Eel life history
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[[image:leptouk.jpg|thumb|right|350px|Leptocephalus larva of an ocean eel]]
The '''[[eel]]''' is a long, thin bony fish of the order [[Anguilliformes]]. Because fishermen never caught anything they recognized as young eels, the life cycle of the [[eel]] was a mystery for a very long period of scientific history. Although there have been more than 6500 publications about eels, much of its life history remains enigmatic.
The [[European eel]] (''Anguilla anguilla'') was the one most familiar to Western scientists, beginning with [[Aristotle]] who did the first known research on eels. He stated that they are born of "earth worms", which emerged from the mud with no [[fertilization]] needed — they grew from the "guts of wet soil". For a long time, nobody could prove Aristotle wrong. Later scientists believed that the [[eelpout]] ''[[Zoarces viviparus]]'' was the "Mother of Eels" (the translation of the German name "Aalmutter").
In [[1777]], the Italian [[Carlo Mondini]] found the creature's [[gonads]] and proved that eels are [[fish]]. In 1876, the young Austrian student [[Sigmund Freud]] dissected hundreds of eels in search for the male sex organs. He had to concede failure in his first published research paper, and turned to other issues in frustration.[http://www.expertensprechen.de/]
[http://www.berlinonline.de/berliner-zeitung/archiv/.bin/dump.fcgi/2004/1020/feuilleton/0027/index.html] [http://www.wno.org/newpages/sci02b.html]
Until [[1893]], larval eels — transparent, leaflike two-[[inch]] (five [[centimetre|cm]]) creatures of the open ocean — were considered a separate [[species]], ''Leptocephalus brevirostris'' (from the [[Greek language|Greek]] ''leptocephalus'' meaning "thin- or flat-head"). In 1886, French zoologist [[Yves Delage]] kept leptocephali alive in a laboratory tank in [[Roscoff]] until they matured into eels, and in 1896 [[Italy|Italian]] zoologist [[Giovanni Battista Grassi]] observed the transformation of a [[Leptocephalus]] into a round glass eel in the [[Mediterranean Sea]], and recognized the importance of salt water to the process. Despite this discovery, the name ''leptocephalus'' is still used for larval eel.
==Search for the spawning grounds==
{{weasel}}
[[Denmark|Danish]] professor [[Johannes Schmidt (biologist)|Johannes Schmidt]], from [[1904]] onwards, directed many expeditions in the Mediterranean Sea and the [[North Atlantic]], largely financed by the [[J. C. Jacobsen|Carlsberg Foundation]]. He postulated from the similarity of all ''leptocephali'' he found that they all must originate from the same parent species. The further into the [[Atlantic Ocean]] he propelled research ships, the smaller the ''leptocephali'' he caught. Finally, in [[1922]], he ended up south of [[Bermuda]] in the [[Sargasso Sea]] where he succeeded in catching the smallest eel-larvae ever seen.
However, Schmidt was unable to observe the spawning directly, nor did he find ready-to-spawn adults. From the size distribution of the leptocephali he collected, Schmidt formulated this part of the life history of the eel:
[[image:anguillamuk.jpg|thumb|left|275px|Distribution and size of leptocephali larvae of the European Eel, ''Anguilla anguilla''.]]
The larvae of European eels travel with the [[Gulf Stream]] across the ocean and, after one to three years, their leptocephali reach a size of 75 - 90 [[millimetre|mm]] before they reach the coasts of [[England]] and Europe. The common name for this recruiment stage of eels is ''glass eel'', based on the transparency of the body. One famous place for large-scale collection of glass eels (for deli-food and stocking) is [[Epney]] at the [[River Severn|Severn]] in England. Glass eels are also eaten as food in Spain. Once they recruit to coastal areas the migrate up rivers and streams, overcoming all sorts of natural challenges — sometimes by piling up their bodies by the tens of thousands to climb over obstacles — and they reach even the smallest of creeks ([[media:glasseelu.mov|short glasseel migration video]]).
