Mastigoteuthis pyrodes Young, Mastigoteuthis hastula Berry, Mastigoteuthis inermis Rancurel, Mastigoteuthis iselini MacDonald and Clench, Mastigoteuthis latipinna Sasaki, Mastigoteuthis okutanii Salcedo-Vargas, Mastigoteuthis tyroi Salcedo-Vargas, Neoteuthis Naef, Neoteuthis thielei Naef, Alluroteuthis Odhner, Alluroteuthis antarcticus Odhner, Narrowteuthis Young and Vecchione, Nototeuthis dimegacotyle Nesis and Nikitina, Octopoteuthis sicula Rppell, Taningia Joubin, Taningia danae Joubin, Octopoteuthis danae Joubin, Octopoteuthis deletron Young, Octopoteuthis indica Naef, Octopoteuthis megaptera Verrill, Octopoteuthis nielseni Robson, Octopoteuthis rugosa Clarke, Illex illecebrosus Lesueur, Illex argentinus Castellanos, Illex coindetii Verany, Illex oxygonius Roper, Lu and Mangold, Ommastrephes dOrbigny, in Ommastrephes bartramii Lesueur, Dosidicus Steenstrup, Dosidicus gigas dOrbigny Eucleoteuthis Berry, Eucleoteuthis luminosa Sasaki, Hyaloteuthis Gray, Hyaloteuthis pelagica Bosc, Ornithoteuthis Okada, Key to the species of Ornithoteuthis.

Ornithoteuthis volatilis Sasaki, Ornithoteuthis antillarum Adam, Sthenoteuthis Verrill, Key to the species of Sthenoteuthis. Sthenoteuthis oulaniensis Lesson, Sthenoteuthis pteropus Steenstrup, Todarodes Steenstrup, Todarodes sagittatus Lamarck, Todarodes angolensis Adam, Todarodes filippovae Adam, Todarodes pacificus Steenstrup, Todarodes pusillus Dunning, Martialia Rochebrune and Mabille, Martialia hyadesi Rochebrune and Mabille, Nototodarus Pfeffer, Nototodarus sloanii Gray, Nototodarus gouldi McCoy, Nototodarus hawaiiensis Berry, Todaropsis Girard, Todaropsis eblanae Ball, Onychoteuthis Lichtenstein, Onychoteuthis banksii Leach, Onychoteuthis borealijaponica Okada, Ancistroteuthis Gray, Ancistroteuthis lichtensteini Frussac, Notonykia Nesis, Roeleveld and Nikitina, Notonykia africanae Nesis, Roeleveld and Nikitina, Notonykia nesisi Bolstad, Onykia Lesueur, Onykia carriboea Lesueur, Onykia ingens Smith, Onykia knipovitchi Filippova, Onykia lnnbergi Ishikawa and Wakiya, Onykia robsoni Adam, Onykia robusta Verrill, Kondakovia Filippova, Kondakovia longimana Filippova, Walvisteuthis Nesis and Nikitina, Walvisteuthis virilis Nesis and Nikitina, Onykia appellfi Pfeffer, Onykia intermedia Pfeffer, Onykia platyptera dOrbigny, Onykia verrilli Pfeffer, Promachoteuthis Hoyle, Promachoteuthis megaptera Hoyle, Promachoteuthis sp.

B Young, Vecchione and Roper D Young, Vecchione and Roper Pyroteuthis Hoyle, Pyroteuthis margaritifera Rppel, Pyroteuthis addolux Young, Pyroteuthis serrata Riddell, Pterygioteuthis Fischer, Pterygioteuthis giardi Fischer, Pterygioteuthis gemmata Chun, Pterygioteuthis microlampas Berry, Cephalopod catches have increased steadily in the last 40 years, from about 1 million metric tonnes in to more than 4 million metric tonnes in FAO, This increase confirms a potential development of the fishery predicted by G.

Voss in , in his first general review of the worlds cephalopod resources prepared for FAO. The rapid expansion of cephalopod fisheries in the decade or so following the publication of Vosss review, meant that a more comprehensive and updated compilation was required, particularly for cephalopod fishery biologists, zoologists and students.

Roper, M. Sweeney and C. Nauen was published in to meet this need. For example, this work should be useful for the ever-expanding search for development and utilization of natural products, pharmaceuticals, etc. The catalogue is based primarily on information available in published literature. However, yet-to-be-published reports and working documents also have been used when appropriate, especially from geographical areas where a large body of published information and data are lacking.

We are particularly grateful to colleagues worldwide who have supplied us with fisheries information, as well as bibliographies of local cephalopod literature. This information is supplemented by field observations made by the authors in many parts of the world, both in preparation of the volume, as well as for the current edition. These field visits provided opportunities to examine fresh material at landing sites, markets and laboratories, as well as to obtain first-hand information about local cephalopod fisheries from regional fisheries workers.

Additional examinations of preserved specimens occurred in museums. During the plus years separating the two editions, the rapid development of cephalopod fisheries worldwide and the simultaneous increase in the population of fisheries scientists, their research and publications, made available an enormous amount of new data and research results.

Sometimes it is difficult to evaluate the reliability of published data, especially with regard to the identification of species in areas where the cephalopod fauna has not been sufficiently studied taxonomically. Moreover, the discovery of new species, the more accurate delimitation of known species, or even the introduction of nomenclatural changes, may cause confusion and lead to the use of scientific names that are incorrect by modern standards.

Although great care was exercised to evaluate and correct such published information used in the preparation of this catalogue, some incorrect interpretations may have occurred. Another potential limitation, in the taxonomic literature especially, is that information on the economic importance of species is rather scarce or of a very general nature.

Also, important information may have been overlooked if published only in local fisheries literature that is unavailable on an international scale. All of these potential limitations, however, have been significantly mitigated during the preparation of the new edition because of the availability of on-line fisheries databases and bibliographic search capabilities.

With regard to the limitations mentioned above, we heartily request that readers who detect any errors in the information presented, or who have additional information and data that will enhance the accuracy and utility of this book, please contact and inform one of the authors or the Species Identification and Data Programme SIDP of the Marine Resources Service, Fisheries Resources Division, Fisheries Department, FAO Rome.

The number of cephalopod species that enter commercial fisheries has continued to grow significantly since , as a result of a still-growing market demand and the expansion of fisheries operations to new fishing areas and to deeper waters. It has been suggested that the cephalopod life-strategy may guarantee survival against environmentally stressful conditions, including those caused by heavy fishing. However, as cephalopod fisheries experienced further extensive development, parallel concern developed regarding potential overexploitation.

Thus, a broad consensus emerged among fishery biologists to apply the experience gained from errors made in finfish management to avoid possible failures in cephalopod exploitation. To help prevent potential failures, refined species identification capabilities are required, as well as a more detailed and accurate compilation of information on cephalopod species, distribution, biology, fisheries and catch statistics.

Consequently, FAO recognized that a new edition of the Cephalopods of the World catalogue was required. To achieve this expanded goal, several authors with particular areas of specialization were assembled to enhance the accuracy, coverage and utility of this revised catalogue.

In our attempt to make this document as comprehensive and as useful as possible, the taxonomic coverage of this edition of the catalogue is organized into 3 levels of interest: Level 1: species of cephalopods currently exploited commercially and species utilized at the subsistence and artisanal levels; Level 2: species of occasional and fortuitous interest to fisheries; this includes species considered to have a potential value to fisheries, based on criteria such as edibility, presumed abundance, accessibility, marketability, etc.

Level 3: species with no current interest to fisheries, which are listed only with the basic information available. The inclusion of such a wide range of species is necessary to provide the most comprehensive inventory of species potentially useful to mankind, regardless of their current.

Species identifications should be attempted only after verification of the family through use of the illustrated key to families. Sizes or measurements might not be completely comparable because they were taken mostly from preserved or fixed specimens, but measurements of commercially important species often come from fresh material. Because of the elasticity of tentacles and arms, total length is not a very accurate measurement. Where both total length and mantle length are given, the respective figures do not necessarily pertain to the same specimen but may have been obtained from different sources.

The available information on the size attained by some species often is very meagre, so the maximum reported size cited here might be considerably smaller than the actual maximum size. Maximum weight is given when available. In cases where only scattered records of occurrence are available, question marks have been used to indicate areas of suspected or unconfirmed distribution. For the sake of exactness actual depth data are reported, as given in the referenced literature.

Information on biological aspects, such as migration, spawning season and area, longevity, prey, and predators, also is included. Due to the dominant role of squids in the marine environment, this section is especially detailed in this volume. Data on utilization fresh, dried, cooked, frozen, canned, etc. Here, too, the quality and quantity of the available information varies considerably among the species, and it is reported in as much detail as possible in relation to the squids significance to the fisheries.

The present compilation is necessarily incomplete, since only a fraction of the local names applied to specific entities actually is published. In many cases, local names are available only for species that support traditional fisheries.

Apart from possible omissions due to limitations of literature available, some of the names included may be somewhat artificial, i. 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.

The type genus within each family is treated first, then all remaining genera are listed alphabetically. The type species within each genus is treated first, then all species are listed alphabetically. Level 1 includes the most important species for fisheries utilization, and it consists of detailed information in all 12 categories listed below. Level 2, which comprises those species of occasional or potential interest to fisheries, consists of whatever information is available and appropriate for the 12 categories.

Level 3, those species for which there is no current direct or indirect interest to fisheries, consists of basic information i. The format within the species sections includes the first two levels of treatment Level 1 and Level 2 presented together. Species included in Level 3 are presented at the end of each family.

Consequently, each major group and family is introduced with general descriptive remarks, illustrations of diagnostic features, highlights of the biology and relevance to fisheries. The information that pertains to each species in Levels 1 and 2 is arranged by categories as follows: 1 scientific name; 2 synonymy; 3 misidentifications; 4 FAO names; 5 diagnostic features with illustrations; 6 maximum known size; 7 geographical distribution with map; 8 habitat and biology; 9 interest to fisheries; 10 local names; 11 remarks 12 literature.

Wherever possible, these names are selected based on vernacular names or parts of names already in existence within the areas where the species is fished. FAO species names, of course, are not intended to replace local species names, but they are considered necessary to. Cephalopods of the World 11 Remarks: Important information concerning the species, but not specifically linked to any of the previous categories, is given here.

