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Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct
marine reptile Marine reptiles are reptiles which have become secondarily adapted for an aquatic or semiaquatic life in a marine environment. The earliest marine reptile mesosaurus (not to be confused with mosasaurus), arose in the Permian period during ...
s. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. ...
in 1842, although the term is now used more for the parent
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
of the Ichthyosauria). Ichthyosaurs thrived during much of the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
era; based on
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 ...
evidence, they first appeared around 250 million years ago ( Ma) and at least one species survived until about 90 million years ago, 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'', t ...
. During the
Early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which is a ...
epoch In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured. The moment of epoch is usually decided by ...
, ichthyosaurs and other
ichthyosauromorphs The Ichthyosauromorpha are an extinct clade of marine reptiles consisting of the Ichthyosauriformes and the Hupehsuchia, living during the Mesozoic. The node clade Ichthyosauromorpha was first defined by Ryosuke Motani ''et al.'' in 2014 as t ...
evolved from a group of unidentified land reptiles that returned to the sea, in a development similar to how the mammalian land-dwelling ancestors of modern-day
dolphin A dolphin is an aquatic mammal within the infraorder Cetacea. Dolphin species belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontoporiidae ( ...
s and whales returned to the sea millions of years later, which they gradually came to resemble in a case of
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 ...
. Ichthyosaurs were particularly abundant in the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch. ...
and
Early Jurassic The Early Jurassic Epoch (in chronostratigraphy corresponding to the Lower Jurassic Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-Jurassic extinction event, 201.3 Ma&nb ...
periods, until they were replaced as the top aquatic predators by another marine reptilian group, the
Plesiosauria 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 ...
, in the later Jurassic and
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous ( chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pr ...
, though previous views of ichthyosaur decline during this period are probably overstated. Ichthyosaurs diversity declined due to environmental volatility caused by climatic upheavals in the early Late Cretaceous, becoming extinct around the Cenomanian-Turonian boundary approximately 90 million years ago. Science became aware of the existence of ichthyosaurs during the early nineteenth century, when the first complete skeletons were found in England. In 1834, the order Ichthyosauria was named. Later that century, many excellently preserved ichthyosaur fossils were discovered in Germany, including soft-tissue remains. Since the late twentieth century, there has been a revived interest in the group, leading to an increased number of named ichthyosaurs from all continents, with over fifty valid genera being now known. Ichthyosaur species varied from in length. Ichthyosaurs resembled both modern fish and dolphins. Their limbs had been fully transformed into flippers, which sometimes contained a very large number of digits and phalanges. At least some species possessed a dorsal fin. Their heads were pointed, and the jaws often were equipped with conical teeth to catch smaller prey. Some species had larger, bladed teeth to attack large animals. The eyes were very large, for deep diving. The neck was short, and later species had a rather stiff trunk. These also had a more vertical tail fin, used for a powerful propulsive stroke. The vertebral column, made of simplified disc-like vertebrae, continued into the lower lobe of the tail fin. Ichthyosaurs were air-breathing, warm-blooded, and bore live young. They may have had a layer of
blubber Blubber is a thick layer of vascularized adipose tissue under the skin of all cetaceans, pinnipeds, penguins, and sirenians. Description Lipid-rich, collagen fiber-laced blubber comprises the hypodermis and covers the whole body, excep ...
for insulation.


