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| | | | Tuna and tuna-like species are very important economically and a significant source of food, with the so-called principal market tuna species the most significant in terms of catch weight and trade. These pages are a collection of Fact Sheets providing detailed information on tuna and tuna-like species. | Taxonomic Classification | [ Family: Scombridae ] : Scombrids [ Family: Istiophoridae Family: Xiphiidae ] : Billfishes | Upper systematics of tunas and tuna-like species Scombrids and billfishes belong to the suborder of the Scombroidei which position is shown below: | | Phylum Chordata | | | Subphylum Vertebrata | | | Superclass Gnathostomata | | | Class Osteichthyes | | | Subclass Actinopterygii | | | Infraclass Teleostei | | | Superorder Acanthopterygii | | | Order Perciformes | | | Suborder Scombroidei | | | See more on Scombrids... | ScombridsThe scombrids belong to the family of the Scombridae which is comprised of 15 genera and 50 species. These species are assigned to 2 sub-families: the Gasterochismatinae with only 1 species and the Scombrinae divided into 4 tribes: | | the mackerels (Scombrini) | | | the seerfishes or Spanish mackerels (Scomberomorini) | | | the bonitos (Sardini) | | | the tunas (Thunnini) | The tribe Thunnini (tunas for ichthylogists) contains 14 species in 4 genera: | | Thunnus (8 species) | | | Katsuwonus (1 species) | | | Euthynnus (3 species) | | | Auxis (2 species) | However, in several languages, the word "tuna" is used for members of both the tribe Thunnini and the tribe Sardini (8 species).Origin of the word "tuna"The word "tuna" is applied to certain members of the family Scombridae, a group of marine fishes including tunas, bonitos, mackerels, seerfishes and the butterfly kingfish. However, for ichthyologists, tuna refers to any of the 14 species of the tribe Thunnini within the family Scombridae (Klawe, 1977). The word "tuna" seems to have come into use in the second half of the last century, but it is not clear why it replaced the older name "tunny". It may have been brought to southern California by fishermen originated from Europe, either from the Dalmatian coast of the former Yugoslavia or from the Iberian Peninsula (Klawe, 1976). Principal market tuna species Among the 14 species of tuna, skipjack, yellowfin, bigeye and albacore constitute more than 80% of the world catch since 1950. Three other species, Atlantic bluefin, Pacific bluefin and southern bluefin tuna, are also commercially important due to the high prices paid for them. These above-mentioned species are often referred as "principal market tuna species". | | hide this box | | See more on Billfishes... | BillfishesThe billfishes include 2 families: the Xiphiidae with one species, the swordfish (Xiphias gladius) and the Istiophoridae with 12 species within 4 genera. | | hide this box | | See more on "tuna common names"... | Scientific and common names in English, French and Spanish of tunas and tuna-like species | Scientific name | Common names, acknowleged by FAO, in: English French Spanish; 3-alpha code | Thunnini Thunnus alalunga Thunnus albacares Thunnus atlanticus Thunnus maccoyii Thunnus obesus Thunnus thynnus Thunnus tonggol Katsuwonus pelamis Euthynnus affinis Euthynnus alleteratus Euthynnus lineatus Auxis rochei Auxis thazard | Tunas Albacore Yellowfin tuna Blackfin tuna Southern bluefin tuna Bigeye tuna Northern bluefin tuna Longtail tuna Skipjack Kawakawa Little tunny Black skipjack Bullet tuna Frigate tuna | Thons Germon Albacore Thon à nageoires noires Thon rouge du Sud Thon obèse Thon rouge Thon mignon Listao Thonine orientale Thonine commune Thonine noire Bonitou Auxide | Atún Atún blanco Rabil Atún aleta negra Atún del Sur Patudo Atún Atún tongol Listado Bacoreta oriental Bacoreta Barrilete negro Melvera Melva | ALB YFT BLF SBF BET BFT LOT SKJ KAW LTA BKJ BLT FRI | Sardini Allothunnus fallai Cybiosarda elegans Gymnosarda unicolor Orcynopsis unicolor Sarda australis Sarda chiliensis Sarda orientalis Sarda sarda | Bonitos Slender tuna Leaping bonito Dogtooth tuna Plain bonito Australian bonito Eastern Pacific bonito Indo-Pacific bonito Atlantic bonito | Bonites Thon élégant Bonite à dos tacheté Bonite à gros yeux Palomette Bonite bagnard Bonite du Pac. oriental Bonite orientale Bonite à dos rayé | Bonitos Atún lanzón Bonito saltador Tasarte ojón Tasarte Bonito austral Bonito del Pac. oriental Bonito mono Bonito atlántico | SLT LEB DOT BOP BAU BEP BIP BON | Scientific name | Common names, acknowleged by FAO, in: English French Spanish; 3-alpha code | Grammatorcynus bicarinatus Grammatorcynus bilineatus | Shark mackerel Double-lined mackerel | Thazard requin Thazard-kusara | Carite cazón Carite cazón pintado | SHM DBM | Scomberomorini Acanthocybium solandri Scomberomorus brasiliensis Scomberomorus cavalla Scomberomorus commerson Scomberomorus concolor Scomberomorus guttatus Scomberomorus koreanus Scomberomorus lineolatus Scomberomorus maculatus Scomberomorus multiradius Scomberomorus munroi Scomberomorus niphonius Scomberomorus plurilineatus Scomberomorus queenslandicus Scomberomorus regalis Scomberomorus semifasciatus Scomberomorus sierra Scomberomorus sinensis Scomberomorus tritor | Seerfishes Wahoo Serra Spanish mackerel King mackerel Narrow-barred king mack. Monterey Spanish mackerel Indo-Pacific king mackerel Korean seerfish Streaked seerfish Atlantic Spanish mackerel Papuan seerfish Australian spotted mackerel Japanese Spanish mackerel Kanadi kingfish Queensland school mackerel Cero Broad-barred king mackerel Pacific sierra Chinese seerfish West African Spanish mackerel | Thazards Thazard-bâtard Thazard serra Thazard barre Thazard rayé Thazard Monterey Thazard ponctué Thazard coréen Thazard cirrus