Tuna
Tuna | |
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Tunas (from top): albacore, Atlantic bluefin, skipjack, yellowfin, bigeye | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Actinopterygii |
Order: | Scombriformes |
Family: | Scombridae |
Subfamily: | Scombrinae |
Tribe: | Thunnini Starks, 1910 |
Genera | |
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A tuna (also called tunny) is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae (mackerel) family. The Thunnini comprise 15 species across five genera,[1] the sizes of which vary greatly, ranging from the bullet tuna (max. length: 50 cm (1.6 ft), weight: 1.8 kg (4 lb)) up to the Atlantic bluefin tuna (max. length: 4.6 m (15 ft), weight: 684 kg (1,508 lb)). The bluefin averages 2 m (6.6 ft), and is believed to live up to 50 years.
Tuna, opah, and mackerel sharks are the only species of fish that can maintain a body temperature higher than that of the surrounding water. An active and agile predator, the tuna has a sleek, streamlined body, and is among the fastest-swimming pelagic fish – the yellowfin tuna, for example, is capable of speeds of up to 75 km/h (47 mph).[2] Found in warm seas, it is extensively fished commercially, and is popular as a game fish. As a result of overfishing, stocks of some tuna species, such as the southern bluefin tuna, are close to extinction.[3]
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Commercial fish |
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Large predatory |
Forage |
Demersal |
Mixed |
Etymology
The term "tuna" ultimately derives from Thunnus, the Middle Latin form of the Template:Lang-grc – which is in turn derived from θύνω (thýnō), "rush, dart along".[4][5]
However, the immediate source for the word tuna in English is American Spanish < Spanish atún < Andalusian Arabic at-tūn, assimilated from al-tūn التون [Modern Arabic التن] : 'tuna fish' < Greco-Latin thunnus mentioned above.[6]
Taxonomy
The Thunnini tribe is a monophyletic clade comprising 15 species in five genera:
- family Scombridae
- tribe Thunnini: the tunas
- genus Allothunnus: slender tunas
- genus Auxis: frigate tunas
- genus Euthynnus: little tunas
- genus Katsuwonus: skipjack tunas
- genus Thunnus: albacores, true tunas
- subgenus Thunnus (Thunnus): bluefin group
- subgenus Thunnus (Neothunnus): yellowfin group
- tribe Thunnini: the tunas
- family Scombridae
The cladogram is a tool for visualizing and comparing the evolutionary relationships between taxa, and is read left-to-right as if on a timeline. The following cladogram illustrates the relationship between the tunas and other tribes of the family Scombridae. For example, the cladogram illustrates that the skipjack tunas are more closely related to the true tunas than are the slender tunas (the most primitive of the tunas), and that the next nearest relatives of the tunas are the bonitos of the Sardini tribe.[1]
The Tunas: Thunnini tribe, within the Family Scombridae | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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Cladogram: Tunas are classified into the tribe Thunnini (bottom-center in the above diagram) – one of four tribes in the family Scombridae.[1] |
True tuna species
The "true" tunas are those that belong to the genus Thunnus. Until recently, it was thought that there were seven Thunnus species, and that Atlantic bluefin tuna and Pacific bluefin tuna were subspecies of a single species. In 1999, Collette established that based on both molecular and morphological considerations, they are in fact distinct species.[7][8]
The genus Thunnus is further classified into two subgenera: Thunnus (Thunnus) (the bluefin group), and Thunnus (Neothunnus) (the yellowfin group).