[[Image:Glasseelskils.jpg|thumb|right|275px|glass eels at the transition between ocean and freshwater; the skin is still transparent and the red [[gill]]s and the heart are visible; length ca. 8 cm]]
They can move themselves over wet grass and dig through wet sand to reach upstream headwaters and ponds, thus colonising the continent. In freshwater they develop pigmentation, turn into ''elvers'' (young eels) and feed on creatures like small [[crustacean]]s, [[worm]]s and [[insect]]s. They grow up in 10 or 14 years to a length of 60 to 80 cm. In this stage they are now called ''yellow eels'' because of their golden pigmentation.
[[image:rostrata.jpg|thumb|right|250px|Juvenile eels, length ca. 25 cm]]
In July some individuals mature and then they migrate back towards the sea, crossing even wet grasslands at night to reach rivers that lead to the sea. Eel migration out of their freshwater growth habitats from various parts of Europe, or through the [[Baltic Sea]] in the Danish belts have been the basis of traditional fisheries with characteristic trapnets ([[Bundgarn]]).
How the adults make the 6,000 [[kilometre|km]] (4,000 [[mile]]) open ocean journey back to their spawning grounds north of the [[Antilles]], [[Haiti]], and [[Puerto Rico]] remains unknown. By the time they leave the continent they stop feeding, so they have to rely on stored energy alone. The body undergoes other dramatic changes as well: the [[eye]]s start to enlarge in size, the eye pigments change for optimal vision in dim blue clear ocean light, and the sides of their bodies turn silvery, to create a countershading pattern to make them difficult to see by predators during their long open ocean migration. These migrating eels are typically called "Silver Eels" or "Big Eyes".
The German fisheries biologist [[Friedrich Wilhelm Tesch]], an eel expert and author of the book "The Eel" (ISBN 0-632-06389-0), conducted many expeditions with high-tech instrumentation to follow eel migration, first down the Baltic, then along the coasts of [[Norway]] and England, but finally the transmitter signals were lost at the [[Continental_Shelf|continental shelf]] when the batteries ran out. According to Schmidt a travel speed in the ocean of 15 km per day can be assumed, so a silver eel would need 140 to 150 days to reach the Sargasso Sea from around Scotland and in about 165 to 175 when leaving from the English Channel.
Tesch — like Schmidt — kept on trying to persuade sponsors to give more funding for expeditions. His proposal was to release fifty Silver Eels from Danish waters with transmitters that will detach from the eels each second day, float up and broadcast position, depth and temperature to satellite receivers, possibly jointly with an equivalent release experiment from the countries of the western coast of the Atlantic. However, only preliminary experiments such as these have so far been performed.
Today our some of the only knowledge about the fate of individual silver eels once they leave the continental shelf is based on three eels found in the stomachs of deep sea fishes, that include [[whales]] caught off [[Ireland]] and off the [[Azores]] and some experiments on the physiology of eels in the laboratory.
[[image:Rostratamuk.jpg|thumb|left|275px|Distribution and size of leptocephali larvae of the American eel, ''Anguilla rostrata'']]
There is another Atlantic Eel species: the [[American eel]], ''Anguilla rostrata''. First it was believed European and American eels were the same species due to their similar appearance and behavior, but research has shown that they differ in chromosome count and various molecular genetic markers, and in the number of vertebrae, ''Anguilla anguilla'' counting 110 to 119 and ''Anguilla rostrata'' 103 to 110.
The spawning grounds for the two species are in an overlapping area of the southern Sargasso Sea, with ''A. rostrata'' apparently being more westward than ''A. anguilla'', and with some some spawning by the American eel possibly even occurring off the Yukatan Peninsuala outside of the [[Gulf of Mexico]], but this has not been confirmed. After spawning in the Sargasso Sea and moving to the west, the ''leptocephali'' of the American eel exit the Gulf Stream earlier than the European eel and begin migrating into the estuaries along the east coast of North America between February and late April at an age of about one year and a length of about 60 mm.