History of discoveries


Early finds

The first known illustrations of ichthyosaur bones, vertebrae, and limb elements were published by the Welshman
Edward Lhuyd Edward Lhuyd FRS (; occasionally written Llwyd in line with modern Welsh orthography, 1660 – 30 June 1709) was a Welsh naturalist, botanist, linguist, geographer and antiquary. He is also named in a Latinate form as Eduardus Luidius. Lif ...
in his ''Lithophylacii Brittannici Ichnographia'' of 1699. Lhuyd thought that they represented fish remains. In 1708, the Swiss naturalist
Johann Jakob Scheuchzer Johann Jakob Scheuchzer (2 August 1672 – 23 June 1733) was a Swiss scholar born at Zürich. Herbarium deluvianum Zürich, Zwingli-Platz ( Grossmünster) : Former home of Konrad von Mure († 1280) and the house, where Johann Jakob Scheuc ...
described two ichthyosaur vertebrae assuming they belonged to a man drowned in the Universal Deluge. In 1766, an ichthyosaur jaw with teeth was found at
Weston Weston may refer to: Places Australia * Weston, Australian Capital Territory, a suburb of Canberra * Weston, New South Wales * Weston Creek, a residential district of Canberra * Weston Park, Canberra, a park Canada * Weston, Nova Scotia ...
near
Bath Bath may refer to: * Bathing, immersion in a fluid ** Bathtub, a large open container for water, in which a person may wash their body ** Public bathing, a public place where people bathe * Thermae, ancient Roman public bathing facilities Pla ...
. In 1783, this piece was exhibited by the
Society for Promoting Natural History A society is a group of individuals involved in persistent social interaction, or a large social group sharing the same spatial or social territory, typically subject to the same political authority and dominant cultural expectations. Societ ...
as those of a crocodilian. In 1779, ichthyosaur bones were illustrated in
John Walcott John is a common English name and surname: * John (given name) * John (surname) John may also refer to: New Testament Works * Gospel of John, a title often shortened to John * First Epistle of John, often shortened to 1 John * Second E ...
's ''Descriptions and Figures of Petrifications''. Towards the end of the eighteenth century, British fossil collections quickly increased in size. Those of the naturalists
Ashton Lever Sir Ashton Lever FRS (5 March 1729 – 28 January 1788) was an English collector of natural objects, in particular the Leverian collection.John Hunter were acquired in their totality by museums; later, it was established that they contained dozens of ichthyosaur bones and teeth. The bones had typically been labelled as belonging to fish, dolphins, or crocodiles; the teeth had been seen as those of sea lions. The demand by collectors led to more intense commercial digging activities. In the early nineteenth century, this resulted in the discovery of more complete skeletons. In 1804, Edward Donovan at
St Donats St Donats ( cy, Sain Dunwyd) is a village and community in the Vale of Glamorgan in south Wales, located just west of the small town of Llantwit Major. The community includes the village of Marcross and the hamlets of Monknash and East and We ...
uncovered a ichthyosaur specimen containing a jaw, vertebrae, ribs, and a shoulder girdle. It was considered to be a giant lizard. In October 1805, a newspaper article reported the find of two additional skeletons, one discovered at Weston by Jacob Wilkinson, the other, at the same village, by Reverend Peter Hawker. In 1807, the last specimen was described by the latter's cousin, Joseph Hawker. This specimen thus gained some fame among geologists as 'Hawker's Crocodile'. In 1810, near
Stratford-upon-Avon Stratford-upon-Avon (), commonly known as just Stratford, is a market town and civil parish in the Stratford-on-Avon district, in the county of Warwickshire, in the West Midlands region of England. It is situated on the River Avon, north-we ...
, an ichthyosaur jaw was found that was combined with
plesiosaur The Plesiosauria (; Ancient Greek, Greek: πλησίος, ''plesios'', meaning "near to" and Sauria, ''sauros'', meaning "lizard") or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauro ...
bones to obtain a more complete specimen, indicating that the distinctive nature of ichthyosaurs was not yet understood, awaiting the discovery of far better fossils.


