Leedsichthys
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''Leedsichthys'' is an extinct genus of pachycormid fish that lived in the oceans of the Middle to Late
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
.Liston, JJ (2004). An overview of the pachycormiform ''Leedsichthys''. In: Arratia G and Tintori A (eds) Mesozoic Fishes 3 - Systematics, Paleoenvironments and Biodiversity. Verlag Dr. Friedrich Pfeil, München, pp 379–390. It is the largest
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or h ...
, and amongst the largest fish known to have ever existed. The first remains of ''Leedsichthys'' were identified in the nineteenth century. Especially important were the finds by the British collector Alfred Nicholson Leeds, after whom the genus was named "Leeds' fish" in 1889. The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
is ''Leedsichthys problematicus''. ''Leedsichthys''
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s have been found in England, France, Germany and Chile. In 1999, based on the Chilean discoveries, a second species was named ''Leedsichthys notocetes'', but this was later shown to be indistinguishable from ''L. problematicus''. ''Leedsichthys'' fossils have been difficult to interpret because the skeletons were not completely made of bone. Large parts consisted of cartilage that did not fossilize. On several occasions the enigmatic large partial remains have been mistaken for stegosaurian
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
bones. As the vertebrae are among the parts that have not been preserved, it is hard to determine the total body length. Estimates have varied significantly. At the beginning of the twentieth century, a length of was seen as plausible, but by its end ''Leedsichthys'' was sometimes claimed to have been over long. Recent research has lowered this to about for the largest individuals. Skull bones have been found indicating that ''Leedsichthys'' had a large head with bosses on the skull roof. Fossilised bony fin rays show large elongated pectoral fins and a tall vertical tail fin. The gill arches were lined by gill rakers, equipped by a unique system of delicate bone plates, that filtered
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
from the sea water, the main food source. Along with its close pachycormid relatives '' Bonnerichthys'' and '' Rhinconichthys'', ''Leedsichthys'' is part of a lineage of large-sized
filter-feeder Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
s that swam the Mesozoic seas for over 100 million years, from the middle Jurassic until the end of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
. Pachycormids might represent an early branch of
Teleostei Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Te ...
, the group most modern bony fishes belong to; in that case ''Leedsichthys'' is the largest known teleost fish.