The spawning area of the [[Japanese eel]], ''Anguilla japonica'', has also been precisely located to be to the west of the [[Suruga seamount]] ([http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=pubmed&cmd=Retrieve&dopt=AbstractPlus&list_uids=16495988 Oceanic biology: spawning of eels near a seamount - K. Tsukamoto (23 February 2006); Nature 439(7079):929]) and their leptocephali are then transported to the west to East Asia by the North Equatorial Current.
==Decline of the glasseels==
No one yet knows the reasons, but beginning in the mid-[[1980s]], glasseel arrival in the spring dropped drastically — in Germany to 10% and in France to 14% of their previous levels — from even conservative estimates. Data from [[Maine]] and other [[North America]]n coasts showed similar declines, although not as drastic.
In [[1997]] [[Europe]]an demand for eels could not be met for the first time ever, and dealers from Asia bought all they could. The traditional European stocking programs could not compete any longer: each week the price for a [[kilogram]] of glasseel went up another [[US$]]30. Even before the [[1997]] generation hit the coasts of Europe, dealers from [[China]] alone placed advance orders for more than 250,000 kg, some bidding more than $1,100 per kg. Asian elvers have sold in Hong Kong for as much as $5,000 to $6,000 a kilogram at times when $1,000 would buy the same amount of American glasseels with [http://web.archive.org/web/20001203123400/www.ecoscope.com/eelnews.htm gunfights at their catching sites]. Such a kilogram, consisting of 5000 glasseels, may bring at least $60,000 and as much as $150,000 after they leave an Asian fish farm. In New Jersey over 2000 licences for glasseel catch were issued and reports of 38 kg per night and fisherman have been made, although the average catch is closer to 1 kg.
The demand for adult eels has continued to grow, [[as of 2003]]. [[Germany]] imported more than $50 million worth of eels in [[2002]]. In Europe 25 million kg are consumed each year, but in [[Japan]] alone more than 100 million kg were consumed in [[1996]]. As the European eels become less available, worldwide interest in American eels has increased dramatically.
New high-tech eel aquaculture plants are appearing in [[Asia]] with detrimental effects on the native Japanese eel, ''[[Anguilla japonica]]''. Traditional eel aquaculture operations rely on wild-caught elvers, but experimental hormone treatments in Japan have led to artificially spawned eels. Eggs from these treated eels have a diameter of about 1 mm, and each female can produce 2 to 10 million eggs.
==Threats to eels==
There are strong concerns that the European eel population might be devastated by a new threat: ''[[Anguillicola crassus]]'', a foreign parasitic [[nematode]]. This parasite from East Asia (the original host is ''Anguilla japonica'') appeared in European eel populations in the early [[1980s]]. Since [[1995]] it also appeared in the [[United States]] ([[Texas]] and [[South Carolina]]), most likely due to uncontrolled aquaculture eel shipments. In Europe, eel populations are already from 30% to 100% infected with the nematode. Recently it was shown that this parasite inhibits the function of the [[swimbladder]] as a [[hydrostatic]] organ (Wuertz et al. 1996). As an open ocean voyager, eels need the carrying capacity of the swimbladder (which makes up 3–6% of the eel's bodyweight) to cross the ocean on stored energy alone.
Because the eels are [[Fish migration|catadromous]] (living in fresh water but spawning in the sea), dams and other river obstructions can block their ability to reach inland feeding grounds. Since the [[1970s]] an increasing number of [[eel ladder]]s have been constructed in North America and Europe to help the fish bypass obstructions.
[[image:rostrataluk.jpg|thumb|right|320px|Glasseel on the online ''in situ'' microscope at the [[Longterm Ecological Observatory|LEO]] project.]]