The first complete skeletons

In 1811, in
Lyme Regis Lyme Regis is a town in west Dorset, England, west of Dorchester and east of Exeter. Sometimes dubbed the "Pearl of Dorset", it lies by the English Channel at the Dorset–Devon border. It has noted fossils in cliffs and beaches on the Herita ...
, along what is now called the
Jurassic Coast The Jurassic Coast is a World Heritage Site on the English Channel coast of southern England. It stretches from Exmouth in East Devon to Studland Bay in Dorset, a distance of about , and was inscribed on the World Heritage List in mid-Decemb ...
of
Dorset Dorset ( ; archaically: Dorsetshire , ) is a county in South West England on the English Channel coast. The ceremonial county comprises the unitary authority areas of Bournemouth, Christchurch and Poole and Dorset. Covering an area of , ...
, the first complete ichthyosaur skull was found by Joseph Anning, the brother of
Mary Anning Mary Anning (21 May 1799 – 9 March 1847) was an English fossil collector, dealer, and palaeontologist who became known around the world for the discoveries she made in Jurassic marine fossil beds in the cliffs along the English Channe ...
, who in 1812 while still a young girl, secured the torso of the same specimen. Their mother, Molly Anning, sold the combined piece to squire Henry Henley for £23. Henley lent the fossil to the London Museum of Natural History of William Bullock. When this museum was closed, the
British Museum The British Museum is a public museum dedicated to human history, art and culture located in the Bloomsbury area of London. Its permanent collection of eight million works is among the largest and most comprehensive in existence. It docu ...
bought the fossil for a price of £47.5s; it still belongs to the collection of the now independent
Natural History Museum A natural history museum or museum of natural history is a scientific institution with natural history scientific collection, collections that include current and historical records of animals, plants, Fungus, fungi, ecosystems, geology, paleon ...
and has the inventory number BMNH R.1158. It has been identified as a specimen of '' Temnodontosaurus platyodon''. In 1814, the Annings' specimen was described by Professor
Everard Home Sir Everard Home, 1st Baronet, FRS (6 May 1756, in Kingston upon Hull – 31 August 1832, in London) was a British surgeon. Home was born in Kingston-upon-Hull and educated at Westminster School. He gained a scholarship to Trinity College, C ...
, in the first scientific publication dedicated to an ichthyosaur. Intrigued by the strange animal, Home tried to locate additional specimens in existing collections. In 1816, he described ichthyosaur fossils owned by
William Buckland William Buckland DD, FRS (12 March 1784 – 14 August 1856) was an English theologian who became Dean of Westminster. He was also a geologist and palaeontologist. Buckland wrote the first full account of a fossil dinosaur, which he named ...
and James Johnson. In 1818, Home published data obtained by corresponding with naturalists all over Britain. In 1819, he wrote two articles about specimens found by Henry Thomas De la Beche and Thomas James Birch. A last publication of 1820 was dedicated to a discovery by Birch at Lyme Regis. The series of articles by Home covered the entire anatomy of ichthyosaurs, but highlighted details only; a systematic description was still lacking. Home felt very uncertain how the animal should be classified. Though most individual skeletal elements looked very reptilian, the anatomy as a whole resembled that of a fish, so he initially assigned the creature to the fishes, as seemed to be confirmed by the flat shape of the vertebrae. At the same time, he considered it a transitional form in-between fishes and crocodiles, not in an evolutionary sense, but as regarded its place in the '' scala naturae'', the "Chain of Being" hierarchically connecting all living creatures. In 1818, Home noted some coincidental similarities between the coracoid of ichthyosaurians and the sternum of the
platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or monoty ...
. This induced him to emphasize its status as a transitional form, combining, like the platypus, traits of several larger groups. In 1819, he considered it a form between
newt A newt is a salamander in the subfamily Pleurodelinae. The terrestrial juvenile phase is called an eft. Unlike other members of the family Salamandridae, newts are semiaquatic, alternating between aquatic and terrestrial habitats. Not all aq ...
s, like
Olm The olm or proteus (''Proteus anguinus'') is an aquatic salamander in the family Proteidae, the only exclusively cave-dwelling chordate species found in Europe. In contrast to most amphibians, it is entirely aquatic, eating, sleeping, and br ...
, and lizards; he now gave a formal generic name: ''Proteo-Saurus''. However, in 1817, Karl Dietrich Eberhard Koenig had already referred to the animal as ''Ichthyosaurus'', "fish saurian" from Greek ἰχθύς, ''ichthys'', "fish". This name at the time was an invalid ''
nomen nudum In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate descr ...
'' and was only published by Koenig in 1825, but was adopted by De la Beche in 1819 in a lecture where he named three ''Ichthyosaurus'' species. This text would only be published in 1822, just after De la Beche's friend William Conybeare published a description of these species, together with a fourth one. 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 specimen( ...
was ''Ichthyosaurus communis'', based on a now lost skeleton. Conybeare considered that ''Ichthyosaurus'' had priority relative to ''Proteosaurus''. Although this is incorrect by present standards, the latter name became a "forgotten" ''
nomen oblitum In zoological nomenclature, a ''nomen oblitum'' (plural: ''nomina oblita''; Latin for "forgotten name") is a disused scientific name which has been declared to be obsolete (figuratively 'forgotten') in favour of another 'protected' name. In its p ...
''. In 1821, De la Beche and Conybeare provided the first systematic description of ichthyosaurs, comparing them to another newly identified marine reptile group, the
Plesiosauria 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 ...
. Much of this description reflected the insights of their friend, the anatomist Joseph Pentland. In 1835, the order Ichthyosauria was named by
Henri Marie Ducrotay de Blainville Henri Marie Ducrotay de Blainville (; 12 September 1777 – 1 May 1850) was a French zoologist and anatomist. Life Blainville was born at Arques, near Dieppe. As a young man he went to Paris to study art, but ultimately devoted himself to natu ...
. In 1840,
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. ...
named an order Ichthyopterygia as an alternative concept.