Thunnus, the true tunas Image Common name Scientific name Maximum
lengthCommon
lengthMaximum
weightMaximum
ageTrophic
levelSource IUCN status Thunnus (Thunnus) – the bluefin group Albacore tuna T. alalunga
(Bonnaterre, 1788)1.4 m
(4.6 ft)1.0 m
(3.3 ft)60.3 kg
(133 lb)9–13 yrs 4.31 [9][10] Least Concern[10] Southern bluefin tuna T. maccoyii
(Castelnau, 1872)2.45 m
(8.0 ft)1.6 m
(5.2 ft)260 kg
(570 lb)20–40 yrs 3.93 [11][3] Endangered[3] Bigeye tuna T. obesus
(Lowe, 1839)2.5 m
(8.2 ft)1.8 m
(5.9 ft)210 kg
(460 lb)5–16 yrs 4.49 [12][13] Vulnerable[13] Pacific bluefin tuna T. orientalis
(Temminck & Schlegel, 1844)3.0 m
(9.8 ft)2.0 m
(6.6 ft)450 kg
(990 lb)15–26 yrs 4.21 [14][15] Near Threatened[15] Atlantic bluefin tuna T. thynnus
(Linnaeus, 1758)4.6 m
(15 ft)2.0 m
(6.6 ft)684 kg
(1,508 lb)35–50 yrs 4.43 [16][17] Least Concern[17] Thunnus (Neothunnus) – the yellowfin group Blackfin tuna T. atlanticus
(Lesson, 1831)1.1 m
(3.6 ft)0.7 m
(2.3 ft)22.4 kg
(49 lb)4.13 [18] Least concern[19] Longtail tuna,
northern bluefin tuna,
tongol tunaT. tonggol
(Bleeker, 1851)1.45 m
(4.8 ft)0.7 m
(2.3 ft)35.9 kg
(79 lb)18 years 4.50 [20][21] Data deficient[21] Yellowfin tuna T. albacares
(Bonnaterre, 1788)2.4 m
(7.9 ft)1.5 m
(4.9 ft)200 kg
(440 lb)5–9 yrs 4.34 [22][23] Least Concern[23]
Other tuna species
The Thunnini tribe also includes seven additional species of tuna across four genera. They are:
Other tuna species Common name Scientific name Maximum
lengthCommon
lengthMaximum
weightMaximum
ageTrophic
levelSource IUCN status Slender tuna Allothunnus fallai
(Serventy, 1948)1.05 m
(3.4 ft)0.86 m
(2.8 ft)13.7 kg
(30 lb)3.74 [24] Least concern[25] Bullet tuna Auxis rochei
(Risso, 1810)0.5 m
(1.6 ft)0.35 m
(1.1 ft)1.8 kg
(4.0 lb)5 years 4.13 [26][27] Least concern[27] Frigate tuna Auxis thazard
(Lacépède, 1800)0.65 m
(2.1 ft)0.35 m
(1.1 ft)1.7 kg
(3.7 lb)5 years 4.34 [28] Least concern[29] Mackerel tuna,
KawakawaEuthynnus affinis
(Cantor, 1849)1.0 m
(3.3 ft)0.6 m
(2.0 ft)13.6 kg
(30 lb)6 years 4.50 [30][31] Least concern[31] Little tunny Euthynnus alletteratus
(Rafinesque, 1810)1.2 m
(3.9 ft)0.8 m
(2.6 ft)16.5 kg
(36 lb)10 years 4.13 [32] Least concern[33] Black skipjack tuna Euthynnus lineatus
(Kishinouye, 1920)0.84 m
(2.8 ft)0.6 m
(2.0 ft)11.8 kg
(26 lb)3.83 [34][35] Least concern[35] Skipjack tuna Katsuwonus pelamis
(Linnaeus, 1758)1.1 m
(3.6 ft)0.8 m
(2.6 ft)34.5 kg
(76 lb)6–12 yrs 3.75 [36][37] Least concern[37]
Biology
Description
The tuna is a sleek and streamlined fish, adapted for speed. It has two closely spaced dorsal fins on its back; The first is "depressible" – it can be laid down, flush, in a groove that runs along its back. Seven to ten yellow finlets run from the dorsal fins to the tail, which is lunate – curved like a crescent moon – and tapered to pointy tips. The caudal peduncle, to which the tail is attached, is quite thin, with three stabilizing horizontal keels on each side. The tuna's dorsal side is generally a metallic dark blue, while the ventral side, or underside, is silvery or whitish, for camouflage.[38]
Physiology
Thunnus are widely but sparsely distributed throughout the oceans of the world, generally in tropical and temperate waters at latitudes ranging between about 45° north and south of the equator.[39] All tunas are able to maintain the temperature of certain parts of their body above the temperature of ambient seawater. For example, bluefin can maintain a core body temperature of 25–33 °C (77–91 °F), in water as cold as 6 °C (43 °F). However, unlike "typical" endothermic creatures such as mammals and birds, tuna do not maintain temperature within a relatively narrow range.[40][41]