[[Image:Glasseelkils.gif|thumb|right|320px|]]
In [[New Jersey]], an ongoing project monitors the glasseel migration with an online ''in situ'' microscope. As soon as more funding becomes available, it will be possible to log into the system via a [[Longterm Ecological Observatory]] (LEO) site.
==Further reading==
*Tesch, F-W (2003) The eel. Blackwell Science, Oxford (UK). 1 - 408pp.
*Wallace, Karen (1993) Think of an Eel, Walker Books (UK) - Children's picture book describing the life cycle of the eel.
*Wenner, C.A. (1978). Anguillidae. In W. Fischer (ed.) FAO species identification sheets for fishery purposes. West Atlantic (Fishing Area 31). volume 1. [pag. var.]. FAO, Rome.
*Smith, C.L. (1997). National Audubon Society field guide to tropical marine fishes of the Caribbean, the Gulf of Mexico, Florida, the Bahamas, and Bermuda. Alfred A. Knopf, Inc., New York. 720 p.
*Robins, Richard C., Reeve M. Bailey, Carl E. Bond, James R. Brooker, Ernest A. Lachner, et al. 1980. A List of Common and Scientific Names of Fishes from the United States and Canada, Fourth Edition. American Fisheries Society Special Publication, no. 12. American Fisheries Society. Bethesda, Maryland, USA. 174.
*Robins, Richard C., Reeve M. Bailey, Carl E. Bond, James R. Brooker, Ernest A. Lachner, et al. 1980. A List of Common and Scientific Names of Fishes from the United States and Canada, Fourth Edition. American Fisheries Society Special Publication, no. 12. American Fisheries Society. Bethesda, Maryland, USA. 174.
*Robins, C.R. and G.C. Ray (1986). A field guide to Atlantic coast fishes of North America. Houghton Mifflin Company, Boston, U.S.A. 354 p.
*Page, L.M. and B.M. Burr (1991). A field guide to freshwater fishes of North America north of Mexico. Houghton Mifflin Company, Boston. 432 p.
*Ogden, J.C., J.A. Yntema, and I. Clavijo (1975). An annotated list of the fishes of St. Croix, U.S. Virgin Islands. Spec. Publ. No. 3.
*Nigrelli, R.F. (1959). Longevity of fishes in captivity, with special reference to those kept in the New York Aquarium. p. 212-230. In G.E.W. Wolstehnolmen and M. O'Connor (eds.) Ciba Foundation Colloquium on Ageing: the life span of animals. Vol. 5., Churchill, London.
*Nielsen, J.G. and E. Bertelsen (1992). Fisk i grønlandske farvande. Atuakkiorfik, Nuuk. 65 s.
*Nelson, Joseph S., Edwin J. Crossman, H. Espinosa-Pérez, L. T. Findley, C. R. Gilbert, et al., eds. 2004. Common and scientific names of fishes from the United States, Canada, and Mexico, Sixth Edition. American Fisheries Society Special Publication, no. 29. American Fisheries Society. Bethesda, Maryland, USA. ix + 386. ISBN 1-888569-61-1.
*Murdy, Edward O., Ray S. Birdsong, and John A. Musick 1997. Fishes of Chesapeake Bay. Smithsonian Institution Press. Washington, DC, USA. xi + 324. ISBN 1-56098-638-7.
*Lim, P., Meunier, F.J., Keith, P. and Noël, P.Y. (2002). Atlas des poissons et des crustacés d'eau douce de la Martinique. Patrimoines Naturels, 51: 120p. Paris: MNHN.
*Kenny, J.S. (1995). Views from the bridge: a memoir on the freshwater fishes of Trinidad. Julian S. Kenny, Maracas, St. Joseph, Trinidad and Tobago. 98 p.
*Jessop, B.M. (1987). Migrating American eels in Nova Scotia. Trans. Amer. Fish. Soc. 116:161-170.
*International Game Fish Association (1991). World record game fishes. International Game Fish Association, Florida, USA.