Discovery and naming

During the 1880s, the gentleman farmer Alfred Nicholson Leeds collected large fish fossils from loam pits near Peterborough, England. In May 1886 these were inspected by
John Whitaker Hulke John Whitaker Hulke FRCS FRS FGS (6 November 1830 – 19 February 1895) was a British surgeon, geologist and fossil collector. He was the son of a physician in Deal, who became a Huxleyite despite being deeply religious. Hulke became Huxley's ...
, who in 1887 partially reported them as the back plates of the stegosaurian ''
Omosaurus ''Omosaurus'' is a dubious genus of extinct crurotarsan reptile, possibly a phytosaur, from the Late Triassic (Carnian) of North Carolina. Only scant remains are known, which makes ''Omosaurus'' hard to classify. The type, and only species, ' ...
''. On 22 August 1888, the American dinosaur expert Professor
Othniel Charles Marsh Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of Paleontology in Yale College and President of the National Academy of Sciences. He was one of the preeminent scientists in the field of paleontology. Among ...
visited Leeds' farm at Eyebury and quickly concluded that the presumed dinosaurian armour in fact represented the skull bones of a giant fish. Within two weeks British fish expert
Arthur Smith Woodward Sir Arthur Smith Woodward, FRS (23 May 1864 – 2 September 1944) was an English palaeontologist, known as a world expert in fossil fish. He also described the Piltdown Man fossils, which were later determined to be fraudulent. He is not relate ...
examined the specimens and began to prepare a formal description published in 1889.Woodward, Smith, A (1889). Preliminary notes on some new and little-known British Jurassic fishes. Geological Magazine Decade 3 Volume 6: 448–455. In it he named the species ''Leedsichthys problematicus''. The generic name ''Leedsichthys'' means "Leeds' fish", from Greek ἰχθύς, ''ichthys'', "fish". The fossils found by Leeds gave the fish the specific epithet ''problematicus'' because the remains were so fragmented that they were extremely hard to recognize and interpret. After a second publication in 1889, objections were raised against the perceived "
barbaric A barbarian (or savage) is someone who is perceived to be either uncivilized or primitive. The designation is usually applied as a generalization based on a popular stereotype; barbarians can be members of any nation judged by some to be less ...
" nature of the generic name, which simply attached a non-Latinised British family name to a Classical Greek word. Woodward therefore in 1890 changed the genus name to ''Leedsia'', resulting in a ''Leedsia problematica''. However, by modern standards this is a non-valid junior synonym. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
specimen, BMNH P.6921, had been found in a layer of the Oxford Clay Formation dating from the
Callovian In the geologic timescale, the Callovian is an age and stage in the Middle Jurassic, lasting between 166.1 ± 4.0 Ma (million years ago) and 163.5 ± 4.0 Ma. It is the last stage of the Middle Jurassic, following the Bathonian and preceding the ...
, about 165 million years old. It consists of 1133 disarticulated elements of the skeleton, mostly fin ray fragments, probably of a single individual. Another specimen, BMNH P.6922, contains additional probable fragmentary remains of ''Leedsichthys''. Woodward also identified a specimen previously acquired from the French collector Tesson, who had in 1857 found them in the Falaises des Vaches Noires of
Normandy Normandy (; french: link=no, Normandie ; nrf, Normaundie, Nouormandie ; from Old French , plural of ''Normant'', originally from the word for "northman" in several Scandinavian languages) is a geographical and cultural region in Northwestern ...
, BMNH 32581, as the gill rakers of ''Leedsichthys''. Another specimen bought in 1875 from the collection of William Cunnington, BMNH 46355, he failed to recognise.Liston, J.J., 2010, "The occurrence of the Middle Jurassic pachycormid fish ''Leedsichthys''", ''Oryctos'' 9: 1-36 Leeds continued to collect ''Leedsichthys'' fossils that subsequently were acquired by British musea. In March 1898, Leeds reported to have discovered a tail which he on 17 March 1899 sold for £25 to the
British Museum of Natural History The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum ...
, which exhibited it as specimen BMNH P.10000; a new inventory number range was begun for the occasion. Already in July 1898, the front of probably the same animal had been bought, BMNH P.11823. On 22 July 1905 specimen BMNH P.10156 was acquired, a gill basket. In January 1915 Leeds sold specimens GLAHM V3362, a pectoral fin, and GLAHM V3363, the remainder of the same skeleton with 904 elements, to the Hunterian Museum of
Glasgow Glasgow ( ; sco, Glesca or ; gd, Glaschu ) is the most populous city in Scotland and the fourth-most populous city in the United Kingdom, as well as being the 27th largest city by population in Europe. In 2020, it had an estimated popul ...