For example, in some cases the taxonomic status of certain scientific names requires further discussion. Other nomenclatural problems are discussed in this section, such as the use of subspecies names. Additional references are included in the bibliography. In the case of a few uncommon species, only systematic papers are available. The massive amount of literature relevant to fisheries for many species of squids required that appendices be compiled for this Volume. The appendix includes a list of publications useful to gain an understanding of the species biology, ecology and fisheries.

Publications are listed by authors name, date of publication and key words for the publications contents. For practical purposes we separate the cephalopods into several groups, without assigning or implying taxonomic relationships. Figure 1 diagrams several of the classification schemes currently under discussion.

Cephalopods include exclusively marine animals that live in all oceans of the world with the exception of the Black Sea, from the Arctic Sea to the Antarctic Ocean and from the surface waters down into the deep sea. Cephalopods first appeared as a separate molluscan taxonomic entity, the nautiloids, in the Upper Cambrian period over million years ago , but more than half of these ancestors were already extinct by the end of the Silurian, million years ago, when only the nautiluses survived.

Meanwhile, other forms arose in the late Palaeozoic between and million years ago , including those of the Subclass Coleoidea, but most of them became extinct by the end of the Mesozoic, about million years ago. The only members of the subclass Coleoidea that exist today are the forms that developed in the Upper Triassic and Lower Jurassic between and million years ago. Although there is a long fossil record of many different groups, all living cephalopods belong to two subclasses: the Coleoidea, which includes the major groups known as squids, cuttlefishes sensu lato, octopods and vampires, and the Nautiloidea, containing two genera, Nautilus and Allonautilus, the only surviving cephalopods with an external shell.

At the present time the status and understanding of the Systematics and Classification of the Recent Cephalopoda is under considerable discussion. The families of living cephalopods are, for the most part, well resolved and relatively well accepted. Species-level taxa usually can be placed in well-defined families. The higher classification, however, still is not resolved.

The classification above the family level is controversial and a broad consensus still needs to be achieved. This situation is not unexpected for a group of organisms that has undergone explosive research attention in recent decades. These terms are also used to indicate the major groups. The term cuttlefishes also is used sensu lato to indicate the following groups: Cuttlefishes, Bobtail squids, Bottletail squids, Pygmy squids and the Rams horn squid.

Cuttlefishes, along with Nautiluses were treated in Volume 1 Jereb and Roper, Octopods will be treated in Volume 3. This second volume of the Catalogue is focused on Squids. Salinity is considered to be a limiting factor in squids distribution; they are generally restricted to salinity concentr ations bet ween 27 and However, Lolliguncula brevis, which lives and reproduces in waters of 17, demonstrates a capacity for a higher degree of salinity tolerance Hendrix et al. Some species.

The habitat depth range extends from the intertidal to over 5 m. Many species of oceanic squids undergo diel vertical migrations: they occur at depths of about to m during the day, then at the onset of twilight and increasing darkness, they ascend into the uppermost m for the night. A deeper-living layer of diel migrators occurs from about 1 m to m during the daytime.

The abundance of squids varies, depending on genera, habitat and season, from isolated individuals, small schools with a few dozen individuals, to huge schools of neritic and oceanic species with millions of specimens.

General charcteristics The size of adult squids ranges from less than 10 mm mantle length e. The largest specimens may weigh over kg, but the average size of commercial species is to mm mantle length and about 0. Squids are soft-bodied, bilaterally symmetrical animals with a well-developed head and a body that consists of the muscular mantle, the mantle cavity that houses the internal organs, and the external fins.

The head bears an anterior circum-oral surrounding the mouth crown of mobile appendages arms, tentacles. The mouth, at the interior base of the arm crown, has a pair of chitinous jaws the beaks and, as in other molluscs, a chitinous tongue-like radula band of teeth.

The ancestral mollusc shell is reduced to a rigid structure composed of chitin, the gladius or pen, sometimes quite thin and flexible. The loss of the external shell allowed the development of a powerful muscular mantle that became the main locomotory organ for fast swimming, via water jettisoned from the funnel. The funnel also known as siphon, an archaic term correctly applied to some other molluscs, but not to modern, extant cephalopods is a unique, multifunctional, muscular structure that aids in respiration and expulsion of materials, in addition to locomotion.

Oxygenated water is drawn through the mantle opening around the head neck into the mantle cavity, where it bathes the gills for respiration. Muscular mantle contraction expels the deoxygenated water from the mantle cavity through the ventrally located funnel. The discharge jet. Female reproductive products eggs, egg masses also are discharged through the funnel. Squids produce ink, a dark, viscous fluid also expelled through the funnel.

The ink may take the form of a mucoidal pseudomorph false body to decoy potential predators, or of a cloud to obscure the escaping cephalopod. One pair of gills ctenidia is present, for respiration, i. Squids may use anerobic muscle layers, and cutaneous respiration also occurs. The circulatory system The circulatory system is distinctive within the Mollusca. It is a closed system blood contained within vessels , similar in many respects to that of vertebrates, that fulfills the demand for the more efficient circulation required by an active locomotory system.

The system is composed of a principal, or systemic, heart, two branchial hearts and developed arterial, venous and capillary systems that supply blood to the muscles and organs. The oxygenated blood passes from the gills through the efferent branchial vessels to the systemic heart, where it is expelled from the ventricle through three aortas: the cephalic or dorsal aorta, which supplies the head and the anterior part of the gut; the posterior, minor or abdominal aorta that supplies the mantle and fins along with the posterior part of the gut and the funnel; and the gonadal aorta that develops gradually with sexual maturation of the animal.

The blood is collected through sinuses and capillaries into the veins, through which it passes to the branchial hearts that pump it through the filaments of the gills. The circulating respiratory pigment used for oxygen transport is copper-containing haemocyanin, a system of rather lower efficiency than the iron-containing haemoglobin of vertebrates.

Blood sinuses in living squids are much reduced and replaced functionally by muscles. The circulatory system therefore has to work against the peripheral muscle-induced pressure, which increases with increasing activity maximum during jet-swimming. It also has to cope with the resistance of the small diameter of the final capillary blood vessels, and the low oxygen carrying capacity of the blood less than 4. In spite of these limitations, the system has other functional modifications see for example Wells and Smith, ; Martin and Voight, that achieve the capacity to deliver oxygen at a rate comparable to that of active fishes, enabling squids to accomplish extraordinary swimming, attack and escape performances.

The excretory system The excretory system also differs markedly from that of other molluscs and, along with the closed circulatory system and the branchial circulation, enables unique relationships between blood and the final secretion, the urine. The excretory system consists basically of the renal sac with the renal appendages organs comparable to vertebrate kidneys , the pericardial glands, the branchial hearts and the gills.

Squids are ammoniotelic, whereby ammonium ions are continuously released by the gill epithelium and by renal appendages into the surrounding water. Ammonium ions are used by buoyant squids to replace denser chloride ions in fluids in the coelom and in the body tissues. Because this solution is less dense and hence more buoyant than seawater, it provides lift for neutral or positive buoyancy. Cephalopods of the World The nervous system The nervous system is highly developed, with a large brain and peripheral connections, contrasting with the original molluscan circumesophageal nerve ring.

Among its most remarkable features is the giant fibre system that connects the central nervous system with the mantle muscles. This system consists of three orders of cells and fibres and ensures the immediate and simultaneous contraction of mantle, fins and retractor muscles of both sides, rather than an anterior to posterior sequential contraction that would be counter-productive for water movement expulsion.

Also remarkable is the eye development of squids, for which vision plays a major role in life. Their eyes are large, have a design generally similar to that of fishes and other vertebrates e. This is extremely important for hunters that rely on sight, and it is accomplished by connections of the eye muscles with the statocysts, a bilateral mechanism similar to the vestibulo-optic system of fishes.

The statocyst system provides squids with information on their orientation, as well as changes in position and direction of movement. It is a highly developed system that consists of two separate cavities located bilaterally in the cartilaginous skull, posteroventral to the brain. The statocysts contain nervous cells and receptors differentiated to detect both linear acceleration, with the aid of calcareous stones called statoliths, and angular acceleration. Some squids also have extra-ocular photoreceptors photosensitive vesicles about which little is known; in mesopelagic squids they appear to monitor light intensity in order to enable the animals to match their counter-illumination with the ambient light with their own photophores light-producing organs.

Squids are provided with numerous mechano- and chemoreceptors and recent evidence indicates that in some species, e. Loligo vulgaris, ciliate cells form lines in several parts of the body, a system analogous to the lateral-line system in fishes. Squids are able to change colour by using a complex system of chromatophores under nervous control. The chromatophores are pigment-filled sacs present in the skin, and capable of remarkable expansion and contraction. This system responds virtually instantaneously to contemporary situations in the environment, and it is critical for survival.

Squid species also have iridocytes shiny, reflective platelets in the skin. Squids behaviour includes rapid changes in overall colour and colour pattern and many deep-sea forms camouflage themselves by producing bioluminescent light from photophores which eliminate their silhouettes against the down-welling sunlight in the dimly-lit mid-depths. This capability enables squids to inhabit open water, even in the great depths in the ocean, the greatest volume of living space on earth.

Feeding Squids are voracious, active predators that feed upon crustaceans, fishes and other cephalopods. A common hunting technique involves extremely rapid shooting forward of the tentacles to capture the prey, while in some oegopsid squids the tentacles may be used like long, sucker-covered fishing lures. The captured prey is brought to the mouth and killed by bites of the strong, chitinous beaks, equipped with powerful muscles.

Reproduction Squids are dioecious separate sexes and many species, though not all, exhibit external sexual dimorphism, either in morphological or morphometric differences. Females frequently are larger than males and males of most species possess one, occasionally two, modified arm s the hectocotylus for transferring spermatophores to females during mating.

The males of some species also exhibit modifications to other arms, in addition to the hectocotylus. The hectocotylus may be simple or complex and can consist of modified suckers, papillae, membranes, ridges and grooves, flaps. The one or two nuptual limbs function to transfer the spermatophores tubular sperm packets from the males reproductive tract to an implantation site on the female. The spermatophores may be implanted inside the mantle cavity where they may penetrate the ovary , into the oviducts themselves, around the mantle opening on the neck, on the head, in a pocket under the eye, around the mouth or in other locations.