Thazard atlantique Thazard papou Thazard australien Thazard oriental Thazard Kanadi Thazard de Queensland Thazard franc Thazard tigre Thazard sierra Thazard nébuleux Thazard blanc | Carites Peto Serra Carite lucio Carite estriado Carite de Monterey Carite del Indo-Pacifico Carite coreano Carite rayado Carite atlántico Carite papuense Carite australiano Carite oriental Carite canadí Carite de Queensland Carite chinigua Carite tigre Carite sierra Carite indochino Carite lusitánico | WAH BRS KGM COM MOS GUT KOS STS SSM PAP ASM NPH KAK QUM CER BBM SIE CHY MAW | Scombrini Rastrelliger brachysoma Rastrelliger faughni Rastrelliger kanagurta Scomber australasicus Scomber japonicus Scomber scombrus | Mackerel Short mackerel Island mackerel Indian mackerel Spotted chub mackerel Chub mackerel Atlantic mackerel | Maquereaux Maquereau trapu Maquereau des îles Maquereau des Indes Maquereau tacheté Maquereau espagnol Maquereau commun | Caballa Caballa rechoncha Caballa isleña Caballa de la India Caballa pintoja Estorino Caballa del Atlántico | RAB RAF RAG MAA MAS MAC | Gasterochismatinae Gasterochisma melampus | Butterfly kingfish | Thon papillon | Atún chauchera | BUK | Scientific name | Common names, acknowleged by FAO, in: English French Spanish; 3-alpha code | Xiphias gladius Istiophorus albicans Istiophorus platypterus Makaira indica Makaira mazara Makaira nigricans Tetrapterus albidus Tetrapterus angustirostris Tetrapterus audax Tetrapterus belone Tetrapterus georgei Tetrapterus pfluegeri | Swordfish Atlantic sailfish Indo-Pacific sailfish Black marlin Indo-Pacific blue marlin Atlantix blue marlin Atlantic white marlin Shortbill spearfish Striped marlin Mediterranean spearfish Roundscale spearfish Longbill spearfish | Espadon Voilier de l'Atantique Voilier de l'Indo-Pacifique Makaire noir Makaire bleu de l'Indo-Pacifique Makaire bleu de l'Atlantique Makaire blanc de l'Atlantique Makaire à rostre court Marlin rayé Marlin de la Méditerranée Makaire épée Makaire bécune | Pez espada Pez vela del Atlántico Pez vela del Indo-Pacífico Aguja negra Aguja azul del Indo-Pacífico Aguja azul del Atlántico Aguja blanca del Atlántico Marlín trompa corta Marlín rayado Marlín del Mediterráneo Marlín peto Aguja picuda | SWO SAI SFA BLM BLZ BUM WHM SSP MLS MSP RSP SPF | | | hide this box | | Fossil records The first fossil records of Scombridae are dated from the beginning of the Eocene epoch (60-40 million years ago) during the Tertiary period. As for the Istiophoridae, the oldest fossils are dated from the upper Cretaceous epoch (70-90 million years ago) during the Secondary period. For the Xiphiidae, the oldest fossil records are dated from the Paleocene epoch of the lower Tertiary period, i.e. 57-65 million years ago (Berg, 1958). | Biological characteristics | Diagnostic Features | | Morphology Morphology of larvae It is often difficult or impossible to identify larvae and, in some cases, early juveniles by anatomical characteristics or colour patterns. Biochemical or genetic methods can be used to distinguished the larvae of the various species (Elliott and Ward, 1995). Morphology of juveniles and adults Characteristics common to both scombrids and billfishes Both scombrids and billfishes have two distinct dorsal fins, generally separated, the first one supported by spines and the second only by soft rays. The pelvic fins are inserted below the base of the pectoral fins. The caudal fin is deeply notched. All scombrids and billfishes except swordfish have a pair of caudal keels on the middle of the caudal peduncle at the base of the caudal fin. The swordfish has only a large median caudal keel. The more advanced members of the Scombridae family also have a large median keel anterior to the pair of caudal keels. The bodies of all the Scombroidei are robust, elongate and streamlined. The first dorsal and first anal fins of all scombrids and billfishes, except swordfish, can fold down into grooves and the pectoral and pelvic fins into depressions when the fish is swimming rapidly. The scombrids and billfishes, all have four gill arches on each side. The gill filaments are ossified as "Gill rays". | | See more on Scombrids... | ScombridsThe scombrids are characterised by the presence of at least four finlets behind the dorsal and the anal fins. The pelvic fins are smaller than the pectoral fins or of equal size. Except for the primitive butterfly kingfish, which body is covered by large cycloid scales, the body of all the scombrids is naked or covered with small to moderate-sized scales. The bonitos (Sardini) are intermediate between the seerfishes and the tunas. As is the case for the tunas, they have a well-developed corselet of scales, but they lack the two longitudinal ridges on the upper surface of the tongue. The most primitive Scombrinae are the mackerels (Scombrini), the seerfishes (Scomberomorini) and the two-line mackerels (Grammatorcynus spp.). The mackerels have only two caudal keels, whereas the seerfishes and the two-line mackerels have a larger median keel in front of the pair of keels. The tunas are the most highly evolved of the scombrids. They are unique among bony fish in having heat exchanger systems that allow them to regulate their body temperature, as can birds and mammals (see the thermoregulation). | | hide this box | | See more on Billfishes... | BillfishesThe billfishes are characterised by their rostrum, an extension of the upper jaw, which extends much beyond the lower jaw. Billfishes rostrum | | | The rostrum has a flat, sword-like cross section for the swordfish and a rounded, spear-like cross section for the Istiophoridae. The swordfish is also characterised by the absence of pelvic fins and scales. Its dorsal fins are well separated, and has only one large median caudal