Tunas achieve endothermy by conserving the heat generated through normal metabolism. In all tunas, the heart operates at ambient temperature, as it receives cooled blood, and coronary circulation is directly from the gills.[42] The rete mirabile ("wonderful net"), the intertwining of veins and arteries in the body's periphery, allows nearly all of the metabolic heat from venous blood to be "re-claimed" and transferred to the arterial blood via a counter-current exchange system, thus mitigating the effects of surface cooling.[43] This allows the tuna to elevate the temperatures of the highly-aerobic tissues of the skeletal muscles, eyes and brain,[40][42] which supports faster swimming speeds and reduced energy expenditure, and which enables them to survive in cooler waters over a wider range of ocean environments than those of other fish.[41]
Also unlike most fish, which have white flesh, the muscle tissue of tuna ranges from pink to dark red. The red myotomal muscles derive their color from myoglobin, an oxygen-binding molecule, which tuna express in quantities far higher than most other fish. The oxygen-rich blood further enables energy delivery to their muscles.[40]
For powerful swimming animals like dolphins and tuna, cavitation may be detrimental, because it limits their maximum swimming speed.[44] Even if they have the power to swim faster, dolphins may have to restrict their speed, because collapsing cavitation bubbles on their tail are too painful. Cavitation also slows tuna, but for a different reason. Unlike dolphins, these fish do not feel the bubbles, because they have bony fins without nerve endings. Nevertheless, they cannot swim faster because the cavitation bubbles create a vapor film around their fins that limits their speed. Lesions have been found on tuna that are consistent with cavitation damage.[44]
Fishing industry
Commercial fishing
Tuna is an important commercial fish. The International Seafood Sustainability Foundation (ISSF) compiled a detailed scientific report on the state of global tuna stocks in 2009, which includes regular updates. According to the ISSF, the most important species for commercial and recreational tuna fisheries are yellowfin (Thunnus albacares), bigeye (T. obesus), bluefin (T. thynnus, T. orientalis, and T. macoyii), albacore (T. alalunga), and skipjack (Katsuwonus pelamis).[39]
Based on catches from 2007, the report states:
Between 1940 and the mid-1960s, the annual world catch of the five principal market species of tunas rose from about 300 thousand tons to about 1 million tons, most of it taken by hook and line. With the development of purse-seine nets, now the predominant gear, catches have risen to more than 4 million tons annually during the last few years. Of these catches, about 68 percent are from the Pacific Ocean, 22 percent from the Indian Ocean, and the remaining 10 percent from the Atlantic Ocean and the Mediterranean Sea. Skipjack makes up about 60 percent of the catch, followed by yellowfin (24 percent), bigeye (10 percent), albacore (5 percent), and bluefin the remainder. Purse-seines take about 62 percent of the world production, longline about 14 percent, pole and line about 11 percent, and a variety of other gears the remainder.[39]
The Australian government alleged in 2006 that Japan had illegally overfished southern bluefin by taking 12,000 to 20,000 tonnes per year instead of the agreed upon 6,000 tonnes; the value of such overfishing would be as much as US$2 billion.[45] Such overfishing has severely damaged bluefin stocks.[46] According to the WWF, "Japan's huge appetite for tuna will take the most sought-after stocks to the brink of commercial extinction unless fisheries agree on more rigid quotas".[47] Japan's Fisheries Research Agency counters that Australian and New Zealand tuna fishing companies under-report their total catches of southern bluefin tuna and ignore internationally mandated total allowable catch totals.[48]