Females of a few species also develop gender-s pecific struc tures e. Mating often is preceded or accompanied by courtship behaviour that involves striking chromatophore patterns and display. Copulatory behaviour varies significantly among species, in colour and textural display, proximity of male and female, duration of display and spermatophore transfer, and the location of implantation of the spermatophores on the female.

The gonads form a single mass at the posterior end of the mantle cavity, and female gonoducts may be paired in oegopsids or single, as in other squids. The reproductive systems are highly complex structures with ducts, glands and storage organs. Female squids have nidamental glands and loliginids have accessory nidamental glands, as well. Spermatophores are produced in the multi-unit spermatophoric gland and stored in the Needhams sac, from which they are released through the terminal part of the duct, the penis.

This term is not strictly accurate,. Locomotion Locomotion is achieved by a combination of jet propulsion and flapping or undulating the fins on the mantle. The fins on the mantle also provide balance and steering during jet propulsion. The number and size of spermatophores vary greatly, depending on the species and group for reviews on spermatophore structures and function see Mann et al.

Once in contact with seawater, the so called sper matophori c react ion begins. T he spermatophores evert, with the resultant extrusion of the sperm packet caused by the penetration of water inside the spermatophoric cavity, where the osmotic pressure is higher. The resulting extruded sperm packet is named spermatangium or sperm bulb or body. Sperm are able to survive several months once stored in the female, at least in some species, and fertilization of mature ova may take place either in the ovary, the mantle cavity or the arm cone formed by the outstretched arms while the eggs are laid.

Fertilized eggs are embedded in one or more layers of protective coatings produced by the nidamental glands and generally are laid as egg masses. Egg masses may be benthic or pelagic. Eggs of neritic, inshore squids, except in Sepioteuthis, generally are very small only a few millimetres in diameter and frequently are laid in finger-like pods each containing from a few to several hundred eggs.

Deposited in multi-finger masses sometimes called sea mops , these eggs are attached to rocks, shells or other hard substrates on the bottom in shallow waters. Many oceanic squids lay their eggs into large sausage-shaped or spherical gelatinous masses containing tens or even hundreds of thousands of eggs that drift submerged in the open sea.

This has led to changes in our conceptions about the physiology and ecology of many species, but more research is required before a full understanding is achieved see Jereb et al. Principal results obtained from the research generally confirm a very high growth rate in squids, comparable to that of the fastest-growing fishes.

The life expectancy of most squids appears to range from a few months to one or two years, and many small oceanic squids, such as pyroteuthids may complete their life cycles in less than six months. Recent evidence, however, suggests that larger species of squids, for example the giant squid Architeuthis spp. All squids die after their spawning period. Systematics status The total number of living species of squids that currently are recognized is more than ; are listed in the present volume.

The status of the systematics of squids has changed in the last 30 years, as research and associated scientific discussions have increased substantially. However, phylogenetic relationships among many families remain uncertain, and new species are described fairly frequently as new habitats are explored and as families are gradually better-understood.

Growth and life history Development of squid embryos is direct, without true metamorphic stages. However, hatchlings undergo gradual changes in proportions during development and the young of some species differ from the adults. Thus, the term paralarva has been introduced for these early stages of cephalopods that differ morphologically and ecologically from older stages.

The paralarvae of many deep-sea species of squids occur in the upper m of the open ocean; then they exhibit an ontogenetic descent, gradually descending to deeper depths with increasing size until the adult depth is attained. Time of embryonic development varies widely, from a few days to many months, depending on the species and the temperature conditions. Hatching may occur synchronously from a single clutch or be extended over a period of 2 or 3 weeks.

In spite of the large number of studies and research carried out on squids, especially in recent decades, the life history of many species still is unknown, and our knowledge of the life cycles of the members of this interesting group remains fragmentary. Information comes from studies in the field as well as from observations in the laboratory. However, little is known of life history for species that are not targets of regular fisheries, and only a few squid species have been reared successfully in the laboratory.

Studies and monitoring of growth are complicated by the high variability in individual growth rates. This makes it difficult to apply conventional methods, e. Determination of age also is difficult, because squids have few hard structures that show daily marks rings that enable direct estimates of age.

In the last Conclusions Squids are important experimental animals in biomedical research with direct applications to human physiology and neurology, for example. Because of their highly developed brains and sensory organs, they are valuable in behavioural and comparative neuro-anatomical studies.

In addition, the extremely large single nerve axons of some squids, the largest in the animal kingdom, are used extensively in neuro-physiological research. The bite of squid can be painful at the least to humans, or secondarily infected, or, rarely, lethal. A documented threat by squids to humans is from the large ommastrephid squid, Dosidicus gigas, which forms large aggressive schools that are known to have attacked fishermen that have fallen in the water, causing several confirmed deaths.

Scuba divers also have been attacked. Therefore, squids must be handled carefully. Of the total cephalopod catch of over 4 million tonnes reported for by FAO statistics FAO, , over 3 million tonnes were squids, i. The impressive increase in squid production during the last 25 years is due. Cephalopods of the World mainly to the discovery and increasing exploitation of squid resources in the southwest Atlantic, principally for Illex argentinus, as well as an increase in the production of other major squid target species, mainly Todarodes pacificus in the northwest Pacific and Dosidicus gigas in the eastern Pacific.

Illex argentinus catches exceeded 1 million tonnes in , a record peak which placed this species at the eleventh position in value of the total world marine-species production for that year. Numerous fishing techniques and methods to capture squids have been developed over time. These were extensively reviewed, for example, by Rathjen , [] and Roper and Rathjen They include lures, jigs, lampara nets, midwater trawls and otter trawls. Jigging is the most widely used method, which accounts for almost half of the world squid catch, primarily ommastrephids, but also a few loliginids.

This technique is employed primarily at night, when many species of squids are attracted to the fishing vessel by lights. Figure 3 shows the distribution of the worlds light fishery for some of the most important squid species. Jigs, which feature numerous, variously-arranged, barbless hooks Plate I, 6 , are lowered and retrieved by jigging machines that simulate the constant swimming behaviour of natural prey, inducing.

While simple hand-jigging machines are still used in small-scale, artisanal fisheries, large modern vessels for industrial fishing activities are equipped with scores of automated, computer-controlled jigging machines, each capable of catching several tonnes per night Plate I, 1 and 5. Trawling is the secondmost productive fishery method to catch squids Plate I, 2. Formerly, almost all squids were caught as bycatch in trawl fisheries for finfishes and shrimps.

Trawling is a very efficient technique to catch species, but soft-bodied animals like cephalopods are often damaged by the other species in the catch, particularly in benthic and epibenthic otter trawls. Even in fisheries in which squid-specific trawling occurs, the huge catches of squids per tow often result in crushed and damaged product.

Consequently, trawled squid product generally is less valuable than jig-caught squids. However, modern oceanic trawlers can process on board many metric tonnes of cephalopods per day, which helps insure a high-quality product. Bottom trawling can be very dangerous for benthic habitats because of the physical damage it causes to the seabed and associated fauna and because of its lack of selectivity.

Consequently, less intense exploitation by this traditional fishing technique and an approach toward diversification of methods and redistribution of the fisheries through different areas were encouraged and still are highly recommended, especially in situations where small-scale fisheries still exist and new, more efficient methods can be implemented.

Nearshore, neritic squids frequently are caught by purse seines, lift nets, beach seines, etc. Products range from fresh food, eaten raw as sashimi in Japan and, in recent years, worldwide, and fresh-cooked, as well as various types of processed product dried, canned, frozen, reduced to meal, etc. The high protein and low fat content of cephalopods make them an important and healthy element in the human diet. Considering the present level of exploitation of the commercially-fished squid populations, a further increase in such fishery production is likely to occur, first by expansion of the fisheries into the less-fished regions of the oceans, e.

There, a standing stock of squid biomass as high as million tonnes was estimated by scientists, based on an estimate of 30 million tonnes consumed by vertebrate predators see Rodhouse et al. Therefore, a priority for the future research in the field of Antarctic cephalopod biology will be to assess the squid biomass there, quantitatively and qualitatively, with the objective of determining and developing a sustainable fishery production.

However, polar squids probably are longer living and slower growing than species currently harvested. Therefore, caution must be exercised in assumptions and decisions for management of polar squid fisheries. In the future, it is likely that attention will be focused on finding other species and families to replace fish stocks that become severely reduced by overfishing.

Even though clear evidence reveals the existence of large cephalopod resources available for exploitation in the open oceans, based on the estimated consumption by predators see Clarke, b; Piatkowski et al. Research is being carried out on how to remove this factor on a commercial scale, but results will take time and catches will need to be processed before marketing and utilization.

A number of ommastrephid squids that lack ammonium are considered to be underexploited. T hese include: Sthenoteuthis pteropus, Ommastrephes bartramii, Martialia hyadesi, Todarodes sagittatus, Sthenoteuthis oualaniensis, Nototodarus philippinensis, Dosidicus gigas, and the circumpolar, sub-Antarctic Todarodes filippovae.

Exploitation of these species would provide large tonnages of high quality cephalopods and would require only minor development in catching techniques. However, it will be necessary to determine where these species congregate for feeding and spawning activities.

An analysis of biomass, production and potential catch for the Ommastrephidae species is presented in Nigmatullin Although a number of other oceanic squid families have large populations and high quality flesh, they are not currently exploited on a commercial scale except for a few seasonal fisheries.

These include members of the families Thysanoteuthidae, Gonatidae and Pholidoteuthidae, for example. Increased exploitation of these groups, however, would also require some research and development of catching techniques. Commercial exploitation of the cosmopolitan family Histioteuthidae also could be considered, since at least one large commercial-level catch has been made in the North Atlantic see Okutani, personal communication, in Clarke, a.

However, the increased exploitation of these oceanic squid species might have unpredictable, far-reaching negative effects on the. Therefore, great caution must be exercised in developing this kind of fishery. Almost all of our knowledge of the general biology of cephalopods, in fact, is limited to the shelf-living species, as well as to those ommastrephids that move onto the shelf at certain seasons.