keel. The Istiophoridae have long, rigid, tapering pelvic fins. Their bodies are covered with small, elongate, bony scales. Their first dorsal fin has a long base and terminates close to the origin of the second dorsal fin. | | hide this box | | Geographical Distribution | | Tropical and temperate tunas Because of different distributions due to their specific thermal tolerances and because of exploitation by different fisheries, a distinction is made between tropical and temperate tunas. Tropical tunas are found in waters with temperatures greater than 18° C (although they can dive in colder waters) whereas temperate tuna are found in waters as cold as 10°C, but can also be found in tropical waters (Brill, 1994). | | Tropical tunas: skipjack and yellowfin | | | Intermediate tunas: bigeye | | | Temperate tunas: albacore, Pacific bluefin, Atlantic bluefin and southern bluefin | | Tropical and temperate tunas | | Tropical tunas Temperate tunas | Scombrids Tunas prefer oceanic waters, and 3 of the 8 species of Thunnus are found worldwide except in the Arctic Ocean. Bonitos and little tunas ( Euthynnus spp.) are primarily coastal fishes, but the distribution of individual species is often widespread. The frigate and bullet tunas ( Auxis spp.) are probably both oceanic and coastal (Olson and Boggs, 1986). The slender tuna and the butterfly kingfish have circum-global distributions in the Southern Ocean. Most mackerels and seerfishes have restricted ranges of distribution. Exceptions are the Spanish mackerel and the wahoo which are found worldwide. Oceanic and neritic tunas Oceanic tunas Neritic tunas | | | Billfishes Billfishes are widely distributed, at least, throughout the oceans in which they occur. The exception are the Mediterranean spearfish, which occurs only in the Mediterranean Sea, and perhaps the roundscale spearfish, which occur in the northeastern Atlantic Ocean around the Canary and Madeira Islands and in the western Mediterranean Sea. However, only the swordfish is cosmopolitan. All other Istiophoridae are being confined to the Atlantic Ocean or to the Indian and Pacific Oceans. Occurrence of the tuna, bonito and billfish species in the different oceans | Common names | Scientific name | Areas of occurrence | | Tunas and bonitos | | Skipjack | Katsuwonus pelamis | worldwide | | Yellowfin tuna | Thunnus albacares | worldwide | | Bigeye tuna | Thunnus obesus | worldwide | | Albacore tuna | Thunnus alalunga | worldwide | | Atlantic bluefin tuna | Thunnus thynnus | Atlantic Ocean | | Pacific bluefin tuna | Thunnus orientalis | Pacific Ocean | | Southern bluefin tuna | Thunnus maccoyii | southern parts of Atlantic, Indian and Pacific Ocean | | Longtail tuna | Thunnus tonggol | Indian Ocean, western Pacific Ocean | | Blackfin tuna | Thunnus atlanticus | western Atlantic Ocean | | Kawakawa | Euthynnus affinis | Indian, western and central Pacific Oceans | | Black skipjack | Euthynnus lineatus | eastern Pacific Ocean | | Little tunny | Euthynnus alleteratus | Atlantic Ocean | | Bullet tuna | Auxis rochei | worldwide | | Frigate tuna | Auxis thazard | Indian and Pacific Oceans | | Slender tuna | Allothunnus fallai | Southern Ocean | | Billfishes | | Swordfish | Xiphias gladius | worldwide | | Atlantic sailfish | Istiophorus albicans | Atlantic Ocean | | Indo-Pacific sailfish | Istiophorus platypterus | Indian and Pacific Oceans | | Black marlin | Makaira indica | Indian and Pacific Oceans | | Indo-Pacific blue marlin | Makaira mazara | Indian and Pacific Oceans | | Atlantic blue marlin | Makaira nigricans | Atlantic Ocean | | Atlantic white marlin | Tetrapterus albidus | Indian and Pacific Oceans | | Striped marlin | Tetrapterus audax | Indian and Pacific Oceans | | Habitat and Biology | | Ecological nicheTunas are pelagic marine fish, spending their entire lives relatively near the surface of tropical, subtropical and temperate oceans and seas. Scombrids and billfishes live primarily in the water layers above the thermocline, but are able to dive to depth of several hundred meters (see the Vertical distribution section). Tuna species attaining only small sizes and juveniles of those attaining large sizes are encountered in epipelagic waters (from the surface to the thermocline) whereas large tunas tend to be mesopelagic and are found also in deeper and cooler waters. | | Epipelagic tunas: skipjack and bonitos | | | Mesopelagic tunas (adults): albacore, bigeye and bluefin | | | Tunas that are found at both depth ranges: yellowfin | Some tunas are found in both offshore and coastal waters and others entirely, or almost entirely, in coastal waters. | | Mid-ocean species: yellowfin and bigeye | | | Coastal species: tonggol | | | Species found in both waters: skipjack, albacore, Pacific bluefin, Atlantic bluefin and southern bluefin | Seerfishes are generally restricted to coastal waters and enter estuaries to feed. One species, the Chinese seerfish moves long distances in freshwater up the Mekong River in China. Tuna and their environment Important environmental parameters for tuna are the sea surface temperature, the quantity of dissolved oxygen in the water and the salinity. Lower thermal boundaries vary between 10°C for temperate tunas and 18°C for tropical tunas (see above; Brill, 1994). The minimum oxygen requirement is estimated between 2 to 2.7 ml/l for principal market tuna species except for bigeye tuna which can tolerate oxygen concentrations as low as 0.6 ml/l (Sharp, 1978 ; Lowe, 2000). Most tunas tend to concentrate along thermal discontinuities such as oceanic fronts (Sund, 1981). Vertical distributionThe vertical distribution of most species of tunas is influenced by the thermal and oxygen structure of the water column. Tuna species attaining only small sizes and juveniles