In recent years, opening day fish auctions at Tokyo's Tsukiji fish market have seen record-setting prices for bluefin tuna, reflecting market demand. In each of 2010, 2011, 2012 and 2013, new record prices have been set for a single fish – the current record is 155.4 million japanese yen (US$1.76 million) for a 221 kg (487 lb) bluefin, or a unit price of JP¥ 703,167/kg (US$3,603/lb). The opening auction price for 2014 plummeted to less than 5% of the previous year's price, which had drawn complaints for climbing "way out of line".[49] A summary of record-setting auctions are shown in the following table (highlighted values indicate new world records):
Record bluefin tuna auctions at Tokyo's Tsukiji fish market | ||||||
---|---|---|---|---|---|---|
(highlighted field indicates new record price for a single fish) | ||||||
Year | Total weight |
Total sale | Unit price | Source | ||
( JP ¥ ) | ( US $ ) | ( ¥ / kg ) | ( $ / lb ) | |||
2001 | 202 kg (445 lb) |
¥20.2 million | $173,600 | ¥100,000 / kg | $386 / lb | [50] |
2010 | 232 kg (511 lb) |
¥16.28 million | $175,000 | ¥70,172 / kg | $343 / lb | [51] |
2011 | 342 kg (754 lb) |
¥32.49 million | $396,000 | ¥95,000 / kg | $528 / lb | [50] |
2012 | 269 kg (593 lb) |
¥56.49 million | $736,000 | ¥210,000 / kg | $1,247 / lb | [52] |
2013 | 221 kg (487 lb) |
¥155.4 million | $1.76 million | ¥703,167 / kg | $3,603 / lb | [53] |
In November 2011, a different record was set when a fisherman in Massachusetts caught an 881-pound tuna. It was captured inadvertently using a dragnet. Due to the laws and restrictions on tuna fishing in the United States, federal authorities impounded the fish because it was not caught with a rod and reel. Because of the tuna's deteriorated condition as a result of the trawl net, the fish sold for just under $5,000.[54]
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Tuna being weighed on Greek quay-side
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Tuna at Tsukiji fish market, Tokyo
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Tuna cut in half for processing at Tsukuji fish market
Fishing methods
External videos | |
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Tuna pole and line fishing BBC Two |
Besides for edible purposes, many tuna species are caught frequently as game, often for recreation or for contests in which money is awarded based on weight. Larger specimens are notorious for putting up a fight while hooked, and have been known to injure people who try to catch them, as well as damage their equipment.
- Phoenician technique for trapping and catching Atlantic bluefin tuna called Almadraba, still used today in Portugal, Spain, Morocco & Italy which uses a maze of nets. In Sicily, the same method is called Tonnara.
- Fish farming (Cage system)[55]
- Tuna ranching
- Longline fishing
- Purse seines
- Pole and line
- Harpoon gun
- Big game fishing
- Fish aggregating device
Association with whaling
In 2005, Nauru, defending its vote from Australian criticism at that year's meeting of the International Whaling Commission, argued that some whale species have the potential to devastate Nauru's tuna stocks, and that Nauru's food security and economy relies heavily on fishing.[56] Despite this, Nauru does not permit whaling in its own waters and does not allow other fishing vessels to take or intentionally interact with marine mammals in its Exclusive Economic Zone. In 2010 and 2011 Nauru supported Australian proposals[57] for a western Pacific-wide ban on tuna purse-seining in the vicinity of marine mammals – a measure which was agreed by the Western and Central Pacific Fisheries Commission at its eighth meeting in March 2012.