. Leeds had a rival, the collector Henry Keeping, who in 1899 tricked pit workers into selling dorsal fin rays by misinforming them that Leeds had lost interest in such finds. Keeping again sold these to the
University of Cambridge The University of Cambridge is a public collegiate research university in Cambridge, England. Founded in 1209 and granted a royal charter by Henry III in 1231, Cambridge is the world's third oldest surviving university and one of its most pr ...
where they were catalogued as specimen CAMSM J.46873. In September 1901, they were examined by the German palaeontologist Friedrich von Huene, who identified them as tail spikes, ''Schwanzstacheln'', of ''Omosaurus'', the second time ''Leedsichthys'' remains were mistaken for stegosaurian bones; Leeds himself was able to disabuse von Huene the same year. In 2001, students at the Dogsthorpe Star Pit discovered a major new British specimen that they nicknamed "Ariston" after a 1991 commercial for the
Indesit Indesit Company (; ) was an Italian company based in Fabriano, Ancona. It was one of the leading European manufacturers and distributors of major domestic appliances (washing machines, dryers, dishwashers, fridges, freezers, cookers, hoods, ov ...
Ariston
washing machine A washing machine (laundry machine, clothes washer, washer, or simply wash) is a home appliance used to wash laundry. The term is mostly applied to machines that use water as opposed to dry cleaning (which uses alternative cleaning fluids and ...
that claimed it went "on and on and on" — likewise the bones of ''Leedsichthys'' seemed to endlessly continue into the face of the loam pit. From 2002 until 2004 "Ariston" or specimen PETMG F174 was excavated by a team headed by Jeff Liston; to uncover the remains it was necessary to remove ten thousand
tonne The tonne ( or ; symbol: t) is a unit of mass equal to 1000  kilograms. It is a non-SI unit accepted for use with SI. It is also referred to as a metric ton to distinguish it from the non-metric units of the short ton ( United State ...
s of loam forming an overburden of thickness. The find generated considerable media attention, inspiring an episode of the
BBC #REDIRECT BBC #REDIRECT BBC Here i going to introduce about the best teacher of my life b BALAJI sir. He is the precious gift that I got befor 2yrs . How has helped and thought all the concept and made my success in the 10th board exam. ...
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''
Sea Monsters Sea monsters are beings from folklore believed to dwell in the sea and often imagined to be of immense size. Marine monsters can take many forms, including sea dragons, sea serpents, or tentacled beasts. They can be slimy and scaly and are oft ...
'' series, "The Second Most Deadly Sea", and a
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documentary titled ''The Big Monster Dig'', both containing computer-generated animated reconstructions of ''Leedsichthys''. Liston subsequently dedicated a dissertation and a series of articles to ''Leedsichthys'', providing the first extensive modern osteology of the animal.Liston, J.J., 2007, ''A Fish Fit For Ozymandias?: The Ecology, Growth and Osteology of'' Leedsichthys ''(Pachycormidae, Actinopterygii)'', Unpublished PhD Thesis. 464 pp. Faculty of BioMedical & LifeSciences, University of Glasgow, Scotland Apart from the British discoveries, finds of a more fragmentary nature continued to be made in Normandy, France. In July 1982, Germany became an important source of ''Leedsichthys'' fossils when two groups of amateur palaeontologists, unaware of each other's activities, began to dig up the same skeleton at Wallücke. Remarkably, parts of it were again incorrectly identified as stegosaurian material, of '' Lexovisaurus''. From 1973 onwards, fragmentary ''Leedsichthys'' fossils were uncovered in
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...
. In March 1994, a more complete specimen was found, SMNK 2573 PAL. In 1999 the Chilean finds were named as a second species, ''Leedsichthys notocetes'', the "Southern Sea Monster".Martill, D.M., Frey, E., Caceras, R.P. & Diaz, G.C., 1999, "The giant pachycormid ''Leedsichthys'' (Actinopterygii) in the southern hemisphere: further evidence for a Jurassic Atlanto-Pacific marine faunal province", ''Neues Jahrbuch für Geologie und Paläontologie'', Monatshefte 1999: 243-256 However, Liston later concluded that the presumed distinguishing traits of this species, depressions on the gill rakers, were artefacts caused by erosion; ''Leedsichthys notocetes'' would be a junior synonym of ''Leedsichthys problematicus''.Liston, J.J., 2013, "The plasticity of gill raker characteristics in suspension feeders: Implications for Pachycormiformes", In: G. Arratia, H.-P. Schultze & M. V. H. Wilson (eds.) ''Mesozoic Fishes 5 – Global Diversity and Evolution'' pp 121-143, Verlag Dr. Friedrich Pfeil, München