of those attaining large sizes tend to live near the surface, whereas adults of large species are found in deeper waters. The use of deep longlines showed that bigeye can be found at depths as great as 300 m (Suzuki et al., 1977). Albacore are also caught under FADs at depths to about 200 m (Bard et al., 1998). Acoustic telemetry has shown that billfishes are found near to the surface during the day, descending more frequently to greater depths at night (Block et al., 1992a). This is in contrast to the large daily vertical movements of the swordfish, which descends to depths as great as 600 m during the day (Carey and Robison, 1981). | | Schooling behavior Tunas use schooling to their advantage when they forage. Some tunas form parabolic-shaped schools to encircle their prey. Most tunas school according to size. Juveniles of tunas attaining large sizes are, therefore, often associated with tunas attaining only small sizes, such as skipjack or bonito. Schools of large adults consist of a few scattered individuals. Schooling offers protection for juvenile tunas by confusing predators and reducing the likelihood that any single fish will become a victim to a predator. Atlantic bluefin tuna can form giant schools spread over several nautical miles when migrating into the Mediterranean Sea to spawn during the summer. As is the case with the other fishes, the structure of tuna schools is maintained by the lateral line.Migration and other movementsAll tunas and tuna-like fishes move constantly to search for food and to keep water passing over their gills. Migrations are seasonal movements, often over long distances, for the purpose of feeding or reproduction. Temperate tunas, i.e. albacore, Atlantic bluefin, Pacific bluefin, and southern bluefin migrate long distances between temperate waters, where they feed, and tropical waters, where they spawn without moving among different oceans. Southern bluefin tuna also migrates among the Atlantic, Indian and Pacific Oceans. The spawning of the three species of bluefin is restricted to relative small areas of tropical waters. Tropical tunas, i.e. skipjack and yellowfin, are less migratory in terms of long-distance directional movements, although several tagged yellowfin released in the western Atlantic have been recaptured in the eastern Atlantic. Bigeye have some of the characteristics of both temperate and tropical tunas. They apparently do not make trans-oceanic migrations, but like the temperate tunas, they migrate back and forth between feeding grounds in temperate waters and their spawning grounds in tropical waters. When they are not making directional migration, tunas move nearly all the time in search of areas where the food is most abundant. Fishermen are sometimes able to predict on the basis of oceanic conditions where the fish are likely appear and then, they can transfer their operations to those areas. Little is known of the movements of billfishes, but apparently, they make seasonal migrations between temperate waters, where they feed, and tropical waters, where they spawn. Swimming Tunas are excellent swimmers, and their bodies are designed for high performance at both sustainable and burst swimming speeds (Dickson, 1995). Tunas must swim constantly to satisfy their oxygen requirements and consequently stay alive. The direction of movements of some species, such as skipjack, seem to be dictated solely by the availability of food. The movement of other species, such as the three species of bluefin, seem to be influenced by both the distribution of food and the need to return to their ancestral spawning grounds at the proper time. Tunas can move up to 15 km per night in order to forage on organisms that swim upward from deeper waters at that time. | | See more on Scombrids... | Tunas have higher cruising speeds than do other active fishes, including other scombroids (Beamish, 1978; Block et al., 1992b). The morphology of the body and caudal fin of tunas is optimal for prolonged, high-speed swimming. Similar body designs are found also in cetaceans, carangids, certain sharks and even the extinct reptilian ichthyosaurs. Webb (1984) lists the following morphological adaptations: | | a lunate tail of large span, but relatively small chord to maximize the thrust | | | a narrow caudal peduncle to provide for locally-large amplitude displacements and to control the angle of attack | | | a large anterior body depth and mass to reduce recoil energy losses | | | a relatively-rigid streamlined body to both minimize the drag and maximize the thrust | | | hide this box | | Long-range swimming The net distances travelled by tunas and billfishes (shortest distances between the locations of release and recapture) exceed those of any other fish, as shown by the following records obtained from tagging studies (from Joseph et al., 1988): | | 10,790 km for a Pacific bluefin tuna (from southeast of Japan to off Baja California) | | | 10,680 km for a black marlin (from off Baja California to Norfolk Island in the South Pacific Ocean) | | | 9,500 km for a skipjack tuna (from off Baja California to the Marshall Islands) | | | 8,500 km for an albacore tuna (from off California to off Japan) | | | 7,700 km for an Atlantic bluefin tuna (accross the Atlantic Ocean) | In addition, net movements of more than 5,000 km have been recorded for yellowfin tuna, bigeye tuna, blue marlin, striped marlin and sailfish. Short-range, fast swimming Scombrids and billfishes are adapted to fast swimming. The champions are, of course, the most highly evolved scombrids, the bonitos (Sardini) and the tuna (Thunnini) and the billfishes. | They are able to exhibit startling bursts of speed, often exceeding one body length per second. The record (for all bony fishes) belongs to