Even so, many gaps still exist in our knowledge about their life cycles, especially as far as the relationships among species are concerned e. Some populations of harvested species have shown sudden, occasionally catastrophic, declines before adequate biological data could be gathered and analysed. Squid stocks experienced true collapses at least in two well-known and documented cases. These were the northwest Pacific Todarodes pacificus fishery failure in the s and the northwest Atlantic Illex illecebrosus fishery collapse in the s.

While the T. These collapses are thought to have occurred mainly as a consequence of temporarily unfavourable environmental conditions or actual long-term environmental changes, probably aggravated by heavy fishing pressure Dawe and Warren, A significant challenge thus exists to deepen our knowledge and learn the details of distribution, life history and biology of exploited species in order to allow rational utilization of the stocks. The necessity for research as a key factor towards attaining this goal has been stressed by many authors e.

Lipinski et al. Therefore, squids are potentially very good indicator species to predict or reflect changes in environmental conditions, both locally and on a broader scale see Pierce et al. Perhaps even more significant is the challenge that exists for future exploitation of new species or populations. The role of squids in the ecosystem, in fact, is more complex than it was thought to be only a few decades ago.

Squids can be considered subdominant predators that tend to increase in biomass when other species, particularly their predators and competitors for food, become depleted, as a result of a combination of heavy or excessive fishing, other human impacts, oceanogr aphic fluc tuations and competition for food see Caddy, , and Caddy and Rodhouse, for a detailed analysis of the transition from finfish-targeted fisheries to cephalopod-targeted fisheries.

In turn, squids are major food items in the diets of innumerable species of fishes, toothed whales e. Muscular squids derive their energy from crustaceans, fishes and other cephalopods. At the same time, they are a very efficient food storage for the large, oceanic predators, by rapidly converting oceanic resources into high energy food.

On the other hand, neutrally buoyant ammoniacal squids, which probably greatly outnumber the muscular squids in biomass, also provide food to many of the same. Cephalopods of the World predators, but not over the continental shelf and with consistently lower energy per unit body mass. We know virtually nothing about the details of feeding, growth, life cycles, periodicities, distribution and spawning in ammoniacal species. In spite of our relatively incomplete knowledge, it is now clear that squids are a dominant component within marine ecosystems and that their abundance ultimately may influence the abundance of their predator and prey populations.

Studies of the effects of consumption of important pelagic squids and fishes by predatory fishes on the northeastern shelf of the United States Overholtz et al. Consistent with our present knowledge. Taking into consideration these factors, increasing effort should be focused on improving scientific knowledge of this group.

Squid catches need increased monitoring, especially in those areas of major environmental fluct uations and where fisher ies management is complicated by multiple countries exploiting the same resource. Cooperation, collaboration and commitment are required to better understand these important and fascinating animals. Abyssal The great depths of the ocean: from 2 to 6 m. Accessory nidamental glands Glands of unknown function; consist of tubules containing symbiotic bacteria.

Found in all decapodiformes except oegopsid squids. Adult A female that has mature eggs these frequently are stored in the oviducts , or a male that has produced spermatophores these are stored in Needhams sac. Afferent blood vessel Artery vessel carrying blood toward an organ.

Afferent nerve Nerve carrying impulses toward the brain or specific ganglia. Anal flaps A pair of fleshy papillae involved in directing releases of ink, 1 flap situated at each side of the anus Fig. Anterior Toward the head-end or toward the arm-tips of cephalopods. Anterior salivary glands Glands on or in the buccal mass that aid in preliminary digestion.

Anterior suboesophageal mass See Brachial lobe. Antitragus Knob that projects inward from the posterior surface of the central depression in the funnel-locking apparatus of some squids Fig. Anus Terminal opening of the digestive tract, in the anterior mantle cavity, sometimes extending to inside the funnel, through which digestive waste products, as well as ink, are expelled.

Apomorphic Derived from a more ancestral condition. Loosely considered the advanced condition. Arm One of the circumoral appendages of cephalopods. Arms are designated by the numbers I to IV, starting with I as the dorsal or upper pair. In squids each appendage of the fourth ancestral pair is modified to form a tentacle.

Arm formula Comparative length of the 4 pairs of arms expressed numerically in decreasing order: the largest arm is indicated first and the shortest last, e. Bathypelagic The deep midwater region of the ocean. Beak One of the 2 chitinous jaws of squids bound in powerful muscles. The dorsal beak is referred to as the upper beak and it inserts within the lower ventral beak to tear tissue with a scissors-like cutting action. Belemnoidea A fossil group of cephalopods that is thought to be the sister group of the Coleoidea.

Belemnoids are distinguished by the presence of hook-like structures on the arms rather than suckers. Benthopelagic A free-swimming animal that lives just above the ocean floor but rarely rests on the ocean floor. Bilateral symmetry The symmetry exhibited by an organism or an organ if only one plane can divide the animal structure into 2 halves that are mirror images of each other.

Granite is a visibly granular, plutonic rock, formed in a depth of some kilometers. Granites are fine- to coarse-grained, color is pink, red, grey or yellowish. The essentials are crystallized and visible with the naked eye.

The color is caused by the feldspars and part of the accessories. Essential s are: quartz, orthoclase, plagioclase and biotite. Accessorie s are: allanite, apatite, ilmenite, magnetite, pyrite, tourmaline, zircon. Accidentals are: garnet, hornblende, muscovite, pyroxene.

BITCOINS WALLET IPHONE 5C

Histioteuthis bonnellii Ferussac, Histioteuthis hoylei Goodrich, Histioteuthis miranda Berry, Histioteuthis reversa Verrill, Histioteuthis arcturi Robson, Histioteuthis atlantica Hoyle, Histioteuthis celetaria celetaria Voss, Histioteuthis celetaria pacifica Voss, Histioteuthis corona berryi Voss, Histioteuthis corona cerasina Nesis, Histioteuthis corona corona Voss and Voss, Histioteuthis corona inermis Taki, Histioteuthis eltaninae Voss, Histioteuthis heteropsis Berry, Histioteuthis macrohista Voss, Histioteuthis meleagroteuthis Chun, Histioteuthis oceani Robson, Lycoteuthis Pfeffer, Lycoteuthis lorigera Steenstrup, Lycoteuthis springeri Voss, Nematolampas Berry, Nematolampas regalis Berry, Nematolampas venezuelensis Arocha, Selenoteuthis Voss, Selenoteuthis scintillans Voss, Lampadioteuthis megaleia Berry, Magnapinna Vecchione and Young, Magnapinna pacifica Vecchione and Young, Magnapinna atlantica Vecchione and Young, Magnapinna talismani Fischer and Joubin, Mastigoteuthis Verrill, Mastigoteuthis agassizii Verrill, Mastigoteuthis atlantica Joubin, Mastigoteuthis cordiformis Chun, Mastigoteuthis danae Joubin, Mastigoteuthis dentata Hoyle, Mastigoteuthis famelica Berry, Mastigoteuthis glaukopis Chun, Mastigoteuthis hjorti Chun, Mastigoteuthis magna Joubin, Mastigoteuthis microlucens Young, Lindgren and Vecchione Mastigoteuthis psychrophila Nesis, Mastigoteuthis pyrodes Young, Mastigoteuthis hastula Berry, Mastigoteuthis inermis Rancurel, Mastigoteuthis iselini MacDonald and Clench, Mastigoteuthis latipinna Sasaki, Mastigoteuthis okutanii Salcedo-Vargas, Mastigoteuthis tyroi Salcedo-Vargas, Neoteuthis Naef, Neoteuthis thielei Naef, Alluroteuthis Odhner, Alluroteuthis antarcticus Odhner, Narrowteuthis Young and Vecchione, Nototeuthis dimegacotyle Nesis and Nikitina, Octopoteuthis sicula Rppell, Taningia Joubin, Taningia danae Joubin, Octopoteuthis danae Joubin, Octopoteuthis deletron Young, Octopoteuthis indica Naef, Octopoteuthis megaptera Verrill, Octopoteuthis nielseni Robson, Octopoteuthis rugosa Clarke, Illex illecebrosus Lesueur, Illex argentinus Castellanos, Illex coindetii Verany, Illex oxygonius Roper, Lu and Mangold, Ommastrephes dOrbigny, in Ommastrephes bartramii Lesueur, Dosidicus Steenstrup, Dosidicus gigas dOrbigny Eucleoteuthis Berry, Eucleoteuthis luminosa Sasaki, Hyaloteuthis Gray, Hyaloteuthis pelagica Bosc, Ornithoteuthis Okada, Key to the species of Ornithoteuthis.

Ornithoteuthis volatilis Sasaki, Ornithoteuthis antillarum Adam, Sthenoteuthis Verrill, Key to the species of Sthenoteuthis. Sthenoteuthis oulaniensis Lesson, Sthenoteuthis pteropus Steenstrup, Todarodes Steenstrup, Todarodes sagittatus Lamarck, Todarodes angolensis Adam, Todarodes filippovae Adam, Todarodes pacificus Steenstrup, Todarodes pusillus Dunning, Martialia Rochebrune and Mabille, Martialia hyadesi Rochebrune and Mabille, Nototodarus Pfeffer, Nototodarus sloanii Gray, Nototodarus gouldi McCoy, Nototodarus hawaiiensis Berry, Todaropsis Girard, Todaropsis eblanae Ball, Onychoteuthis Lichtenstein, Onychoteuthis banksii Leach, Onychoteuthis borealijaponica Okada, Ancistroteuthis Gray, Ancistroteuthis lichtensteini Frussac, Notonykia Nesis, Roeleveld and Nikitina, Notonykia africanae Nesis, Roeleveld and Nikitina, Notonykia nesisi Bolstad, Onykia Lesueur, Onykia carriboea Lesueur, Onykia ingens Smith, Onykia knipovitchi Filippova, Onykia lnnbergi Ishikawa and Wakiya, Onykia robsoni Adam, Onykia robusta Verrill, Kondakovia Filippova, Kondakovia longimana Filippova, Walvisteuthis Nesis and Nikitina, Walvisteuthis virilis Nesis and Nikitina, Onykia appellfi Pfeffer, Onykia intermedia Pfeffer, Onykia platyptera dOrbigny, Onykia verrilli Pfeffer, Promachoteuthis Hoyle, Promachoteuthis megaptera Hoyle, Promachoteuthis sp.