Popularisation during the 19th century

The discovery of a hitherto unsuspected extinct group of large marine reptiles generated much publicity, capturing the imagination of both scientists and the public at large. People were fascinated by the strange build of the animals, especially the large scleral rings in the eye sockets, of which it was sometimes erroneously assumed these would have been visible on the living animal. Their bizarre form induced a feeling of alienation, allowing people to realise the immense span of time passed since the era in which the ichthyosaur swam the oceans. Not all were convinced that ichthyosaurs had gone extinct: Reverend George Young found a skeleton in 1819 at
Whitby Whitby is a seaside town, port and civil parish in the Scarborough borough of North Yorkshire, England. Situated on the east coast of Yorkshire at the mouth of the River Esk, Whitby has a maritime, mineral and tourist heritage. Its East Clif ...
; in his 1821 description, he expressed the hope that living specimens could still be found. Geologist
Charles Lyell Sir Charles Lyell, 1st Baronet, (14 November 1797 – 22 February 1875) was a Scottish geologist who demonstrated the power of known natural causes in explaining the earth's history. He is best known as the author of ''Principles of Geolo ...
to the contrary, assumed that the Earth was eternal so that in the course of time the ichthyosaur might likely reappear, a possibility lampooned in a famous caricature by De la Beche. Public awareness was increased by the works of the eccentric collector Thomas Hawkins, a
pre-Adamite The pre-Adamite hypothesis or pre-Adamism is the theological belief that humans (or intelligent yet non-human creatures) existed before the biblical character Adam. Pre-Adamism is therefore distinct from the conventional Abrahamic belief that Ada ...
believing that ichthyosaurs were monstrous creations by the devil: ''Memoirs of Ichthyosauri and Plesiosauri'' of 1834 and ''The Book of the Great Sea-Dragons'' of 1840. The first work was illustrated by
mezzotint Mezzotint is a monochrome printmaking process of the '' intaglio'' family. It was the first printing process that yielded half-tones without using line- or dot-based techniques like hatching, cross-hatching or stipple. Mezzotint achieves tonalit ...
s by John Samuelson Templeton. These publications also contained scientific descriptions and represented the first textbooks of the subject. In the summer of 1834, Hawkins, after a taxation by
William Buckland William Buckland DD, FRS (12 March 1784 – 14 August 1856) was an English theologian who became Dean of Westminster. He was also a geologist and palaeontologist. Buckland wrote the first full account of a fossil dinosaur, which he named ...
and
Gideon Mantell Gideon Algernon Mantell MRCS FRS (3 February 1790 – 10 November 1852) was a British obstetrician, geologist and palaeontologist. His attempts to reconstruct the structure and life of ''Iguanodon'' began the scientific study of dinosaurs: in ...
, sold his extensive collection, then the largest of its kind in the world, to the British Museum. However, curator Koenig quickly discovered that the fossils had been heavily restored with plaster, applied by an Italian artist from
Lucca Lucca ( , ) is a city and ''comune'' in Tuscany, Central Italy, on the Serchio River, in a fertile plain near the Ligurian Sea. The city has a population of about 89,000, while its province has a population of 383,957. Lucca is known as one ...
; of the most attractive piece, an ''Ichthyosaurus'' specimen, almost the entire tail was fake. It turned out that Professor Buckland had been aware of this beforehand, and the museum was forced to reach a settlement with Hawkins, and gave the fake parts a lighter colour to differentiate them from the authentic skeletal elements. Ichthyosaurs became even more popular in 1854 by the rebuilding at
Sydenham Hill Sydenham Hill forms part of a longer ridge and is an affluent locality in southeast London. It is also the name of a road which runs along the northeastern part of the ridge, demarcating the London Boroughs of Southwark, Bromley, and Lewisham ...
of
the Crystal Palace The Crystal Palace was a cast iron and plate glass structure, originally built in Hyde Park, London, to house the Great Exhibition of 1851. The exhibition took place from 1 May to 15 October 1851, and more than 14,000 exhibitors from around th ...
, originally erected at the world exhibition of 1851. In the surrounding park, life-sized, painted, concrete statues of extinct animals were placed, which were designed by
Benjamin Waterhouse Hawkins Benjamin Waterhouse Hawkins (8 February 1807 – 27 January 1894) was an English sculptor and natural history artist renowned for his work on the life-size models of dinosaurs in the Crystal Palace Park in south London. The models, accurately ...
under the direction of Richard Owen. Among them were three models of an ichthyosaur. Although it was known that ichthyosaurs had been animals of the open seas, they were shown basking on the shore, a convention followed by many nineteenth century illustrations with the aim, as Conybeare once explained, of better exposing their build. This led to the misunderstanding that they really had an amphibious lifestyle. The pools in the park were at the time subjected to tidal changes, so that fluctuations in the water level at intervals submerged the ichthyosaur statues, adding a certain realism. Remarkably, internal skeletal structures, such as the scleral rings and the many phalanges of the flippers, were shown at the outside.