Fossil range

The fossil remains of ''Leedsichthys'' have been found in the
Callovian In the geologic timescale, the Callovian is an age and stage in the Middle Jurassic, lasting between 166.1 ± 4.0 Ma (million years ago) and 163.5 ± 4.0 Ma. It is the last stage of the Middle Jurassic, following the Bathonian and preceding the ...
of
England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...
and northern
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, the Oxfordian of
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...
, and the Callovian and upper
Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxford ...
of France.Liston, JJ (2008a). ''Leedsichthys'' des Vaches Noires ... au peigne fin (translation by M-C Buchy) L’Écho des Falaises (=Ech.des Fal.) No.12: 41–49, 2008 ISSN 1253-6946. These occurrences span a temporal range of at least five million years. A complete and isolated gill raker from the
Vaca Muerta The Vaca Muerta Formation, commonly known as Vaca Muerta (Spanish for ''Dead Cow''), is a geologic formation of Late Jurassic to Early Cretaceous age, located in the Neuquén Basin in northern Patagonia, Argentina. It is well known as the host ro ...
formation of
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
(MOZ-Pv 1788), has been assigned to the genus and dates to the early Tithonian.


Description

Although the remains of over seventy individuals have been found, most of them are partial and fragmentary. The skeleton of ''Leedsichthys'' is thus only imperfectly known. This is largely caused by the fact that many skeletal elements, including the front of the skull and the vertebral centra, did not ossify but remained cartilage. Furthermore, those that did ossify were gradually hollowed out during the lifetime of the animal by resorption of the inner bone tissue. In the fossil phase, compression flattened and cracked these hollow structures, making it extraordinarily difficult to identify them or determine their original form. The head was probably relatively large and wide but still elongated. The snout is completely unknown.
Frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, pa ...
s are absent. The skull roof is rather robust with bosses on the parietals, continuing sideways over the dermopterotica, and the postparietals. The parietals have a notch on the front midline. A dermosphenoticum is present above the eye socket. The jaws are toothless. Behind the jaw joint a robust hyomandibula is present. The gill basket rests on paired hypohyalia. At least the first two gill arches have ossified hypobranchialia, the lower parts of the gill arch; a third hypobranchiale was likely present. The hypobranchials are attached at their lower ends at an angle of 21,5° via a functional joint that possibly served to increase the gape of the mouth, to about two feet. All five gill arches have ossified ceratobranchialia with a triangular cross-section, the middle sections of the arches. The hypobranchials are fused with their ceratobranchials. The fifth gill arch is fused with the front parts of the basket. Higher epibranchialia and pharyngobranchialia are present but poorly known. The fourth arches are supported by a midline fourth basibranchiale. An ossified operculum is present.Liston, J.J., 2008, "A review of the characters of the edentulous pachycormiforms ''Leedsichthys'', ''Asthenocormus'' and ''Martillichthys'' nov. gen.", In: ''Mesozoic Fishes 4 Homology and Phylogeny'', G. Arratia, H.-P. Schultze & M. V. H. Wilson (eds.): pp. 181–198, 10 figs., 1 tab. © 2008 by Verlag Dr. Friedrich Pfeil, München, Germany – The gill arches are equipped with rows of parallel 3-to-12-centimetre-long (1.2-to-4.7-inch-long)
gill rakers Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of t ...
, in life probably attached to the ceratobranchials via soft tissue. On the top of each raker one or two rows of dozens of low "teeth" are present. When there are two rows, they are placed on the edges of the upper surface and separated by a deep trough, itself separated from an internal hollow space by a transverse septum. The teeth or " fimbriations" are obliquely directed towards the front and the top. They are grooved at their sides, the striations continuing over the sides of the raker. Detailed study of exquisitely preserved French specimens revealed to Liston that these teeth were, again via soft tissue, each attached to delicate 2-millimetre-long (0.08-inch-long) bony plates, structures that had never before been observed among living or extinct fishes. An earlier hypothesis that the striations would function as sockets for sharp "needle teeth", as with the
basking shark The basking shark (''Cetorhinus maximus'') is the second-largest living shark and fish, after the whale shark, and one of three plankton-eating shark species, along with the whale shark and megamouth shark. Adults typically reach in leng ...
, was hereby refuted. The rakers served to filter
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
, the main food supply of ''Leedsichthys'', from the sea water. Large parts of the ''Leedsichthys'' fossils consist of bony finrays. ''Leedsichthys'' has two pectoral fins that probably were located rather low on the body. They are large, very elongated — about five times longer than wide — and scythe-like, with a sudden kink at the lower end, curving 10° to the rear. Also a dorsal fin is present, although its position is unknown. Pelvic fins at the belly are lacking; also a pelvic plate is absent. However, there are indications for a small triangular anal fin. The vertical tail fin is very large and symmetrical with paired upper and lower lobes; there is a smaller lobe in the middle protruding between them. The rays are unsegmented lepidotrichia, resulting in a rather stiff structure. They are bifurcated at up to three splitting points along their length, so a proximally single ray may have eight distal ends. A row of bony supraneuralia is present behind the head, at each side of the vertebral column. Uroneuralia at the tail are unknown. No bony scales are present.