the sailfish ( Istiophorus spp.), which has been clocked at over 110 km/h. Estimated maximun swimming speed of some tunas and billfishes | Species | Sustained in m/s | SL/s | Burst in m/s | SL/s | | Scombrids | | Thunnus albacares(1) | 0.64 | 1.31 | 20.46 | 31 | | Thunnus obesus (1) | 0.6 | 1.3 | - | - | | Thunnus thynnus (1) | 3.49 | 1.64 | - | - | | Katsuwonus pelamis (1) | 0.84 | 2.15 | 9.41 | 19.6 | | Euthynnus affinis (1) | 0.76 | 2.11 | 5 | 10 | | Auxis rochei (1) | 0.68 | 2.19 | - | - | | Sarda chiliensis (1) | 0.88 | 1.54 | 3.70 | 6.49 | | Sarda sarda (1) | 0.35 | 2.18 | 1.2 | 8.58 | | Acanthocybium solandri (1) | 0.4 | 0.32 | 21.23 | 19.3 | | Scomber japonicus (1) | 0.92 | 2.7 | 2.25 | 8.35 | | Scomber scombrus (2) | 0.98 | 3.27 | 5.4 | 18 | | Billfishes | | Tetrapterus audax (3) | 1.8 | - | - | - | | Makaira indica (3) | - | - | 36.1 | - | | Makaira nigricans (3) | 1 | - | 2.25 | - | | Xiphias gladius (2) | - | - | 24.86 | 11.3 | References (1): Magnuson (1978) (2): Wardle and He (1988) (3): Block et al. (1992b) | | Physiological aspects of swimming In order to swim at high speeds for long periods, tunas are capable of taking in and utilizing large amounts of oxygen. In contrast to other fishes that contract their jaws and opercular muscles to pump water over their gills, tunas and billfishes (and some species of sharks) ram ventilate, that is they swim through the water with their mouths open, which forces water over their gills. This is an efficient way to get a large amount of water flowing through the gills at a low energetic cost, but it has an important drawback: tunas cannot stop swimming, or they will suffocate! They must swim at a speed of at least 0.65 m/s to provide sufficient flow of water over thier gills. The amount of gill surface of tunas is up to 30 times those of other fish, and for some tunas the absorptive surface approaches those of the lungs of mammals of comparable weight (Joseph et al., 1988). This large surface enables the tunas to extract about half of the oxygen present in the water flowing over their gills. To transfer oxygen from the gills to the other tissues, tunas have hearts that are about 10 times the size, relative to the weight of the entire body, of those of other fish. The blood pressure of tunas is about three times those of other fishes, and their hearts pump blood at a rate about three times those of other fish. The blood of tuna has a hematocrit of 40%, a value usually associated with diving mammals. Scombrids and billfishes, like most fish, have two types of muscle, white and red. The white muscles function during short bursts of activity, while the red muscles, which have a relatively large mass, allow the fish to swim at high speeds (up to 45 km/h) for long periods without fatigue, as demonstrated by tagging studies with conventional and sonic tags (Joseph et al., 1988 ; Bushnell and Holland, 1997). The proportion of red muscle is much greater for tunas than for other fish (Dickson, 1995) and their white muscles are capable of working in both aerobic and anaerobic conditions. Therefore, the increase in swimming speed can be portrayed as follows : | | sustained speed | high speed | burst speed | | red muscle in action | yes | | | | white muscle in action | | aerobic condition | yes | yes | | | anaerobic condition | yes | yes | yes | The red muscles are located deep within the body, and appear to be more important at the anterior part of the fish. They extend from the vertebral column to a lateral subcutaneous position. In contrast to other fishes, the proportion of red muscle does not seem to increase with the size of the tuna, probably because of greater muscle efficiency and labor sharing between red and white muscles, to which both endothermy and thermoregulation could contribute (Graham et al., 1983). Muscle | | | Heart and white muscle aerobic capacities are significantly greater in tunas than in billfishes and other scombrids. Recovery from intense activities Furthermore, tunas and billfishes are capable of recovering more quickly than other fish after intense activities, such as that involved in capture of prey. For some tunas, the rates of removal of lactate from the blood and white muscle tissue approximate the rates measured in mammals, which allows the tuna to recover within two hours (Dickson, 1995). | | Thermoregulation in tuna As a consequence of swimming constantly to maintain hydrostatic equilibrium (Magnuson, 1973) and oxygenate the blood (Roberts, 1978), muscular metabolism continuously generates heat as a byproduct. Tunas get rid of this excess, but, on the other hand, the heat can be used by the tuna to enable them to forage in cold waters. Metabolic mechanism for thermoregulation Among all bony fish, the Thunnini are unique in their ability to regulate their body temperatures, due to a complex counter-current heat exhanger system, also called the rete mirabile (miraculous network) (Stevens and Neil, 1978). The only other fishes with this system are some sharks of the family Lamnidae (Collette, 1978). The tuna maintain their body temperatures above that of the ambient water by passing arterial blood through vascular countercurrent heat exchangers. All species of tuna have a lateral rete, consisting of small arteries branching from the lateral subcutaneous arteries and small veins emptying into the lateral veins (Graham et al., 1983). In addition, many species of tuna also have a central rete within the vertebral haemal canal, consisting of arteries from the dorsal aorta and veins to the posterior cardinal veins (Stevens and Neil, 1978). The arterial blood is, then, warmed by the venous blood that flows through the red swimming muscles (Holland et al., 1992). The rete mirabile retains between 70 and 99 % of