Later 19th-century finds

During the nineteenth century, the number of described ichthyosaur genera gradually increased. New finds allowed for a better understanding of their anatomy. Owen had noted that many fossils showed a downward bend in the rear tail. At first, he explained this as a ''post mortem'' effect, a tendon pulling the tail end downwards after death. However, after an article on the subject by Philip Grey Egerton, Owen considered the possibility that the oblique section could have supported the lower lobe of a tail fin. This hypothesis was confirmed by new finds from Germany. In the Posidonia Shale at
Holzmaden Holzmaden is a town in Baden-Württemberg, Germany that lies between Stuttgart and Ulm. Holzmaden is 4 km south-east from Kirchheim unter Teck and 19 km south-east of Esslingen am Neckar. The A 8 runs south from Holzmaden. The town a ...
, dating from the early Jurassic, already in the early nineteenth century, the first ichthyosaur skeletons had been found. During the latter half of the century, the rate of discovery quickly increased to a few hundred each year. Ultimately, over four thousand were uncovered, forming the bulk of ichthyosaur specimens displayed today. The sites were also a '' Konservat-Lagerstätte'', meaning not only the quantity, but also the quality was exceptional. The skeletons were very complete and often preserved soft tissues, including tail and dorsal fins. Additionally, female individuals were discovered with embryos.


20th century

In the early twentieth century, ichthyosaur research was dominated by the German paleontologist
Friedrich von Huene Friedrich von Huene, born Friedrich Richard von Hoinigen, (March 22, 1875 – April 4, 1969) was a German paleontologist who renamed more dinosaurs in the early 20th century than anyone else in Europe. He also made key contributions about v ...
, who wrote an extensive series of articles, taking advantage of an easy access to the many specimens found in his country. The amount of anatomical data was hereby vastly increased. Von Huene also travelled widely abroad, describing many fossils from locations outside of Europe. During the 20th century,
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the C ...
became an important source of new fossils. In 1905, the Saurian Expedition led by
John Campbell Merriam John Campbell Merriam (October 20, 1869 – October 30, 1945) was an American paleontologist, educator, and conservationist. The first vertebrate paleontologist on the West Coast of the United States, he is best known for his taxonomy of ver ...
and financed by
Annie Montague Alexander Annie Montague Alexander (29 December 1867 - 10 September 1950) was an explorer, naturalist, paleontological collector, and philanthropist. She founded the University of California Museum of Paleontology (UCMP) and the Museum of Vertebrate Zoo ...
, found twenty-five specimens in central
Nevada Nevada ( ; ) is a state in the Western region of the United States. It is bordered by Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. Nevada is the 7th-most extensive, ...
, which were under a shallow ocean during the Triassic. Several of these are now in the collection of the University of California Museum of Paleontology. After a slack during the middle of the century, with no new genera being named between the 1930s and the 1970s, the rate of discoveries picked up towards its end. Other specimens are embedded in the rock and visible at Berlin–Ichthyosaur State Park in Nye County. In 1977 the Triassic ichthyosaur ''Shonisaurus'' became the state fossil of Nevada. About half of the ichthyosaur genera today seen as valid were described after 1990. In 1992 Canadian paleontologist Elizabeth Nicholls uncovered the largest known specimen, a '' Shastasaurus''. The new finds have allowed a gradual improvement in knowledge about the anatomy and physiology of what had already been seen as rather advanced "Mesozoic dolphins". Christopher McGowan published a larger number of articles and also brought the group to the attention of the general public. The new method of
cladistics 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 cha ...
provided a means to exactly calculate the relationships between groups of animals, and in 1999, Ryosuke Motani published the first extensive study on ichthyosaur
phylogenetics In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
. In 2003, McGowan and Motani published the first modern textbook on the Ichthyosauria and their closest relatives.