Size

''Leedsichthys'' is the largest known member of the Osteichthyes or bony fishes. The largest extant non-tetrapodomorph bony fish is the
ocean sunfish The ocean sunfish or common mola (''Mola mola'') is one of the largest bony fish in the world. It was misidentified as the heaviest bony fish, which was actually a different species, ''Mola alexandrini''. Adults typically weigh between . The spe ...
, ''Mola mola'', being with a weight of up to two tonnes an
order of magnitude An order of magnitude is an approximation of the logarithm of a value relative to some contextually understood reference value, usually 10, interpreted as the base of the logarithm and the representative of values of magnitude one. Logarithmic di ...
smaller than ''Leedsichthys''. The extant giant oarfish might rival ''Leedsichthys'' in length but is much thinner. The lack of a preserved vertebral column has made it difficult to estimate the exact length of ''Leedsichthys''.
Arthur Smith Woodward Sir Arthur Smith Woodward, FRS (23 May 1864 – 2 September 1944) was an English palaeontologist, known as a world expert in fossil fish. He also described the Piltdown Man fossils, which were later determined to be fraudulent. He is not relate ...
, who described the
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...
in 1889, estimated specimen BMNH P.10000 to be of an around nine metre long individual, by comparing this tail of ''Leedsichthys'', having a preserved height of , with another pachycormid, '' Hypsocormus''. The length of ''Leedsichthys'' was not historically the subject of much attention, the only reference to it being made by Woodward himself when he in 1937 indicated it again as on the museum label of BMNH P.10000. However, in 1986, David Martill compared the bones of ''Leedsichthys'' to a pachycormid that he had recently discovered, '' Asthenocormus''.Martill, DM (1986). The world's largest fish. ''Geology Today'' March–April: 61–63. The unusual proportions of that specimen gave a wide range of possible sizes. Some were as low as , but extrapolating from the gill basket resulted in an estimated length of for ''Leedsichthys'' specimen NHM P.10156 (the earlier BMNH P.10156). Martill considered the higher estimate as a plausible size of the largest individuals. Subsequently, a length of thirty metres (hundred feet) was often mentioned in popular science publications, sometimes one as high as . Liston in his studies concluded to much lower estimates. Documentation of historical finds and the excavation of "Ariston", the most complete specimen ever from the Star Pit near
Whittlesey Whittlesey (also Whittlesea) is a market town and civil parish in the Fenland district of Cambridgeshire, England. Whittlesey is east of Peterborough. The population of the parish was 16,058 at the 2011 Census. History and architecture W ...
,
Peterborough Peterborough () is a cathedral city in Cambridgeshire, east of England. It is the largest part of the City of Peterborough unitary authority district (which covers a larger area than Peterborough itself). It was part of Northamptonshire until ...
, support Woodward's figures of between . With "Ariston" the pectoral fins are apart, indicating a narrow body of no excessive size, even though it was initially thought to have been long. In 2007 Liston stated that most specimens indicated lengths between . A linear extrapolation from the gill basket would be flawed because the gills grow disproportionally in size, having to increase their surface allometrically to ensure the
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...
supply of a body increasing in volume to the third power. The
growth ring Dendrochronology (or tree-ring dating) is the scientific method of dating tree rings (also called growth rings) to the exact year they were formed. As well as dating them, this can give data for dendroclimatology, the study of climate and atmo ...
structures within the remains of ''Leedsichthys'' have indicated that it would have taken 21 to 25 years to reach these lengths, and isolated elements from other specimens showed that a maximum size of just over is not unreasonable. In 2013 a new study by Liston found most specimens to be between 17 and 31 years old. Specimen GLAHM V3363, "Big Meg", could have been between . The exception was NHM P.10156. Its gill basket, with a preserved width of and height of , indicated a body length of and an age of 45 years.Liston, J., Newbrey, M., Challands, T., and Adams, C., 2013, "Growth, age and size of the Jurassic pachycormid ''Leedsichthys problematicus'' (Osteichthyes: Actinopterygii) in: Arratia, G., Schultze, H. and Wilson, M. (eds.) ''Mesozoic Fishes 5 – Global Diversity and Evolution''. Verlag Dr. Friedrich Pfeil, München, Germany, pp. 145–175 The largest individual would have weighed up to .