B Young, Vecchione and Roper D Young, Vecchione and Roper Pyroteuthis Hoyle, Pyroteuthis margaritifera Rppel, Pyroteuthis addolux Young, Pyroteuthis serrata Riddell, Pterygioteuthis Fischer, Pterygioteuthis giardi Fischer, Pterygioteuthis gemmata Chun, Pterygioteuthis microlampas Berry, Cephalopod catches have increased steadily in the last 40 years, from about 1 million metric tonnes in to more than 4 million metric tonnes in FAO, This increase confirms a potential development of the fishery predicted by G.

Voss in , in his first general review of the worlds cephalopod resources prepared for FAO. The rapid expansion of cephalopod fisheries in the decade or so following the publication of Vosss review, meant that a more comprehensive and updated compilation was required, particularly for cephalopod fishery biologists, zoologists and students.

Roper, M. Sweeney and C. Nauen was published in to meet this need. For example, this work should be useful for the ever-expanding search for development and utilization of natural products, pharmaceuticals, etc. The catalogue is based primarily on information available in published literature.

However, yet-to-be-published reports and working documents also have been used when appropriate, especially from geographical areas where a large body of published information and data are lacking. We are particularly grateful to colleagues worldwide who have supplied us with fisheries information, as well as bibliographies of local cephalopod literature.

This information is supplemented by field observations made by the authors in many parts of the world, both in preparation of the volume, as well as for the current edition. These field visits provided opportunities to examine fresh material at landing sites, markets and laboratories, as well as to obtain first-hand information about local cephalopod fisheries from regional fisheries workers.

Additional examinations of preserved specimens occurred in museums. During the plus years separating the two editions, the rapid development of cephalopod fisheries worldwide and the simultaneous increase in the population of fisheries scientists, their research and publications, made available an enormous amount of new data and research results. Sometimes it is difficult to evaluate the reliability of published data, especially with regard to the identification of species in areas where the cephalopod fauna has not been sufficiently studied taxonomically.

Moreover, the discovery of new species, the more accurate delimitation of known species, or even the introduction of nomenclatural changes, may cause confusion and lead to the use of scientific names that are incorrect by modern standards. Although great care was exercised to evaluate and correct such published information used in the preparation of this catalogue, some incorrect interpretations may have occurred.

Another potential limitation, in the taxonomic literature especially, is that information on the economic importance of species is rather scarce or of a very general nature. Also, important information may have been overlooked if published only in local fisheries literature that is unavailable on an international scale.

All of these potential limitations, however, have been significantly mitigated during the preparation of the new edition because of the availability of on-line fisheries databases and bibliographic search capabilities. With regard to the limitations mentioned above, we heartily request that readers who detect any errors in the information presented, or who have additional information and data that will enhance the accuracy and utility of this book, please contact and inform one of the authors or the Species Identification and Data Programme SIDP of the Marine Resources Service, Fisheries Resources Division, Fisheries Department, FAO Rome.

The number of cephalopod species that enter commercial fisheries has continued to grow significantly since , as a result of a still-growing market demand and the expansion of fisheries operations to new fishing areas and to deeper waters. It has been suggested that the cephalopod life-strategy may guarantee survival against environmentally stressful conditions, including those caused by heavy fishing. However, as cephalopod fisheries experienced further extensive development, parallel concern developed regarding potential overexploitation.

Thus, a broad consensus emerged among fishery biologists to apply the experience gained from errors made in finfish management to avoid possible failures in cephalopod exploitation. To help prevent potential failures, refined species identification capabilities are required, as well as a more detailed and accurate compilation of information on cephalopod species, distribution, biology, fisheries and catch statistics.

Consequently, FAO recognized that a new edition of the Cephalopods of the World catalogue was required. To achieve this expanded goal, several authors with particular areas of specialization were assembled to enhance the accuracy, coverage and utility of this revised catalogue.

In our attempt to make this document as comprehensive and as useful as possible, the taxonomic coverage of this edition of the catalogue is organized into 3 levels of interest: Level 1: species of cephalopods currently exploited commercially and species utilized at the subsistence and artisanal levels; Level 2: species of occasional and fortuitous interest to fisheries; this includes species considered to have a potential value to fisheries, based on criteria such as edibility, presumed abundance, accessibility, marketability, etc.

Level 3: species with no current interest to fisheries, which are listed only with the basic information available. The inclusion of such a wide range of species is necessary to provide the most comprehensive inventory of species potentially useful to mankind, regardless of their current. Species identifications should be attempted only after verification of the family through use of the illustrated key to families. Sizes or measurements might not be completely comparable because they were taken mostly from preserved or fixed specimens, but measurements of commercially important species often come from fresh material.

Because of the elasticity of tentacles and arms, total length is not a very accurate measurement. Where both total length and mantle length are given, the respective figures do not necessarily pertain to the same specimen but may have been obtained from different sources.

The available information on the size attained by some species often is very meagre, so the maximum reported size cited here might be considerably smaller than the actual maximum size. Maximum weight is given when available. In cases where only scattered records of occurrence are available, question marks have been used to indicate areas of suspected or unconfirmed distribution.

For the sake of exactness actual depth data are reported, as given in the referenced literature. Information on biological aspects, such as migration, spawning season and area, longevity, prey, and predators, also is included. Due to the dominant role of squids in the marine environment, this section is especially detailed in this volume. Data on utilization fresh, dried, cooked, frozen, canned, etc.

Here, too, the quality and quantity of the available information varies considerably among the species, and it is reported in as much detail as possible in relation to the squids significance to the fisheries. The present compilation is necessarily incomplete, since only a fraction of the local names applied to specific entities actually is published. In many cases, local names are available only for species that support traditional fisheries.

Apart from possible omissions due to limitations of literature available, some of the names included may be somewhat artificial, i. 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. The type genus within each family is treated first, then all remaining genera are listed alphabetically. The type species within each genus is treated first, then all species are listed alphabetically.

Level 1 includes the most important species for fisheries utilization, and it consists of detailed information in all 12 categories listed below. Level 2, which comprises those species of occasional or potential interest to fisheries, consists of whatever information is available and appropriate for the 12 categories.

Level 3, those species for which there is no current direct or indirect interest to fisheries, consists of basic information i. The format within the species sections includes the first two levels of treatment Level 1 and Level 2 presented together. Species included in Level 3 are presented at the end of each family.

Consequently, each major group and family is introduced with general descriptive remarks, illustrations of diagnostic features, highlights of the biology and relevance to fisheries. The information that pertains to each species in Levels 1 and 2 is arranged by categories as follows: 1 scientific name; 2 synonymy; 3 misidentifications; 4 FAO names; 5 diagnostic features with illustrations; 6 maximum known size; 7 geographical distribution with map; 8 habitat and biology; 9 interest to fisheries; 10 local names; 11 remarks 12 literature.

Wherever possible, these names are selected based on vernacular names or parts of names already in existence within the areas where the species is fished. FAO species names, of course, are not intended to replace local species names, but they are considered necessary to. Cephalopods of the World 11 Remarks: Important information concerning the species, but not specifically linked to any of the previous categories, is given here.

For example, in some cases the taxonomic status of certain scientific names requires further discussion. Other nomenclatural problems are discussed in this section, such as the use of subspecies names. Additional references are included in the bibliography. In the case of a few uncommon species, only systematic papers are available. The massive amount of literature relevant to fisheries for many species of squids required that appendices be compiled for this Volume.

The appendix includes a list of publications useful to gain an understanding of the species biology, ecology and fisheries. Publications are listed by authors name, date of publication and key words for the publications contents. For practical purposes we separate the cephalopods into several groups, without assigning or implying taxonomic relationships. Figure 1 diagrams several of the classification schemes currently under discussion.

Cephalopods include exclusively marine animals that live in all oceans of the world with the exception of the Black Sea, from the Arctic Sea to the Antarctic Ocean and from the surface waters down into the deep sea. Cephalopods first appeared as a separate molluscan taxonomic entity, the nautiloids, in the Upper Cambrian period over million years ago , but more than half of these ancestors were already extinct by the end of the Silurian, million years ago, when only the nautiluses survived.

Meanwhile, other forms arose in the late Palaeozoic between and million years ago , including those of the Subclass Coleoidea, but most of them became extinct by the end of the Mesozoic, about million years ago. The only members of the subclass Coleoidea that exist today are the forms that developed in the Upper Triassic and Lower Jurassic between and million years ago.

Although there is a long fossil record of many different groups, all living cephalopods belong to two subclasses: the Coleoidea, which includes the major groups known as squids, cuttlefishes sensu lato, octopods and vampires, and the Nautiloidea, containing two genera, Nautilus and Allonautilus, the only surviving cephalopods with an external shell.

At the present time the status and understanding of the Systematics and Classification of the Recent Cephalopoda is under considerable discussion. The families of living cephalopods are, for the most part, well resolved and relatively well accepted.

Species-level taxa usually can be placed in well-defined families. The higher classification, however, still is not resolved. The classification above the family level is controversial and a broad consensus still needs to be achieved.

This situation is not unexpected for a group of organisms that has undergone explosive research attention in recent decades. These terms are also used to indicate the major groups. The term cuttlefishes also is used sensu lato to indicate the following groups: Cuttlefishes, Bobtail squids, Bottletail squids, Pygmy squids and the Rams horn squid.

Cuttlefishes, along with Nautiluses were treated in Volume 1 Jereb and Roper, Octopods will be treated in Volume 3. This second volume of the Catalogue is focused on Squids. Salinity is considered to be a limiting factor in squids distribution; they are generally restricted to salinity concentr ations bet ween 27 and However, Lolliguncula brevis, which lives and reproduces in waters of 17, demonstrates a capacity for a higher degree of salinity tolerance Hendrix et al. Some species. The habitat depth range extends from the intertidal to over 5 m.

Many species of oceanic squids undergo diel vertical migrations: they occur at depths of about to m during the day, then at the onset of twilight and increasing darkness, they ascend into the uppermost m for the night. A deeper-living layer of diel migrators occurs from about 1 m to m during the daytime.

The abundance of squids varies, depending on genera, habitat and season, from isolated individuals, small schools with a few dozen individuals, to huge schools of neritic and oceanic species with millions of specimens.