Evolutionary history


Origin

The origin of the ichthyosaurs is contentious. Until recently, clear transitional forms with land-dwelling vertebrate groups had not yet been found, the earliest known species of the ichthyosaur lineage being already fully aquatic. In 2014, a small basal ichthyosauriform from the upper Lower Triassic was described that had been discovered in China with characteristics suggesting an amphibious lifestyle. In 1937,
Friedrich von Huene Friedrich von Huene, born Friedrich Richard von Hoinigen, (March 22, 1875 – April 4, 1969) was a German paleontologist who renamed more dinosaurs in the early 20th century than anyone else in Europe. He also made key contributions about v ...
even hypothesised that ichthyosaurs were not reptiles, but instead represented a lineage separately developed from amphibians. Today, this notion has been discarded and a consensus exists that ichthyosaurs are
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are distin ...
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct therapsid ...
s, having descended from terrestrial egg-laying amniotes during the late
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
or the earliest Triassic. However, establishing their position within the amniote evolutionary tree has proven difficult, due to their heavily derived morphology obscuring their ancestry. Several conflicting hypotheses have been posited on the subject. In the second half of the 20th century, ichthyosaurs were usually assumed to be of the Anapsida, seen as an early branch of "primitive" reptiles. This would explain the early appearance of ichthyosaurs in the fossil record, and also their lack of clear affinities with other reptile groups, as anapsids were supposed to be little specialised. This hypothesis has become unpopular for being inherently vague because Anapsida is an unnatural,
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
group. Modern exact quantitative cladistic analyses consistently indicate that ichthyosaurs are members of the clade
Diapsida Diapsids ("two arches") are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The group first appeared about three hundred million years ag ...
. Some studies showed a basal, or low, position in the diapsid tree. More analyses result in their being
Neodiapsida Neodiapsida is a clade, or major branch, of the reptilian family tree, typically defined as including all diapsids apart from some early primitive types known as the araeoscelidians. Modern reptiles and birds belong to the neodiapsid subclade S ...
, a derived diapsid subgroup. Until the 1980s, a close relationship was assumed between the Ichthyosauria and the
Sauropterygia Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosauria became ...
, another marine reptile group, within an overarching
Euryapsida __NOTOC__ Euryapsida is a polyphyletic (unnatural, as the various members are not closely related) group of sauropsids that are distinguished by a single temporal fenestra, an opening behind the orbit, under which the post-orbital and squamosal bo ...
. Today, however, this is usually rejected, with the Euryapsida being seen as an unnatural polyphyletic assemblage of reptiles that happen to share some adaptations to a swimming lifestyle. An exception is a study in 1997 by John Merck, which showed
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...
archosauromorph euryapsids.


Affinity with the Hupehsuchia

Since 1959, a second enigmatic group of ancient sea reptiles is known, the
Hupehsuchia Hupehsuchia is an order of diapsid reptiles closely related to ichthyosaurs. The group was short-lasting, with a temporal range restricted to the late Olenekian age, spanning only a few million years of the Early Triassic. The order gets its nam ...
. Like the Ichthyopterygia, the Hupehsuchia have pointed snouts and show
polydactyly Polydactyly or polydactylism (), also known as hyperdactyly, is an anomaly in humans and animals resulting in supernumerary fingers and/or toes. Polydactyly is the opposite of oligodactyly (fewer fingers or toes). Signs and symptoms In human ...
, the possession of more than five fingers or toes. Their limbs more resemble those of land animals, making them appear as a transitional form between these and ichthyosaurs. Initially, this possibility was largely neglected because the Hupehsuchia have a fundamentally different form of propulsion, with an extremely stiffened trunk. The similarities were explained as a case of convergent evolution. Furthermore, the descent of the Hupehsuchia is no less obscure, meaning a possible close relationship would hardly clarify the general evolutionary position of the ichthyosaurs. In 2014, ''
Cartorhynchus ''Cartorhynchus'' (meaning "shortened snout") is an extinct genus of early ichthyosauriform marine reptile that lived during the Early Triassic epoch, about 248 million years ago. The genus contains a single species, ''Cartorhynchus lenticarpu ...
'' was announced, a small species with a short snout, large flippers, and a stiff trunk. Its lifestyle might have been amphibious. Motani found it to be more basal than the Ichthyopterygia and named an encompassing clade
Ichthyosauriformes The Ichthyosauriformes are a group of marine reptiles, belonging to the Ichthyosauromorpha, that lived during the Mesozoic. The stem clade Ichthyosauriformes was in 2014 defined by Ryosuke Motani and colleagues as the group consisting of all ic ...
. The latter group was combined with the Hupesuchia into the
Ichthyosauromorpha The Ichthyosauromorpha are an extinct clade of marine reptiles consisting of the Ichthyosauriformes and the Hupehsuchia, living during the Mesozoic. The node clade Ichthyosauromorpha was first defined by Ryosuke Motani ''et al.'' in 2014 as t ...
. The ichthyosauromorphs were found to be diapsids. The proposed relationships are shown by this cladogram:


Early Ichthyopterygia

The earliest ichthyosaurs are known from the Early and Early-Middle ( Olenekian and
Anisian In the geologic timescale, the Anisian is the lower stage or earliest age of the Middle Triassic series or epoch and lasted from million years ago until million years ago. The Anisian Age succeeds the Olenekian Age (part of the Lower Triassic ...
) Triassic strata of
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
,
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
, and
Spitsbergen Spitsbergen (; formerly known as West Spitsbergen; Norwegian: ''Vest Spitsbergen'' or ''Vestspitsbergen'' , also sometimes spelled Spitzbergen) is the largest and the only permanently populated island of the Svalbard archipelago in northern No ...
in
Norway Norway, officially the Kingdom of Norway, is a Nordic country in Northern Europe, the mainland territory of which comprises the western and northernmost portion of the Scandinavian Peninsula. The remote Arctic island of Jan Mayen and th ...
, being up to 246 million years old. These first forms included the genera '' Chaohusaurus'', '' Grippia'', and '' Utatsusaurus''. This diversity suggests an even earlier origin. They more resembled finned lizards than the fishes or dolphins to which the later, more familiar species were similar. Their bodies were elongated and they probably used an anguilliform locomotion, swimming by undulations of the entire trunk. Like land animals, their pectoral girdles and pelves were robustly built, and their vertebrae still possessed the usual interlocking processes to support the body against the force of gravity. However, they were already rather advanced in having limbs that had been completely transformed into flippers. They also were probably warm-blooded and
viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...
. These very early "proto-ichthyosaurs" had such a distinctive build compared to "ichthyosaurs proper" that Motani excluded them from the Ichthyosauria and placed them in a basal position in a larger
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
, the Ichthyopterygia. However, this solution was not adopted by all researchers.


Later Triassic forms

The basal forms quickly gave rise to ichthyosaurs in the narrow sense sometime around the boundary between the
Early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which is a ...
and
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma and ...
; the earliest Ichthyosauria in the sense Motani gave to the concept, appear about 245 million years ago. These later diversified into a variety of forms, including the still
sea serpent A sea serpent or sea dragon is a type of dragon sea monster described in various mythologies, most notably Mesopotamian (Tiamat), Judaeo-Christian ( Leviathan), Greek (Cetus, Echidna, Hydra, Scylla), and Norse (Jörmungandr). Mythology and fol ...
-like '' Cymbospondylus'', a problematic form which reached ten metres in length, and smaller, more typical forms like '' Mixosaurus''. The Mixosauria were already very fish-like with a pointed skull, a shorter trunk, a more vertical tail fin, a dorsal fin, and short flippers containing many phalanges. The sister group of the Mixosauria were the more advanced Merriamosauria. By the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch. ...
, merriamosaurs consisted of both the large, classic Shastasauria and more advanced, "dolphin-like" Euichthyosauria. Experts disagree over whether these represent an evolutionary continuum, with the less specialised shastosaurs a paraphyletic grade that was evolving into the more advanced forms, or whether the two were separate clades that evolved from a common ancestor earlier on. Euichthyosauria possessed more narrow front flippers, with a reduced number of fingers. Basal euichthyosaurs were '' Californosaurus'' and '' Toretocnemus''. A more derived branch were the Parvipelvia, with a reduced pelvis, basal forms of which are '' Hudsonelpidia'' and '' Macgowania''. During the
Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by ...
and
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic ...
, Shastosauria reached huge sizes. '' Shonisaurus popularis'', known from a number of specimens from the Carnian of Nevada, was long. Norian Shonisauridae are known from both sides of the Pacific. '' Himalayasaurus tibetensis'' and '' Tibetosaurus'' (probably a
synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
) have been found in
Tibet Tibet (; ''Böd''; ) is a region in East Asia, covering much of the Tibetan Plateau and spanning about . It is the traditional homeland of the Tibetan people. Also resident on the plateau are some other ethnic groups such as Monpa, Tama ...
. These large (10- to 15-m-long) ichthyosaurs have by some been placed into the genus ''Shonisaurus''. The gigantic ''Shonisaurus sikanniensis'' (considered as a shastasaurus between 2011 and 2013) whose remains were found in the Pardonet Formation of
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, for ...
by Elizabeth Nicholls, has been estimated to be as much as in length—if correct, the largest marine reptile known to date. In the Late Triassic, ichthyosaurs attained the peak of their diversity. They occupied many
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s. Some were
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 l ...
s; others were hunters of small prey. Several species perhaps specialised in
suction feeding Aquatic feeding mechanisms face a special difficulty as compared to feeding on land, because the density of water is about the same as that of the prey, so the prey tends to be pushed away when the mouth is closed. This problem was first identifi ...
or were ram feeders; also, durophagous forms are known. Towards the end of the Late Triassic, a decline of variability seems to have occurred. The giant species seemed to have disappeared at the end of the Norian.
Rhaetian The Rhaetian is the latest age of the Triassic Period (in geochronology) or the uppermost stage of the Triassic System (in chronostratigraphy). It was preceded by the Norian and succeeded by the Hettangian (the lowermost stage or earliest age ...
(latest Triassic) ichthyosaurs are known from England, and these are very similar to those of the
Early Jurassic The Early Jurassic Epoch (in chronostratigraphy corresponding to the Lower Jurassic Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-Jurassic extinction event, 201.3 Ma&nb ...
. A possible explanation is an increased competition by
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
s,
Teleost 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. Tel ...
ei, and the first
Plesiosauria 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 ...
. Like the dinosaurs, the ichthyosaurs and their contemporaries, the plesiosaurs, survived the Triassic–Jurassic extinction event, and quickly diversified again to fill the vacant ecological niches of the early Jurassic.