the heat produced by the red muscle fibers, and provide a barrier between the red muscle and the environment (Graham et al., 1983). However, when excessive temperatures have been generated by heavy exercise, tunas appear to be able to control the efficiency of the heat exchangers by closing down some blood vessels of the rete mirabile, allowing heat to dissipate into the colder ambient water (Bushnell and Holland, 1997). Measurements of body temperatures and ambient temperatures with histological analyses of the rete mirabile show that tunas as small as 207 mm in length can maintain their body temperatures more than 3°C above the ambient temperature, and thus can be considered to be endotherms (Graham et al., 1983). Tuna body temperatures are often 10°C greater than those of ambient water. The maximum temperature difference was recorded for an Atlantic bluefin tuna, for which the body temperature was 21.5°C greater than the surrounding water (Graham et al., 1983). The thermoregulatory system cannot conserve heat indefinitely, and when a fish has been foraging in deep, cold water for an extended period, its body temperature decreases. When this happens, it can ascend to warmer water and disengage its thermoregulatory system to allow rapid warming of the tissues (Holland et al., 1992). | Behavioural mechanisms for thermoregulation Combined with the physiological mechanisms, movements into cooler water will facilitate heat dissipation (Bushnell and Holland, 1997). Advantages of thermoregulationThermoregulation allows the tunas to sustain high swimming speeds for long periods and to recover quickly after prolonged exertion (Carey et al., 1971), because most biochemical reactions proceed more rapidly at higher temperatures. Therefore, according to Bushnell and Holland, 1997, elevated body temperatures allow: | | red muscle to contract more quickly, approaching the contraction rate of white muscle and consequently, contributing to high-speed swimming resulting from white muscle contractions | | | more rapid transfer of oxygen from blood to muscle cells | | | more rapid recovery, by enhancing the breakdown of lactic acid | In addition, being "warm bodied" allows the tunas to have a good vision at significant depths by maintaining their brains and eyes at greater than ambient temperatures (Bushnell and Holland, 1997). It also allows the tuna to be more sensitive to thermal gradients (Sharp and Dizon, 1978). Also, because of that, the tuna can forage beneath the thermocline, in deep water, without suffering radical decreases in their core temperature. For example, a bigeye tuna was observed to dive 250 meters in one minute, going from 24°C to 9°C water (Holland et al., 1992). Trophic relations and growth Growth stages The following three stages can be distinguished: | | larvae (recently hatched individuals which are considerably different in appearance from juveniles or adults) | | | juveniles (similar in appearance to adults, but sexually immature) | | | adults (sexually mature fish) | Trophic position of larvae Larvae of tunas and tuna-like fishes live in warm surface waters and feed primarily on the zooplankton including small crustaceans and larvae of crustaceans, fishes, molluscs and jelly-fish. Larvae of tunas and tuna-like species are preyed upon by zooplankton foragers, such as larger larvae and early juveniles of pelagic fishes. Cannibalism is, therefore, an important cause of mortality for tuna larvae. Trophic position of juveniles and adults Tunas and tuna-like fishes in the oceanic food web Tunas and billfishes prey on fish, squid and crustaceans. The larger individuals (wahoo, bonitos, tunas and billfishes), which feed on pelagic fishes, are positioned at the top of the trophic web. The smaller individuals (juvenile tunas and billfishes, mackerels and seerfishes) prey on zooplankton (mainly crustaceans) and constitute part of the ration of large scombroids, sharks and cetaceans. Analyses of stomach contents of yellowfin and skipjack tuna indicate that they feed on small epipelagic fishes between 1 and 10 cm in length (Roger, 1994). Since these prey of yellowfin and skipjack feed directly on zooplankton (mainly copepods), it seems that the tunas are at the top of a short food web, which is probably very efficient from the point of view of energetics. Food web | | Position of tunas and tuna-like fishes in the food web | Food itemsTunas and billfishes are opportunistic feeders. At the species level, they do not have strong preferences for certain types of prey. However, on a regional scale and at a given time, a few species may represent almost all of the food of fish of a specific age group (Cayré et al., 1988). Tunas and billfishes prey on pelagic or epipelagic fishes (including small tunas), crustaceans and molluscs (squids). Coastal tunas feed on neritic and epipelagic prey (Olson and Boggs, 1986). Foraging behavior of juveniles and adults Tunas and billfishes are predators that locate their prey visually. To satisfy their food requirements tunas and billfishes have to swim long distances. Their type of locomotion is, therefore, particularly adapted to the search for prey in a large volume of water with the least expenditure of energy. However, they appear less effective than transient predators, such as esocids, in actually capturing the prey (Webb, 1984). To compensate for this, tunas tend to break up schools of prey, producing disorientation and straggling, and/or search for prey in schools (Webb, 1984; Partridge, 1982). Tunas can detect minute traces of scents of oils, proteins and amino-acids of the mucus layer produced by their prey. When prey is detected, some tunas show changes in their behavior consisting of a general increase of activity: increase in swimming speed, change