General charcteristics The size of adult squids ranges from less than 10 mm mantle length e. The largest specimens may weigh over kg, but the average size of commercial species is to mm mantle length and about 0. Squids are soft-bodied, bilaterally symmetrical animals with a well-developed head and a body that consists of the muscular mantle, the mantle cavity that houses the internal organs, and the external fins.

The head bears an anterior circum-oral surrounding the mouth crown of mobile appendages arms, tentacles. The mouth, at the interior base of the arm crown, has a pair of chitinous jaws the beaks and, as in other molluscs, a chitinous tongue-like radula band of teeth. The ancestral mollusc shell is reduced to a rigid structure composed of chitin, the gladius or pen, sometimes quite thin and flexible.

The loss of the external shell allowed the development of a powerful muscular mantle that became the main locomotory organ for fast swimming, via water jettisoned from the funnel. The funnel also known as siphon, an archaic term correctly applied to some other molluscs, but not to modern, extant cephalopods is a unique, multifunctional, muscular structure that aids in respiration and expulsion of materials, in addition to locomotion. Oxygenated water is drawn through the mantle opening around the head neck into the mantle cavity, where it bathes the gills for respiration.

Muscular mantle contraction expels the deoxygenated water from the mantle cavity through the ventrally located funnel. The discharge jet. Female reproductive products eggs, egg masses also are discharged through the funnel. Squids produce ink, a dark, viscous fluid also expelled through the funnel. The ink may take the form of a mucoidal pseudomorph false body to decoy potential predators, or of a cloud to obscure the escaping cephalopod. One pair of gills ctenidia is present, for respiration, i.

Squids may use anerobic muscle layers, and cutaneous respiration also occurs. The circulatory system The circulatory system is distinctive within the Mollusca. It is a closed system blood contained within vessels , similar in many respects to that of vertebrates, that fulfills the demand for the more efficient circulation required by an active locomotory system. The system is composed of a principal, or systemic, heart, two branchial hearts and developed arterial, venous and capillary systems that supply blood to the muscles and organs.

The oxygenated blood passes from the gills through the efferent branchial vessels to the systemic heart, where it is expelled from the ventricle through three aortas: the cephalic or dorsal aorta, which supplies the head and the anterior part of the gut; the posterior, minor or abdominal aorta that supplies the mantle and fins along with the posterior part of the gut and the funnel; and the gonadal aorta that develops gradually with sexual maturation of the animal.

The blood is collected through sinuses and capillaries into the veins, through which it passes to the branchial hearts that pump it through the filaments of the gills. The circulating respiratory pigment used for oxygen transport is copper-containing haemocyanin, a system of rather lower efficiency than the iron-containing haemoglobin of vertebrates. Blood sinuses in living squids are much reduced and replaced functionally by muscles.

The circulatory system therefore has to work against the peripheral muscle-induced pressure, which increases with increasing activity maximum during jet-swimming. It also has to cope with the resistance of the small diameter of the final capillary blood vessels, and the low oxygen carrying capacity of the blood less than 4.

In spite of these limitations, the system has other functional modifications see for example Wells and Smith, ; Martin and Voight, that achieve the capacity to deliver oxygen at a rate comparable to that of active fishes, enabling squids to accomplish extraordinary swimming, attack and escape performances.

The excretory system The excretory system also differs markedly from that of other molluscs and, along with the closed circulatory system and the branchial circulation, enables unique relationships between blood and the final secretion, the urine. The excretory system consists basically of the renal sac with the renal appendages organs comparable to vertebrate kidneys , the pericardial glands, the branchial hearts and the gills.

Squids are ammoniotelic, whereby ammonium ions are continuously released by the gill epithelium and by renal appendages into the surrounding water. Ammonium ions are used by buoyant squids to replace denser chloride ions in fluids in the coelom and in the body tissues.

Because this solution is less dense and hence more buoyant than seawater, it provides lift for neutral or positive buoyancy. Cephalopods of the World The nervous system The nervous system is highly developed, with a large brain and peripheral connections, contrasting with the original molluscan circumesophageal nerve ring. Among its most remarkable features is the giant fibre system that connects the central nervous system with the mantle muscles.

This system consists of three orders of cells and fibres and ensures the immediate and simultaneous contraction of mantle, fins and retractor muscles of both sides, rather than an anterior to posterior sequential contraction that would be counter-productive for water movement expulsion. Also remarkable is the eye development of squids, for which vision plays a major role in life. Their eyes are large, have a design generally similar to that of fishes and other vertebrates e. This is extremely important for hunters that rely on sight, and it is accomplished by connections of the eye muscles with the statocysts, a bilateral mechanism similar to the vestibulo-optic system of fishes.

The statocyst system provides squids with information on their orientation, as well as changes in position and direction of movement. It is a highly developed system that consists of two separate cavities located bilaterally in the cartilaginous skull, posteroventral to the brain. The statocysts contain nervous cells and receptors differentiated to detect both linear acceleration, with the aid of calcareous stones called statoliths, and angular acceleration.

Some squids also have extra-ocular photoreceptors photosensitive vesicles about which little is known; in mesopelagic squids they appear to monitor light intensity in order to enable the animals to match their counter-illumination with the ambient light with their own photophores light-producing organs.

Squids are provided with numerous mechano- and chemoreceptors and recent evidence indicates that in some species, e. Loligo vulgaris, ciliate cells form lines in several parts of the body, a system analogous to the lateral-line system in fishes. Squids are able to change colour by using a complex system of chromatophores under nervous control. The chromatophores are pigment-filled sacs present in the skin, and capable of remarkable expansion and contraction.

This system responds virtually instantaneously to contemporary situations in the environment, and it is critical for survival. Squid species also have iridocytes shiny, reflective platelets in the skin. Squids behaviour includes rapid changes in overall colour and colour pattern and many deep-sea forms camouflage themselves by producing bioluminescent light from photophores which eliminate their silhouettes against the down-welling sunlight in the dimly-lit mid-depths.

This capability enables squids to inhabit open water, even in the great depths in the ocean, the greatest volume of living space on earth. Feeding Squids are voracious, active predators that feed upon crustaceans, fishes and other cephalopods. A common hunting technique involves extremely rapid shooting forward of the tentacles to capture the prey, while in some oegopsid squids the tentacles may be used like long, sucker-covered fishing lures.

The captured prey is brought to the mouth and killed by bites of the strong, chitinous beaks, equipped with powerful muscles. Reproduction Squids are dioecious separate sexes and many species, though not all, exhibit external sexual dimorphism, either in morphological or morphometric differences. Females frequently are larger than males and males of most species possess one, occasionally two, modified arm s the hectocotylus for transferring spermatophores to females during mating.

The males of some species also exhibit modifications to other arms, in addition to the hectocotylus. The hectocotylus may be simple or complex and can consist of modified suckers, papillae, membranes, ridges and grooves, flaps. The one or two nuptual limbs function to transfer the spermatophores tubular sperm packets from the males reproductive tract to an implantation site on the female. The spermatophores may be implanted inside the mantle cavity where they may penetrate the ovary , into the oviducts themselves, around the mantle opening on the neck, on the head, in a pocket under the eye, around the mouth or in other locations.

Females of a few species also develop gender-s pecific struc tures e. Mating often is preceded or accompanied by courtship behaviour that involves striking chromatophore patterns and display. Copulatory behaviour varies significantly among species, in colour and textural display, proximity of male and female, duration of display and spermatophore transfer, and the location of implantation of the spermatophores on the female. The gonads form a single mass at the posterior end of the mantle cavity, and female gonoducts may be paired in oegopsids or single, as in other squids.

The reproductive systems are highly complex structures with ducts, glands and storage organs. Female squids have nidamental glands and loliginids have accessory nidamental glands, as well. Spermatophores are produced in the multi-unit spermatophoric gland and stored in the Needhams sac, from which they are released through the terminal part of the duct, the penis. This term is not strictly accurate,. Locomotion Locomotion is achieved by a combination of jet propulsion and flapping or undulating the fins on the mantle.

The fins on the mantle also provide balance and steering during jet propulsion. The number and size of spermatophores vary greatly, depending on the species and group for reviews on spermatophore structures and function see Mann et al. Once in contact with seawater, the so called sper matophori c react ion begins. T he spermatophores evert, with the resultant extrusion of the sperm packet caused by the penetration of water inside the spermatophoric cavity, where the osmotic pressure is higher.

The resulting extruded sperm packet is named spermatangium or sperm bulb or body. Sperm are able to survive several months once stored in the female, at least in some species, and fertilization of mature ova may take place either in the ovary, the mantle cavity or the arm cone formed by the outstretched arms while the eggs are laid.

Fertilized eggs are embedded in one or more layers of protective coatings produced by the nidamental glands and generally are laid as egg masses. Egg masses may be benthic or pelagic. Eggs of neritic, inshore squids, except in Sepioteuthis, generally are very small only a few millimetres in diameter and frequently are laid in finger-like pods each containing from a few to several hundred eggs.

Deposited in multi-finger masses sometimes called sea mops , these eggs are attached to rocks, shells or other hard substrates on the bottom in shallow waters. Many oceanic squids lay their eggs into large sausage-shaped or spherical gelatinous masses containing tens or even hundreds of thousands of eggs that drift submerged in the open sea. This has led to changes in our conceptions about the physiology and ecology of many species, but more research is required before a full understanding is achieved see Jereb et al.

Principal results obtained from the research generally confirm a very high growth rate in squids, comparable to that of the fastest-growing fishes. The life expectancy of most squids appears to range from a few months to one or two years, and many small oceanic squids, such as pyroteuthids may complete their life cycles in less than six months.

Recent evidence, however, suggests that larger species of squids, for example the giant squid Architeuthis spp. All squids die after their spawning period. Systematics status The total number of living species of squids that currently are recognized is more than ; are listed in the present volume.

The status of the systematics of squids has changed in the last 30 years, as research and associated scientific discussions have increased substantially. However, phylogenetic relationships among many families remain uncertain, and new species are described fairly frequently as new habitats are explored and as families are gradually better-understood. Growth and life history Development of squid embryos is direct, without true metamorphic stages. However, hatchlings undergo gradual changes in proportions during development and the young of some species differ from the adults.