Association with dolphins
Dolphins swim beside several tuna species. These include yellowfin tuna in the eastern Pacific Ocean, but not albacore. Tuna schools are believed to associate themselves with dolphins for protection against sharks, which are tuna predators.[58]
Commercial fishing vessels used to exploit this association by searching for dolphin pods. Vessels would encircle the pod with nets to catch the tuna beneath,[59] however the nets were prone to entangling dolphins, injuring or killing them. Public outcry and new government regulations, which are now monitored by NOAA have led to more "dolphin friendly" methods, now generally involving lines rather than nets. However, there are neither universal independent inspection programs nor verification of "dolphin safeness", so these protections are not absolute. According to Consumers Union, the resulting lack of accountability means claims of tuna that is "dolphin safe" should be given little credence.
Fishery practices have changed to be dolphin friendly, which has caused greater bycatch including sharks, turtles and other oceanic fish. Fishermen no longer follow dolphins, but concentrate their fisheries around floating objects such as fish aggregation devices, also known as FADs, which attract large populations of other organisms. Measures taken thus far to satisfy the public demand to protect dolphins can be potentially damaging to other species as well.[60]
Aquaculture
Increasing quantities of high-grade tuna are reared in net pens and fed bait fish. In Australia, former fishermen raise southern bluefin tuna, Thunnus maccoyii, and another bluefin species.[55] Farming its close relative, the Atlantic bluefin tuna, Thunnus thynnus, is beginning in the Mediterranean, North America and Japan. Hawaiʻi approved permits for the first U.S. offshore farming of bigeye tuna in water 1,300 feet (400 m) deep in 2009.[61]
Japan is the biggest tuna consuming nation and is also the leader in tuna farming research.[62] Japan first successfully farm-hatched and raised bluefin tuna in 1979. In 2002, it succeeded in completing the reproduction cycle and in 2007, completed a third generation.[63][64][65] The farm breed is known as Kindai tuna. Kindai is the contraction of Kinki University in Japanese (Kinki daigaku).[66] In 2009, Clean Seas, an Australian company which has been receiving assistance from Kinki University[67][68][69] managed to breed Southern Bluefin Tuna in captivity and was awarded the second place in World's Best Invention of 2009 by Time magazine.[70][71]
As food
The fresh or frozen flesh of tuna is widely regarded as a delicacy in most areas where it is shipped, being prepared in a variety of ways for the sake of achieving specific flavors or textures. When served as a steak, the meat of most species is known for its thickness and tough texture.
Canned
When tuna is canned and packaged for sale, the product is sometimes called tuna fish. Canned tuna was first produced in Australia in 1903, quickly becoming popular.[72] Tuna is canned in edible oils, in brine, in water, and in various sauces. Tuna may be processed and labeled as "solid", "chunked" or "flaked". When canned, the product is often referred to as "tuna fish", a calque (loan translation) from the German Thunfisch. In the United States, 52% of canned tuna is used for sandwiches; 22% for tuna salads; and 15% for tuna casseroles and dried and prepackaged meal kits, such as General Mills's Tuna Helper line.[73]
Depending upon the color of the flesh of the tuna species, the can is marked as "light" or "white" meat, with "light" meaning a greyish pink color and "white" meaning a light pink color. In the United States, only albacore can legally be sold in canned form as "white meat tuna";[74] in other countries, yellowfin is also acceptable. While in the early 1980s canned tuna in Australia was most likely Southern bluefin, as of 2003[update] it was usually yellowfin, skipjack, or tongol (labelled "northern bluefin" or "longtail").[72]