in swimming pattern, jaw snapping and display of dark stripes on the flanks. Tropical tunas often dive down to or below the thermocline to feed (Holland et al., 1992). It is commonly believed that tunas feed during the day. However, sonic tracking experiments show that some tunas feed also at dusk, when mesopelagic micronecton migrate toward the surface (Bard et al., 1998). | | Size | | Common and maximum sizes of tunas and billfishes Tunas (1) | Scientific name | Common size (in cm) | Maximum size (in cm) | Maximum weigh t (in kg) | | Auxis rochei | 15-35 | 50 | - | | Auxis thazard | 25-40 | 58 | - | | Euthynnus lineatus | 30-65 | 70 | 9 | | Euthynnus alleteratus | 30-80 | 100 | 12 | | Euthynnus affinis | 30-60 | 100 | 13 | | Katsuwonus pelamis | 40-80 | 108 | 33 | | Thunnus atlanticus | 40-70 | 100 | 19 | | Thunnus alalunga | 40-100 | 127 | 40 | | Thunnus tonggol | 40-70 | 130 | 35 | | Thunnus albacares | 60-150 | 200 | 175 | | Thunnus maccoyii | 160-200 | 225 | 160 | | Thunnus obesus | 70-180 | 230 | 200 | | Thunnus thynnus | 80-200 | 300 | 650 | Bonitos (1) | Scientific name | Common size (in cm) | Maximum size (in cm) | Maximum weight (in kg) | | Cybiosarda elegans | 35-45 | 50 | 5 | | Sarda sarda | 30-50 | 85 | 7.5 | | other Sarda * | 30-50 | 100 | - | | Allothunnus fallai | 65-95 | 96 | 10 | | Orcynopsis unicolor | 40-90 | 130 | 13 | | Gymnosarda unicolor | 65-150 | 200 | 131 | * Sarda australis, S. chiliensis, S. orientalis, S. sarda Seerfishes and mackerels (1) | Scientific name | Common size (in cm) | Maximum size (in cm) | Maximum weight (in kg) | | Rastrelliger faughni | - | 20 | 0.75 | | Rastrelliger brachysoma | 15-25 | 35 | - | | Rastrelliger kanagurta | 15-25 | 35 | - | | Grammatorcynus spp. | - | 60 | 3.5 | | Scomber spp. | 15-30 | 40-50 | 1 | | Scomberomorus multiradius | - | 35 | 0.5 | | Scomberomorus concolor | - | 75-80 | - | | Scomberomorus guttatus | - | 75-80 | - | | Scomberomorus niphonius | - | 100 | 4.5 | | Scomberomorus cavalla | 20-70 | 170 | - | | Scomberomorus commerson | 30-90 | 220 | 45 | | Gasterochisma melampus | 74-164 | 164 | - | | Acanthocybium solandri | 100-170 | 210 | 80 | Billfishes (2) | Scientific name | Common size (in cm) | Maximum size (in cm) | Maximum weight (in kg) | | Tetrapterus georgei | - | 160 BL | 21 | | Tetrapterus pfluegeri | - | 200 BL | 45 | | Tetrapterus angustirostris | - | 200 TL | 52 | | Tetrapterus belone | - | 240 BL | 70 | | Tetrapterus albidus | 130-210 BL | 280 TL | 82 | | Tetrapterus audax | 140-280 BL | 350 TL | 200 | | Istiophorus albicans | 125-210 BL | 315 TL | 58 | | Istiophorus platypterus | 140-240 BL | 340 TL | 100 | | Xiphias gladius | 115-190 BL | 445 TL | 540 | | Makaira nigricans | 230-345 TL | 375 TL | 580 | | Makaira indica | 170-210 BL | 448 TL | 700 | | Makaira mazara | 200-300 BL | 447 TL | 900 | Notes for billfishes: BL = Body Length, TL = Total Length References: (1): Collette and Nauen (1983) (2): Nakamura (1985) International Game Fish Association (IGFA) records Tunas and bonitos | Common name | Record size (in cm) | Record weight (in kg) | Location and year of capture | | Sarda chiliensis | - | 6.3 | off Baja California, 1980 | | Euthynnus lineatus | - | 11.8 | off Baja California, 1991 | | Euthynnus alleteratus | 92.7 | 12.2 | off Florida, 1976 | | Katsuwonus pelamis | 99 | 18.9 | off Mauritius island, 1982 | | Thunnus alalunga | 123 | 40 | off Canary Islands, 1972 | | Acanthocybium solandri | - | 72 | off Baja California, 1996 | | Thunnus albacares | 208 | 176.4 | west coast of Mexico, 1977 | | Thunnus obesus | 236 | 197.3 | off Peru, 1957 | | Thunnus thynnus | 304 | 679 | off Nova Scotia, 1979 | Seerfishes and mackerels | Common name | Record size (in cm) | Record weight (in kg) | Location and year of capture | | Scomber japonicus | - | 1.9 | off Baja California, 1986 | Swordfish and billfishes | Common name | Record size (in cm) | Record weight (in kg) | Location and year of capture | | Istiophorus albicans | - | 61.4 | off Nigeria, 1991 | | Tetrapterus albidus | - | 82.5 | off Brazil, 1979 | | Istiophorus platypterus | 327.7 | 100.2 | off Galapagos, 1947 | | Tetrapterus audax | - | 224.0 | off New Zealand, 1986 | | Xiphias gladius | 445 | 536.1 | off Chile, 1953 | | Makaira mazara | - | 624.1 | off Hawaii, 1982 | | Makaira nigricans | - | 636.0 | off Brazil, 1992 | | Makaira indica | 442 | 707.6 | off Peru, 1953 | Source: IGFA (1995) | Growth rates Most scombrids grow rapidly and reach their adult sizes in a few years. Average growth rates vary according to the species, the age and the location. In general, larger tunas grow to about 40 to 55 cm the first year, then the annual growth rate ranges between 20 to 30 cm per year decreasing with age. Tuna species attaining only small sizes grow to 20 to 35 cm in the first year and their annual length increments rapidly decrease to less than 10 cm. In the Atlantic and Indian Oceans, several studies have shown that yellowfin grow rapidly during the first year, slowing their growth during the next one or two years and then having again a fast growth before gradually slowing down as the maximum size is approached. Seerfishes and mackerels have also a fast growth during their first years of life. Sizes of 35 to 45 cm at age 1 year are common. Billfishes can grow to more than 80 cm during their first year of life. After this very fast juvenile growth, adult growth rates are comparable to those of tunas. | | See more on Scombrids... | Growth of juveniles and adults Weights and lengths ranges The maximum weights attained by tunas range from about 1 to 2 kg for bullet and frigate tunas to more than 600 kg for Atlantic bluefin tuna. The maximum lengths attained by tunas range from about 50 cm for bullet and frigate tunas to more than 300 cm for