Thus, the term paralarva has been introduced for these early stages of cephalopods that differ morphologically and ecologically from older stages. The paralarvae of many deep-sea species of squids occur in the upper m of the open ocean; then they exhibit an ontogenetic descent, gradually descending to deeper depths with increasing size until the adult depth is attained. Time of embryonic development varies widely, from a few days to many months, depending on the species and the temperature conditions.

Hatching may occur synchronously from a single clutch or be extended over a period of 2 or 3 weeks. In spite of the large number of studies and research carried out on squids, especially in recent decades, the life history of many species still is unknown, and our knowledge of the life cycles of the members of this interesting group remains fragmentary. Information comes from studies in the field as well as from observations in the laboratory. However, little is known of life history for species that are not targets of regular fisheries, and only a few squid species have been reared successfully in the laboratory.

Studies and monitoring of growth are complicated by the high variability in individual growth rates. This makes it difficult to apply conventional methods, e. Determination of age also is difficult, because squids have few hard structures that show daily marks rings that enable direct estimates of age. In the last Conclusions Squids are important experimental animals in biomedical research with direct applications to human physiology and neurology, for example.

Because of their highly developed brains and sensory organs, they are valuable in behavioural and comparative neuro-anatomical studies. In addition, the extremely large single nerve axons of some squids, the largest in the animal kingdom, are used extensively in neuro-physiological research. The bite of squid can be painful at the least to humans, or secondarily infected, or, rarely, lethal. A documented threat by squids to humans is from the large ommastrephid squid, Dosidicus gigas, which forms large aggressive schools that are known to have attacked fishermen that have fallen in the water, causing several confirmed deaths.

Scuba divers also have been attacked. Therefore, squids must be handled carefully. Of the total cephalopod catch of over 4 million tonnes reported for by FAO statistics FAO, , over 3 million tonnes were squids, i. The impressive increase in squid production during the last 25 years is due.

Cephalopods of the World mainly to the discovery and increasing exploitation of squid resources in the southwest Atlantic, principally for Illex argentinus, as well as an increase in the production of other major squid target species, mainly Todarodes pacificus in the northwest Pacific and Dosidicus gigas in the eastern Pacific.

Illex argentinus catches exceeded 1 million tonnes in , a record peak which placed this species at the eleventh position in value of the total world marine-species production for that year. Numerous fishing techniques and methods to capture squids have been developed over time. These were extensively reviewed, for example, by Rathjen , [] and Roper and Rathjen They include lures, jigs, lampara nets, midwater trawls and otter trawls.

Jigging is the most widely used method, which accounts for almost half of the world squid catch, primarily ommastrephids, but also a few loliginids. This technique is employed primarily at night, when many species of squids are attracted to the fishing vessel by lights. Figure 3 shows the distribution of the worlds light fishery for some of the most important squid species.

Jigs, which feature numerous, variously-arranged, barbless hooks Plate I, 6 , are lowered and retrieved by jigging machines that simulate the constant swimming behaviour of natural prey, inducing. While simple hand-jigging machines are still used in small-scale, artisanal fisheries, large modern vessels for industrial fishing activities are equipped with scores of automated, computer-controlled jigging machines, each capable of catching several tonnes per night Plate I, 1 and 5.

Trawling is the secondmost productive fishery method to catch squids Plate I, 2. Formerly, almost all squids were caught as bycatch in trawl fisheries for finfishes and shrimps. Trawling is a very efficient technique to catch species, but soft-bodied animals like cephalopods are often damaged by the other species in the catch, particularly in benthic and epibenthic otter trawls.

Even in fisheries in which squid-specific trawling occurs, the huge catches of squids per tow often result in crushed and damaged product. Consequently, trawled squid product generally is less valuable than jig-caught squids. However, modern oceanic trawlers can process on board many metric tonnes of cephalopods per day, which helps insure a high-quality product.

Bottom trawling can be very dangerous for benthic habitats because of the physical damage it causes to the seabed and associated fauna and because of its lack of selectivity. Consequently, less intense exploitation by this traditional fishing technique and an approach toward diversification of methods and redistribution of the fisheries through different areas were encouraged and still are highly recommended, especially in situations where small-scale fisheries still exist and new, more efficient methods can be implemented.

Nearshore, neritic squids frequently are caught by purse seines, lift nets, beach seines, etc. Products range from fresh food, eaten raw as sashimi in Japan and, in recent years, worldwide, and fresh-cooked, as well as various types of processed product dried, canned, frozen, reduced to meal, etc. The high protein and low fat content of cephalopods make them an important and healthy element in the human diet. Considering the present level of exploitation of the commercially-fished squid populations, a further increase in such fishery production is likely to occur, first by expansion of the fisheries into the less-fished regions of the oceans, e.

There, a standing stock of squid biomass as high as million tonnes was estimated by scientists, based on an estimate of 30 million tonnes consumed by vertebrate predators see Rodhouse et al. Therefore, a priority for the future research in the field of Antarctic cephalopod biology will be to assess the squid biomass there, quantitatively and qualitatively, with the objective of determining and developing a sustainable fishery production.

However, polar squids probably are longer living and slower growing than species currently harvested. Therefore, caution must be exercised in assumptions and decisions for management of polar squid fisheries. In the future, it is likely that attention will be focused on finding other species and families to replace fish stocks that become severely reduced by overfishing. Even though clear evidence reveals the existence of large cephalopod resources available for exploitation in the open oceans, based on the estimated consumption by predators see Clarke, b; Piatkowski et al.

Research is being carried out on how to remove this factor on a commercial scale, but results will take time and catches will need to be processed before marketing and utilization. A number of ommastrephid squids that lack ammonium are considered to be underexploited. T hese include: Sthenoteuthis pteropus, Ommastrephes bartramii, Martialia hyadesi, Todarodes sagittatus, Sthenoteuthis oualaniensis, Nototodarus philippinensis, Dosidicus gigas, and the circumpolar, sub-Antarctic Todarodes filippovae.

Exploitation of these species would provide large tonnages of high quality cephalopods and would require only minor development in catching techniques. However, it will be necessary to determine where these species congregate for feeding and spawning activities. An analysis of biomass, production and potential catch for the Ommastrephidae species is presented in Nigmatullin Although a number of other oceanic squid families have large populations and high quality flesh, they are not currently exploited on a commercial scale except for a few seasonal fisheries.

These include members of the families Thysanoteuthidae, Gonatidae and Pholidoteuthidae, for example. Increased exploitation of these groups, however, would also require some research and development of catching techniques. Commercial exploitation of the cosmopolitan family Histioteuthidae also could be considered, since at least one large commercial-level catch has been made in the North Atlantic see Okutani, personal communication, in Clarke, a.

However, the increased exploitation of these oceanic squid species might have unpredictable, far-reaching negative effects on the. Therefore, great caution must be exercised in developing this kind of fishery. Almost all of our knowledge of the general biology of cephalopods, in fact, is limited to the shelf-living species, as well as to those ommastrephids that move onto the shelf at certain seasons. Even so, many gaps still exist in our knowledge about their life cycles, especially as far as the relationships among species are concerned e.

Some populations of harvested species have shown sudden, occasionally catastrophic, declines before adequate biological data could be gathered and analysed. Squid stocks experienced true collapses at least in two well-known and documented cases.

These were the northwest Pacific Todarodes pacificus fishery failure in the s and the northwest Atlantic Illex illecebrosus fishery collapse in the s. While the T. These collapses are thought to have occurred mainly as a consequence of temporarily unfavourable environmental conditions or actual long-term environmental changes, probably aggravated by heavy fishing pressure Dawe and Warren, A significant challenge thus exists to deepen our knowledge and learn the details of distribution, life history and biology of exploited species in order to allow rational utilization of the stocks.

The necessity for research as a key factor towards attaining this goal has been stressed by many authors e. Lipinski et al. Therefore, squids are potentially very good indicator species to predict or reflect changes in environmental conditions, both locally and on a broader scale see Pierce et al. Perhaps even more significant is the challenge that exists for future exploitation of new species or populations.

The role of squids in the ecosystem, in fact, is more complex than it was thought to be only a few decades ago. Squids can be considered subdominant predators that tend to increase in biomass when other species, particularly their predators and competitors for food, become depleted, as a result of a combination of heavy or excessive fishing, other human impacts, oceanogr aphic fluc tuations and competition for food see Caddy, , and Caddy and Rodhouse, for a detailed analysis of the transition from finfish-targeted fisheries to cephalopod-targeted fisheries.

In turn, squids are major food items in the diets of innumerable species of fishes, toothed whales e. Muscular squids derive their energy from crustaceans, fishes and other cephalopods. At the same time, they are a very efficient food storage for the large, oceanic predators, by rapidly converting oceanic resources into high energy food.

On the other hand, neutrally buoyant ammoniacal squids, which probably greatly outnumber the muscular squids in biomass, also provide food to many of the same. Cephalopods of the World predators, but not over the continental shelf and with consistently lower energy per unit body mass. We know virtually nothing about the details of feeding, growth, life cycles, periodicities, distribution and spawning in ammoniacal species.

In spite of our relatively incomplete knowledge, it is now clear that squids are a dominant component within marine ecosystems and that their abundance ultimately may influence the abundance of their predator and prey populations. Studies of the effects of consumption of important pelagic squids and fishes by predatory fishes on the northeastern shelf of the United States Overholtz et al.

Consistent with our present knowledge. Taking into consideration these factors, increasing effort should be focused on improving scientific knowledge of this group. Squid catches need increased monitoring, especially in those areas of major environmental fluct uations and where fisher ies management is complicated by multiple countries exploiting the same resource. Cooperation, collaboration and commitment are required to better understand these important and fascinating animals.

Abyssal The great depths of the ocean: from 2 to 6 m. Accessory nidamental glands Glands of unknown function; consist of tubules containing symbiotic bacteria. Found in all decapodiformes except oegopsid squids. Adult A female that has mature eggs these frequently are stored in the oviducts , or a male that has produced spermatophores these are stored in Needhams sac. Afferent blood vessel Artery vessel carrying blood toward an organ.

Afferent nerve Nerve carrying impulses toward the brain or specific ganglia. Anal flaps A pair of fleshy papillae involved in directing releases of ink, 1 flap situated at each side of the anus Fig. Anterior Toward the head-end or toward the arm-tips of cephalopods. Anterior salivary glands Glands on or in the buccal mass that aid in preliminary digestion. Anterior suboesophageal mass See Brachial lobe. Antitragus Knob that projects inward from the posterior surface of the central depression in the funnel-locking apparatus of some squids Fig.