As tunas are often caught far from where they are processed, poor interim conservation can lead to spoilage. Tuna is typically gutted by hand, and later pre-cooked for prescribed times of 45 minutes to three hours. The fish are then cleaned and filleted, canned, and sealed, with the dark lateral blood meat often separately canned for pet food. The sealed can is then heated under pressure (called retort cooking) for 2 to 4 hours.[75] This process kills any bacteria, but retains the histamine that may have been produced by those bacteria. The international standard sets the maximum histamine level at 200 milligrams per kilogram. An Australian study of 53 varieties of unflavored canned tuna found none to exceed the safe histamine level, although some had "off" flavors.[72]
Australian standards once required cans of tuna to contain at least 51% tuna, but these regulations were dropped in 2003.[76][77] The remaining weight is usually oil or water. In the US, the Food and Drug Administration (FDA) regulates canned tuna (see part c).[78]
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Tuna steak served in a French bistro -
Katsuobushi shavings
Nutrition
Nutritional value per 100 g (3.5 oz) | |||||||||||||||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Energy | 830 kJ (200 kcal) | ||||||||||||||||||||||||||||||
0 g | |||||||||||||||||||||||||||||||
8 g | |||||||||||||||||||||||||||||||
29 g | |||||||||||||||||||||||||||||||
| |||||||||||||||||||||||||||||||
Other constituents | Quantity | ||||||||||||||||||||||||||||||
Water | 60 g | ||||||||||||||||||||||||||||||
†Percentages estimated using US recommendations for adults,[79] except for potassium, which is estimated based on expert recommendation from the National Academies.[80] |
Canned light tuna in oil is 29% protein, 8% fat, 60% water, and contains no carbohydrates, while providing 200 calories in a 100 gram reference amount (table). It is a rich source (20% or more of the Daily Value, DV) of phosphorus (44% DV) and vitamin D (45% DV), and a moderate source of iron (11% DV).
Mercury levels
Mercury content in tuna can vary widely. Among those calling for improved warnings about mercury in tuna is the American Medical Association, which adopted a policy that physicians should help make their patients more aware of the potential risks.[81] A study published in 2008 found that mercury distribution in the meat of farmed tuna is inversely related to the lipid content, suggesting that higher lipid concentration within edible tissues of tuna raised in captivity might, other factors remaining equal, have a diluting effect on mercury content.[82] Due to their high position in the food chain and the subsequent accumulation of heavy metals from their diet, mercury levels can be high in larger species such as bluefin and bigeye.[citation needed] Mackeral tuna is one species of tuna that is lower in mercury concentration than skipjack or yellowfin,[83] but this species is known as "black meat" or "dark meat" tuna, which is a lower grade for canning because of the color, unfavorable flavor, and poor yield.[84]
In March 2004, the United States FDA issued guidelines recommending that pregnant women, nursing mothers, and children limit their intake of tuna and other predatory fish.[85] The Environmental Protection Agency provides guidelines on how much canned tuna is safe to eat. Roughly speaking, the guidelines recommend one 6-ounce (170 g) can of light tuna per week for individuals weighing less than 110 pounds (50 kg), and two cans per week for those who weigh more.[86] In 2007 it was reported that some canned light tuna such as yellowfin tuna[87] is significantly higher in mercury than skipjack, and caused Consumers Union and other activist groups to advise pregnant women to refrain from consuming canned tuna.[88] In 2009, a California appeals court upheld a ruling that canned tuna does not need warning labels as the methylmercury is naturally occurring.[89]
A January 2008 report revealed potentially dangerous levels of mercury in certain varieties of sushi tuna, reporting levels "so high that the Food and Drug Administration could take legal action to remove the fish from the market."[90]
Management and conservation