Phylogeny

Woodward initially assigned ''Leedsichthys'' to the Acipenseroidea, considering it related to the sturgeon, having the large gill rakers and branching finrays in common. In 1905, he changed this to the
Pachycormidae Pachycormiformes is an extinct order of marine ray-finned fish known from the Early Jurassic to the end of the Cretaceous. It only includes a single family, Pachycormidae. They were characterized by having serrated pectoral fins (though more rece ...
. The Pachycormidae have a somewhat uncertain position. Often they are considered very basal
Teleostei Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Te ...
 — if so, ''Leedsichthys'' would be the largest known teleost — others see them as members of a
Pachycormiformes Pachycormiformes is an extinct order of marine ray-finned fish known from the Early Jurassic to the end of the Cretaceous. It only includes a single family, Pachycormidae. They were characterized by having serrated pectoral fins (though more rece ...
forming the sister group of the Teleostei, and sometimes they are seen as even more basal
Amiiformes The Amiiformes order of fish has only one extant species, the bowfin (''Amia calva''). These Amiiformes are found in the freshwater systems of North America, in the United States and parts of southern Canada. They live in freshwater streams, riv ...
. In the latter case the extant bowfin, ''Amia calva'', would be the closest living relative of ''Leedsichthys''. Within the Pachycormidae, a
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analysis found ''Leedsichthys'' to be the
sister species In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of ''Asthenocormus'', their clade being the sister group of '' Martillichthys''. This
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
after Friedman ''et al.'' shows a possible position of ''Leedsichtys'' in the evolutionary tree.