Anus Terminal opening of the digestive tract, in the anterior mantle cavity, sometimes extending to inside the funnel, through which digestive waste products, as well as ink, are expelled. Apomorphic Derived from a more ancestral condition. Loosely considered the advanced condition. Arm One of the circumoral appendages of cephalopods.

Arms are designated by the numbers I to IV, starting with I as the dorsal or upper pair. In squids each appendage of the fourth ancestral pair is modified to form a tentacle. Arm formula Comparative length of the 4 pairs of arms expressed numerically in decreasing order: the largest arm is indicated first and the shortest last, e.

Bathypelagic The deep midwater region of the ocean. Beak One of the 2 chitinous jaws of squids bound in powerful muscles. The dorsal beak is referred to as the upper beak and it inserts within the lower ventral beak to tear tissue with a scissors-like cutting action. Belemnoidea A fossil group of cephalopods that is thought to be the sister group of the Coleoidea.

Belemnoids are distinguished by the presence of hook-like structures on the arms rather than suckers. Benthopelagic A free-swimming animal that lives just above the ocean floor but rarely rests on the ocean floor. Bilateral symmetry The symmetry exhibited by an organism or an organ if only one plane can divide the animal structure into 2 halves that are mirror images of each other.

Bioluminescence The production of light by living organisms, sometimes called living light. The light is produced through a chemical reaction that generally takes place in complex organs called photophores or light organs. Brachial Pertaining to the arms. Brachial crown The combination of arms and tentacles that surround the mouth.

Brachial lobe of the brain The anteriormost part of the brain located ventral to the oesophagus. The large axial nerve cords that run down the centres of the arms connect to this lobe. The proper name is anterior suboesophageal mass. Brachial photophore Photophore located on the arms. Brachial pillar A narrow, elongate anterior region on the paralarval or juvenile head of some families, between the eyes and the base of the brachial crown; especially well developed in young cranchiid squids.

Brain Medial portion of the central nervous system that includes the suboesophageal and supraoesophageal masses but generally does not include the large optic lobes. Branchial Pertaining to the gills. Branchial canal A large opening at the base of each gill lamella and between the primary afferent and efferent blood vessels of the gill. Branchial gland Elongate or spheroidal gland adjacent and parallel to the gill attachment to the mantle wall.

Branchial heart A gland at the base of the gill through which afferent blood is pumped to the gill. It also is the site of hemocyanin the blood respiratory pigment synthesis. Brooding Incubation of eggs by the female. A characteristic feature of incirrate octopods, but also found in some squids e. Buccal Pertaining to the mouth.

Buccal connective Thin muscular band that attaches the buccal support of the buccal membrane to the base of buccal membrane. The position of attachment of the connective on the fourth arms was recognized in the early twentieth c entury as an important charact er for phylogenetic relationships among decapodiformes Fig. Buccal crown Umbrella-like structure that surrounds the mouth and in turn is enveloped by the brachial crown. It consists of buccal supports and the buccal membrane.

Buccal lappet A small, subtriangular flap at the tip of each buccal support of the buccal membrane; thought to be homologous with the inner ring of tentacles that surrounds the mouth of nautiluses. May bear suckers Fig. Buccal mass Muscular bulb at the anteriormost part of the digestive system that consists of the mouth, beaks, radula, muscles and pairs of salivary glands. Buccal membrane The muscular membrane that encircles the mouth like an umbrella Fig.

It connects the buccal supports to form the buccal crown. The pigmentation of the buccal membrane often differs from that of the adjacent oral surfaces of the arms. Buccal membrane connectives See Buccal connective Fig. Buccal support Muscular rod fused to buccal membrane as supporting rib; 6 to 8 in number Fig.

A neutrally buoyant object does not rise or sink but maintains its position in the water; a positively buoyant object will rise and a negatively buoyant object will sink. Caecal sac The sac-like, thin-walled posterior portion of the caecum in the digestive tract that lacks the internal, ciliated leaflets characteristic of the anterior portion of the caecum.

Caecum Region of the digestive tract of all cephalopods between the stomach and intestine. It is the primary site of food absorption. Calcified Chalky, calcareous material of calcium salts calcium carbonate , formed by deposition. Carpal knobs Small, rounded, hemispherical, muscular protuberances on the carpus to which carpal suckers from the opposite club adhere during the locking of the clubs Fig.

Carpal-locking apparatus Arrangement of suckers and matching knobs on the carpal region of the tentacular club that permits the 2 clubs to be locked together Fig. Cephalic cartilage Cartilage-like tissue that envelops the p o s t e r i o r pa r t o f t h e b r a i n o f c e p h a l o p o d s a n d encompasses the statocysts. Anteriorly the cartilage thins and entwines with muscular tissue, which makes a well-defined limit difficult to distinguish.

The cartilage has a large central foramen through which the oesophagus passes and minor foramina for nerves and blood vessels. Cephalic vein Large vein that drains blood from the head region; it lies along the ventral surface of the visceral sac, beside or dorsal to the intestine. The cephalic vein terminates by dividing into the 2 vena cavae, each of which passes through the kidney nephridium , the branchial heart and into the gill.

C e p h al o p o d a T he class within the Mollusca, characterized by bilateral symmetry, internal shell or absence of shell except nautiluses , anterior head, appendages and funnel, posterior mantle, mantle cavity with organs, and shell and fins when present. Character state A particular condition of a taxonomic character. For example, the character sucker may include the 2 states: sucker with a horny ring or sucker without a horny ring.

Chemotactile Refers to chemical and touch sensitivity. Chitin ous A horny polysaccharide substance fingernail-like that forms the sucker rings, hooks and beaks. Chorion A tough secreted membrane that encapsules the egg. Chromatophores Pigment-filled muscular sacs in the skin under individual nervous control that collectively provide the background colour, colour patterns and colour dynamics play of cephalopods.

Circumoral appendages The 8 arms plus the 2 tentacles. All arise from the head and encircle the mouth Fig. Clade A monophyletic group. That is, a group whose members share a closer common ancestor with one another than with members of any other group. Coelom An internal body cavity of mesodermal orgin that is lined by an epithelium. Cephalopods have 2 coeloms, the viscero-pericardial coelom and the nephridial coelom. Collar Muscular, flange-like structure that extends from the nuchal cartilage to the funnel; it forms a one-way valve that allows water to enter the mantle cavity but closes as the mantle contracts, thereby forcing exhalent water out through the funnel.

Carpus The proximal zone of small suckers and knobs on the base of the tentacular club in some families Fig. Cartilaginous structures or scales Cartilage-like structures in the skin of certain squids; may be overlapping and scale-like, or multifaceted platelets, knobs or papillae Fig. Cephalopods of the World Cone, conus The spoon-like, cup-like, spiked or simple conical posterior terminus of the gladius; homologous to the phragmacone of fossil squids Fig.

III suckers keel rachis buccal membrane funnel groove oegopsid eye funnel funnel-mantle fusion vane funnel-locking cartilage mantle-locking cartilage mantle photophores fin lobe fin posteriorly concave myopsid eye tentacle II I IV sheathed hooks. Conus field The sides of the conus that continue anteriorly along the vanes of the gladius. Cornea Smooth, thin, turgid, transparent skin without muscles that covers the eyes to protect the eye lenses of myopsid squids Fig.

Counter illumination The production of bioluminescent light by an animal to conceal its silhouette against a lighted background. The process can allow an animal to become virtually invisible under dim directional light. Cretaceous The last period of the Mesozoic Era. Cusp A point or projection on a tooth of the radula or on a cartilagenous tubercule in the skin.

Dactylus The distal, terminal section of the tentacular club, often characterized by suckers of reduced size Fig. Decapodiformes Higher-level taxon that includes all limbed cephalopods Fig. Within the Decapodiformes, typically, two higher taxa are recognized: the Sepioidea, which includes the Sepiidae, Idiosepiidae, Sepiolidae, Spirulidae and Sepiadariidae and the Teuthoidea, which includes Myopsid and Oegopsid squids.

Because of the long history of referring to these cephalopods by the common name decapods, the latter is maintained as the common name for the Decapodiformes. Demersal Organisms that live close to the ocean floor. Diel vertical migration Vertical animal migration during twilight periods. Many mesopelagic animals migrate to shallow depths at sunset, where they spend the night feeding; then they descend at sunrise from near-surface waters to spend the day hiding at greater, darker depths.

Some animals migrate vertically over 1 m, others migrate less than m. Digestive gland Primary organ in cephalopods that secretes digestive enzymes. It is also important in absorption and excretion Fig. Digestive gland duct appendages Outpockets of the ducts leading from the digestive gland that are covered with glandular epithelium Fig.

Distal Away from the central region of the body or point of origin; toward the peripheral parts opposite of proximal. Accidentals are: garnet, hornblende, muscovite, pyroxene. Granite is used as building stone, dimensional stone, ornamental stone. It is transformed into crushed stones, tombstones, decorative cradft objects, rolls and millstones, cladding and flooring in- and outside.

DVD information. Description, technical characteristics, manufacturer-specific recommendations for cleaning, protection und maintenance, quarry operators, wholesalers. Das Standardwerk der Natursteinbranche. Granite Granite is a visibly granular, plutonic rock, formed in a depth of some kilometers.

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Health Loading. Get Help Loading. Extras Loading. This page needs Javascript enabled in order to work properly. Click here for instructions on how to enable it in your browse. Ready to discover your family story? First Name. Last Name. You can see how Bettinger families moved over time by selecting different census years. The most Bettinger families were found in the USA in In there were 14 Bettinger families living in New York.

New York had the highest population of Bettinger families in Use census records and voter lists to see where families with the Bettinger surname lived. Within census records, you can often find information like name of household members, ages, birthplaces, residences, and occupations. United States. Top Male Occupations in Farmer.

Top Female Occupations in Housewife. Census Record There are 10, census records available for the last name Bettinger. Search US census records for Bettinger. Passenger List There are 1, immigration records available for the last name Bettinger.

View all Bettinger immigration records. Draft Card There are 2, military records available for the last name Bettinger.