The main tuna fishery management bodies are the Western and Central Pacific Fisheries Commission, the Inter-American Tropical Tuna Commission, the Indian Ocean Tuna Commission, the International Commission for the Conservation of Atlantic Tunas, and the Commission for the Conservation of Southern Bluefin Tuna.[91] The five gathered for the first time in Kobe, Japan in January 2007. Environmental organizations made submissions[92] on risks to fisheries and species. The meeting concluded with an action plan drafted by some 60 countries or areas. Concrete steps include issuing certificates of origin to prevent illegal fishing and greater transparency in the setting of regional fishing quotas. The delegates were scheduled to meet at another joint meeting in January or February 2009 in Europe.[93]
In 2010, Greenpeace International added the albacore, bigeye tuna, Pacific bluefin tuna, Atlantic bluefin tuna, southern bluefin tuna, and yellowfin tuna to its seafood red list, which are fish "commonly sold in supermarkets around the world, and which have a very high risk of being sourced from unsustainable fisheries."[94][95]
Bluefin tuna have been widely accepted as being severely overfished, with some stocks at risk of collapse.[96][97] According to the International Seafood Sustainability Foundation (a global, nonprofit partnership between the tuna industry, scientists, and the World Wide Fund for Nature), Indian Ocean yellowfin tuna, Pacific Ocean (eastern and western) bigeye tuna, and North Atlantic albacore tuna are all overfished. In April 2009, no stock of skipjack tuna (which makes up roughly 60% of all tuna fished worldwide) was considered to be overfished.[98] However, the BBC documentary South Pacific, which first aired in May 2009, stated that, should fishing in the Pacific continue at its current rate, populations of all tuna species could collapse within five years.[99] It highlighted huge Japanese and European tuna fishing vessels, sent to the South Pacific international waters after overfishing their own fish stocks to the point of collapse.
A 2010 tuna fishery assessment report, released in January 2012 by the Secretariat of the Pacific Community, supported this finding, recommending that all tuna fishing should be reduced or limited to current levels and that limits on skipjack fishing be considered.[100]
Research[101] indicates that increasing ocean temperatures are taking a toll on the tuna in the Indian Ocean, where rapid warming of the ocean has resulted in a reduction of marine phytoplankton. The bigeye tuna catch rates have also declined abruptly during the past half century, mostly due to increased industrial fisheries, with the ocean warming adding further stress to the fish species.[101]
See also
References
- ^ a b c Graham, Jeffrey B.; Dickson, Kathryn A. (2004). "Tuna Comparative Physiology" (PDF). The Journal of Experimental Biology. 207: 4015–4024. doi:10.1242/jeb.01267. Retrieved 20 September 2012.
- ^ Block, Barbara A.; Booth, David; Carey, Francis G. (1992). "Direct measurement of swimming speeds and depth of blue marlin" (PDF). Journal of Experimental Biology. 166. Company of Biologists Ltd.: 267–284. ISSN 0022-0949. Retrieved 19 September 2012.
- ^ a b c Collette, B.; et al. (2021). "Thunnus maccoyii". IUCN Red List of Threatened Species. 2021. Retrieved 29 March 2022.
- ^ θύννος in Liddell, Henry George; Scott, Robert (1940) A Greek–English Lexicon, revised and augmented throughout by Jones, Sir Henry Stuart, with the assistance of McKenzie, Roderick. Oxford: Clarendon Press. In the Perseus Digital Library, Tufts University.
- ^ θύνω in Liddell and Scott.
- ^ Houghton Mifflin Harcourt Publishing Company. "American Heritage Dictionary Entry: tuna". ahdictionary.com.
- ^ Collette, B.B. (1999). "Mackerels, molecules, and morphology". In Séret, B.; Sire, J.Y. (eds.). Proceedings. 5th Indo-Pacific Fish Conference: Nouméa, New Caledonia, 3–8 November 1997. Paris: Société Française d'Ichtyologie [u.a.] pp. 149–164. ISBN 978-2-9507330-5-4.
{{cite conference}}
: Unknown parameter|booktitle=
ignored (|book-title=
suggested) (help) - ^ Tanaka, Y.; Satoh, K.; Iwahashi, M.; Yamada, H. (2006). "Growth-dependent recruitment of Pacific bluefin tuna Thunnus orientalis in the northwestern Pacific Ocean". Marine Ecology Progress Series. 319: 225–235. doi:10.3354/meps319225.