Atlantic bluefin tuna. Seerfishes, mackerels and bonitos are relatively small (less than 1 meter in length), except for some species of seerfishes such as the king mackerel or the narrow-barred king mackerel which grow to more than 170 cm. The smallest billfish is the Mediterranean spearfish, which reaches a maximum length of a little more than 180 cm. The largest billfishes are the black marlin and the Indo-Pacific blue marlin, which reach lengths of more than 4 m and weights of more than 600 kg. | | Maximum sizes of tunas and bonitos | | | hide this box | | Longevity Longevities of tunas vary from a few years for the smaller tunas to 12 to 15 years for the larger tunas. The longevity record for tunas is about 20 years for the Atlantic bluefin tuna (Cort, 1990) and for the southern bluefin tuna. Longevities of 20 years have also been estimated for several species of billfishes. Natural mortality For larger tunas and billfishes, adult natural mortalities range from 0.2 to 0.6. Juvenile natural mortalities are higher. Little is known on natural mortalities of seerfishes and mackerels. Reproduction of tunas and tuna-like fishes Spawning Spawning behavior Tuna spawn in open water close to the surface. Eggs are released by females in several batches. For example, yellowfin tuna in the Pacific spawn nearly every day. However, for some species like bluefin, spawning is more seasonal. Spawning areas and seasonsTunas spawn in areas where the survival of their larvae is greatest. Most species of tunas spawn only in waters where the surface temperatures are greater than 24°C. Tropical tunas appear to spawn in equatorial areas all year around and at higher latitudes during the warm seasons. Albacore and bigeye appear to migrate annually from temperate feeding areas to tropical spawning areas. Bigeye larvae are less abundant than those of other tropical tunas, and are found mainly in equatorial waters in which the temperatures are greater than 28°C (Collette and Nauen, 1983). Atlantic bluefin, Pacific bluefin and southern bluefin tuna exhibit a homing behavior when they mature, and return to restricted areas in the Atlantic, Pacific and Indian Oceans to spawn. It is commonly accepted that there is a homing behavior, but to a lesser extent, in yellowfin in the Atlantic Ocean. Billfishes appear to spawn in warm tropical and subtropical waters. Maturity and fecundity The batch fecundities of most species of tunas range from 2 to 70 million eggs, the lowest fecundity being for albacore and the highest for skipjack tuna and other small-sized tunas. Known batch fecundities of mackerels range from 300 000 to 1 500 000 eggs. Little is known on fecundities of seerfishes. Fecundity of wahoo has been estimated to 6 million eggs. Very little information on reproductive biology of billfishes is available. Swordfish batch fecundity was estimated to 3.9 millions eggs in the Atlantic. Maturity and fecundity parameters of the principal market species of tunas | Scientific name | Size and age at maturity | Annual batch fecundity | | Katsuwonus pelamis | about 3 years and 42 to 45 cm | 7 to 76 million eggs | | Thunnus alalunga | about 5 years and 90 cm | 2 to 3 million eggs | | Thunnus albacares | about 3 years and 100 cm | 4 to 60 million eggs | | Thunnus obesus | about 3 years and 100 cm | 4 to 60 million eggs | | Thunnus thynnus | about 4 years and 105 to 120 cm | 5 to 30 million eggs | Sex ratio It has been shown that for yellowfin, bigeye and albacore, the sex-ratio changes with the age of the fish with a predominance of males for the larger sizes. A predominance of females has also been observed for medium-sized Atlantic bluefin tuna. For skipjack, differences in the numbers of males and females have been observed locally. Predominance of females at older ages is observed for several species of billfishes. | | | Ackowledgment | - Michel Goujon and Jacek Majkowski, who compiled the information contained in this presentation, are grateful to:
- - Marcella Pesce* (consultant involved in the development of FIGIS) for restructuring and editing the information according to FIGIS requirements,
- - Aureliano Gentile (consultant involved in the development of FIGIS) and Yves Jaques (Information Officer of FAO's Fishery Information, Data and Statistic Unit (FIDI)) for incorporating the restructured and edited information into FIGIS and
- - Ignacio de Leiva (Fishery Resources Officer of FAO's Marine Resources Service (FIRM)) for his suggestions for improvements to the presentation.
- *Presently, an expert in the information technology in FAO's Inland Water Resources and Aquaculture Service (FIRI).
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An annotated and illustrated catalogue of marlins, sailfishes, spearfishes and swordfishes known to date. FAO Fish. Synop., 125 (5), Rome: 65 p. Olson, R.J. and H. Boggs, 1986. Apex predation by yellowfin tuna (Thunnus albacares): independent estimates from gastric evacuation and stomach contents, bioenergetics and cesium concentrations. Can. J. Fish. Aquat. Sci. 43 (9):1760-1775. Partridge, B.L., 1982. The structure and function of fish schools. Scient. Amer., 247: 114-123. Roger, C., 1994. The plankton of the tropical western Indian ocean as a biomass indirectly supporting surface tunas (yellowfin, Thunnus albacares and skipjack, Katsuwonus pelamis. Environ. Biol. Fish. 39 (2):161-172. Roberts, J.L., 1978. Ram gill ventilation in fish. pp. 83-88 in : G.D. Sharp and A.D. Dizon (eds), The Physiological Ecology of Tunas, Academic Press, New-York: 485 p. Sharp, G.D., 1978. 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Body form, locomotion and foraging in aquatic vertebrates. Amer. Zool., 24: 107-120. | | | | | |
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