*Greenfield, D.W and J.E Thomerson (1997). Fishes of the continental waters of Belize. University Press of Florida, Florida. 311 p.
*Food and Agriculture Organization (1992). FAO yearbook 1990. Fishery statistics. Catches and landings. FAO Fish. Ser. (38). FAO Stat. Ser. 70:(105):647 p.
*Fish, M.P. and W.H. Mowbray (1970). Sounds of Western North Atlantic fishes. A reference file of biological underwater sounds. The Johns Hopkins Press, Baltimore.
*FAO (1997). Aquaculture production statistics 1986-1995. FAO Fish. Circ. 815, Rev. 9. 195 p.
*Eschmeyer, William N., ed. 1998. Catalog of Fishes. Special Publication of the Center for Biodiversity Research and Information, no. 1, vol 1-3. California Academy of Sciences. San Francisco, California, USA. 2905. ISBN 0-940228-47-5.
*Erdman, D.S. (1984). Exotic fishes in Puerto Rico. p. 162-176. In W.R. Courtney, Jr. and J.R. Stauffer, Jr. (eds.) Distribution, biology and management of exotic fishes. Johns Hopkins University Press, Baltimore, USA.
Comments: Full author list: Nelson, Joseph S., Edwin J. Crossman, Héctor Espinosa-Pérez, Lloyd T. Findley, Carter R. Gilbert, Robert N. Lea, and James D. Williams
Claro, Rodolfo, and Lynne R. Parenti / Claro, Rodolfo, Kenyon C. Lindeman, and L. R. Parenti, eds. 2001. Chapter 2: The Marine Ichthyofauna of Cuba. Ecology of the Marine Fishes of Cuba. Smithsonian Institution Press. Washington, DC, USA. 21-57. ISBN 1-56098-985-8.
*Claro, R. (1994). Características generales de la ictiofauna. p. 55-70. In R. Claro (ed.) Ecología de los peces marinos de Cuba. Instituto de Oceanología Academia de Ciencias de Cuba and Centro de Investigaciones de Quintana Roo.
*Böhlke, J.E. and C.C.G. Chaplin (1993). Fishes of the Bahamas and adjacent tropical waters. 2nd edition. University of Texas Press, Austin.
*Butsch, R.S. (1939). A list of Barbadian fishes. J. B.M.H.S. 7(1):17-31.
*Bussing, W.A. (1998). Peces de las aguas continentales de Costa Rica [Freshwater fishes of Costa Rica]. 2nd Ed. San José Costa Rica: Editorial de la Universidad de Costa Rica. 468 p.
*Banks, R. C., R. W. McDiarmid, A. L. Gardner, and W. C. Starnes 2003. Checklist of Vertebrates of the United States, the U.S. Territories, and Canada.
*Andrew K. Townesmith, ITIS Data Development Technician: Nelson et al. (2004) Occurrence values: A-F:CUM
==External links==
* [http://www.maine.gov/dmr/rm/eel.html The Maine Eel and Elver Fishery Maine Department of Marine Resources]
* [http://filaman.uni-kiel.de/Summary/SpeciesSummary.cfm?ID=35&genusname=Anguilla&speciesname=anguilla Fishbase entry for ''Anguilla anguilla'']
* [http://filaman.uni-kiel.de/Summary/SpeciesSummary.cfm?ID=296&genusname=Anguilla&speciesname=rostrata Fishbase entry for ''Anguilla rostrata'']
*[http://www.ices.dk/marineworld/eel.asp ICES report about eel stock collapse]
*[http://www.glasseel.com/ U.K Glass Eels — a large commercial firm's website, with history and fact pages]
*[http://web.archive.org/web/20001027080113/www.ecoscope.com/eelbase.htm Projekt eelBASE]
*[http://www.paling.nl]
[[Category:Ichthyology]]
[[Category:Anguilliformes]]
[[Category:Commercial fish]]
[[de:Geschichte der Erforschung der Aale]]