- ^ Froese, Rainer; Pauly, Daniel (eds.). "Thunnus alalunga". FishBase. January 2012 version.
- ^ a b Collette, B.; et al. (2021). "Thunnus alalunga". IUCN Red List of Threatened Species. 2021. Retrieved 29 March 2022.
- ^ Froese, Rainer; Pauly, Daniel (eds.). "Thunnus maccoyii". FishBase. January 2012 version.
- ^ Froese, Rainer; Pauly, Daniel (eds.). "Thunnus obesus". FishBase. January 2012 version.
- ^ a b Collette, B.; et al. (2021). "Thunnus obesus". IUCN Red List of Threatened Species. 2021. Retrieved 29 March 2022.
- ^ Froese, Rainer; Pauly, Daniel (eds.). "Thunnus orientalis". FishBase. January 2012 version.
- ^ a b Collette, B.; et al. (2021). "Thunnus orientalis". IUCN Red List of Threatened Species. 2021. Retrieved 29 March 2022.
- ^ Froese, Rainer; Pauly, Daniel (eds.). "Thunnus thynnus". FishBase. January 2012 version.
- ^ a b Collette, B.; et al. (2021). "Thunnus thynnus". IUCN Red List of Threatened Species. 2021. Retrieved 9 January 2015.
- ^ Froese, Rainer; Pauly, Daniel (eds.). "Thunnus atlanticus". FishBase. January 2012 version.
- ^ Collette, B.; et al. (2010). "Thunnus atlanticus". IUCN Red List of Threatened Species. 2010. Retrieved 29 March 2022.
- ^ Froese, Rainer; Pauly, Daniel (eds.). "Thunnus tonggol". FishBase. January 2012 version.
- ^ a b Collette, B.; et al. (2009). "Thunnus tonggol". IUCN Red List of Threatened Species. 2009. Retrieved 29 March 2022.
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Oxygenated blood that has just reached thermal equilibrium with ambient sea water in the gills enters the rete on the arterial side, while warmed, deoxygenated, and carbon dioxide-laden blood enters on the venous end. In the rete, countercurrent flow and the high surface area contact between the two blood supplies facilitate the transfer of nearly all of the metabolic heat in the venous blood to arterial blood, thus conserving muscle temperature. After exiting the rete, arterial blood continues to the red muscle capillary beds, and cooled venous blood flows to the gills where carbon dioxide is excreted and oxygen is loaded.
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Further references
- Clover, Charles. 2004. The End of the Line: How Overfishing Is Changing the World and What We Eat. Ebury Press, London. ISBN 0-09-189780-7
- FAO: Species Catalog Vol. 2 Scombrids of the World. FAO Fisheries Synopsis No. 125, Volume 2. FIR/S125 Vol. 2.ISBN 92-5-101381-0
- FAO: Review of the state of world marine fishery resources: Tuna and tuna-like species – Global, 2005 Rome.
- Majkowski, Jacek (1995) "Tuna and tuna-like species" In: Review of the state of world marine fishery resources, FAO Fisheries technical paper 457, FAO, Rome. ISBN 978-92-5-107023-9.
- Majkowski J, Arrizabalaga H, Carocci F and Murua H (2011) "Tuna and tuna-like species" In: Review of the state of world marine fishery resources, pages 227–244, FAO Fisheries technical paper 569, FAO, Rome. ISBN 978-92-5-107023-9.
- Standard of Identity for Canned Tuna (United States), Code of Federal Regulations: 21 CFR 161.190 – Canned tuna.
- Viñas J and Tudela S (2009) "A validated methodology for genetic identification of tuna species (genus Thunnus)" PLoS One, 4(10): e7606.
Other reading
- Bluefin Tuna, Chinese Cobra and Others Added to Red List of Threatened Species, Scientific American, November 18, 2014
- How Hot Tuna (and Some Sharks) Stay Warm National Science Foundation, October 27, 2005