Paleobiology

Like the largest fish today, the
whale sharks The whale shark (''Rhincodon typus'') is a slow-moving, filter-feeding carpet shark and the largest known extant fish species. The largest confirmed individual had a length of .McClain CR, Balk MA, Benfield MC, Branch TA, Chen C, Cosgrove J, ...
and basking sharks, ''Leedsichthys problematicus'' derived its nutrition as a suspension feeder, using an array of specialised
gill raker Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of the ...
s lining its gill basket to extract
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, small animals, from the water passing through its mouth and across its gills. It is less clear whether also phytoplankton, algae, were part of the diet. ''Leedsichthys'' could have been a ram feeder, making the water pass through its gills by swimming, but could also have actively pumped the water through the gill basket. In 2010, Liston suggested that fossilised furrows discovered in ancient sea floors in Switzerland and attributed to the activity of
plesiosaurs The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared i ...
, had in fact been made by ''Leedsichthys'' spouting water through its mouth to disturb and eat the
benthos Benthos (), also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone. Much is still uncertain about the life cycle of ''Leedsichthys''. Liston's 2013 study suggested a slow, nearly linear, growth. A French study in 1993 of its bone structure concluded however, that the metabolism was rather high. Also problematic is how ''Leedsichthys'' could increase its size quickly during the first year of its life. Teleostei typically lay relatively small eggs and this has been seen as an obstacle for them attaining giant sizes. In 1986, Martill reported the presence of a tooth of the marine crocodile ''
Metriorhynchus ''Metriorhynchus'' is an extinct genus of marine crocodyliform that lived in the oceans during the Late Jurassic. The type species, ''M. brevirostris'' was named in 1829 as a species of ''Steneosaurus'' before being named as a separate genus by ...
'' in a bone of ''Leedsichthys''. The bone would have healed, a sign that the about 3-metre-long (10-foot-long) ''Metriorhynchus'' was actively hunting the much larger fish. However, in 2007 Liston concluded the bone tissue had not in fact healed and that this was probably a case of scavenging. An
apex predator An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic lev ...
of the
Oxford Clay The Oxford Clay (or Oxford Clay Formation) is a Jurassic marine sedimentary rock formation underlying much of southeast England, from as far west as Dorset Dorset ( ; archaically: Dorsetshire , ) is a county in South West England on the ...
seas large enough to attack ''Leedsichthys'' was the
pliosaurid Pliosauridae is a family of plesiosaurian marine reptiles from the Latest Triassic to the early Late Cretaceous (Rhaetian to Turonian stages) of Australia, Europe, North America and South America. The family is more inclusive than the archetypal ...
'' Liopleurodon''. In 1999 Martill suggested that a
climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
at the end of the Callovian led to the extinction of ''Leedsichthys'' in the northern seas, the southern Ocean offering a last refuge during the Oxfordian. However, in 2010 Liston pointed out that ''Leedsichthys'' during the later Kimmeridgian was still present in the north, as testified by Normandian finds. Liston did nevertheless consider in 2007 that the lack of any vertebrate suspension feeders as large as prior to the Callovian stage of the Mesozoicum might indicate that the Callovian had seen a marked change in productivity as regarded
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
populations. Indeed, further studies supported this, viewing ''Leedsichthys'' as the beginning of a long line of large (> in length) pachycormid suspension feeders that continued to flourish well into the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
, such as '' Bonnerichthys'' and '' Rhinconichthys'', and emphasising the
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
ary paths taken by pachycormids and
baleen whales Baleen whales (systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea (whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in the ...
. Recent studies have uncovered some estimations regarding metabolic rate and speed for ''Leedsichthys''. Using data from living teleost fish as a comparison, scientists discovered that ''Leedsichthys'' could have cruised along at potential speeds of while still maintaining oxygenation of its body tissues.Humberto G. Ferrón, Borja Holgado, Jeffrey J. Liston, Carlos Martínez‐Pérez & Héctor Botella, 2018, "Assessing metabolic constraints on the maximum body size of actinopterygians: locomotion energetics of ''Leedsichthys problematicus'' (Actinopterygii, Pachycormiformes)", ''Palaeontology'' 61(5): 775-783


Footnotes


External links


For more on the Star Pit dig of 2002-2003 look here
and also here where the dig was featured i


For a more accurate reconstruction of ''Leedsichthys'' visit Paleocreations.
The artist went on to produce the most accurate reconstruction of ''Leedsichthys'' to date, which was used on 'Fossil Detectives' (see above).
Article on ''Leedsichthys''
by
Darren Naish Darren William Naish is a British vertebrate palaeontologist, author and science communicator. As a researcher, he is best known for his work describing and reevaluating dinosaurs and other Mesozoic reptiles, including '' Eotyrannus'', '' Xenop ...
.
Profile of ''Leedsichthys problematicus'' from the BBC series Sea Monsters
National Geographic News. Published October 1, 2003.

estimated 22 meters = long.
Picture of ''Leedsichthys problematicus''

Article on ''Leedsichthys'' from BBC news
{{Taxonbar, from=Q131114 Pachycormiformes Prehistoric ray-finned fish genera Callovian genus first appearances Oxfordian life Kimmeridgian life Jurassic fish of Europe Jurassic France Fossils of France Jurassic Germany Fossils of Germany Jurassic United Kingdom Fossils of England Prehistoric fish of South America Jurassic Chile Fossils of Chile Fossil taxa described in 1889 Taxa named by Arthur Smith Woodward Jurassic Argentina