Jurassic

During the Early Jurassic, the ichthyosaurs still showed a large variety of species, ranging from in length. From this
epoch In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured. The moment of epoch is usually decided by ...
, the original British discoveries were made, so that among ichthyosaurs, their names are the ones most familiar to the general public. Genera include '' Eurhinosaurus'', '' Ichthyosaurus'', '' Leptonectes'', '' Stenopterygius'', and the large predator '' Temnodontosaurus'', along with the basal parvipelvian '' Suevoleviathan'', which was little changed from its Norian ancestors. The general morphological variability had been strongly reduced, however. Giant forms, suction feeders and durophagous species were absent. All of these animals were streamlined, dolphin-like forms, although the more basal animals were perhaps more elongated than the advanced and compact ''Stenopterygius'' and ''Ichthyosaurus''. The latter belonged to a parvipelvian subgroup, the Thunnosauria. These were fully adapted to an efficient thunniform locomotion, propelling themselves with the end of the tail only, equipped with a vertical tail fin. Their front flippers had more than five fingers. Another parvipelvian branch was the Eurhinosauria such as ''Leptonectes'' and ''Eurhinosaurus'', which were specialised forms having very elongated and pointy snouts. Few ichthyosaur fossils are known from the Middle Jurassic. This might be a result of the poor
fossil record 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 in ...
in general of this epoch. The strata of the Late Jurassic seem to indicate that a further decrease in diversity had taken place. From the Middle Jurassic onwards, almost all ichthyosaurs belonged to the thunnosaurian clade Ophthalmosauridae. Represented by the ''
Ophthalmosaurus ''Ophthalmosaurus'' (meaning "eye lizard" in Greek) is an ichthyosaur of the Jurassic period (165–150 million years ago). Possible remains from the Cretaceous, around 145 million years ago, are also known. It was a relatively medium-sized ichth ...
'' and related genera, they were very similar in general build to ''Ichthyosaurus''. The eyes of ''Ophthalmosaurus'' were huge, and these animals likely hunted in dim and deep water. However, new finds from the Cretaceous indicate that ichthyosaur diversity in the Late Jurassic must have been underestimated.


Cretaceous

Traditionally, ichthyosaurs were seen as decreasing in diversity even further with 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 ...
, though they had a worldwide distribution. All fossils from this period were referred to a single genus: '' Platypterygius''. This last ichthyosaur genus was thought to have become extinct early in the late Cretaceous, during the
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in ...
about 95 million years ago. The ichthyosaurs thus would have disappeared much earlier than other large Mesozoic reptile groups that generally survived until the end of the Cretaceous. Two major explanations were given for this. Firstly, it could have been a matter of chance. The second explanation had the extinction as caused by competition. Less hydrodynamically efficient animals, such as the
Mosasaur Mosasaurs (from Latin ''Mosa'' meaning the ' Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht o ...
idae and long-necked
plesiosaur The Plesiosauria (; Ancient Greek, Greek: πλησίος, ''plesios'', meaning "near to" and Sauria, ''sauros'', meaning "lizard") or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauro ...
s, flourished. The ichthyosaurian overspecialisation could be a contributing factor to their extinction, possibly being unable to 'keep up' with the fast-swimming and highly evasive new
teleost 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. Tel ...
fish, which had become dominant at this time, against which the sit-and-wait ambush strategies of the mosasaurids proved superior. This model thus emphasised evolutionary stagnation, the only innovation shown by ''Platypterygius'' being its ten fingers. Recent studies, however, show that ichthyosaurs were actually far more diverse in the Cretaceous than previously thought. Fragments referred to ''Platypterygius'' in fact represented diverse species. In 2012, at least eight lineages spanned the Jurassic-Cretaceous boundary, among them ''
Acamptonectes ''Acamptonectes'' is a genus of ophthalmosaurid ichthyosaurs, a type of dolphin-like marine reptiles, that lived during the Early Cretaceous around 130 million years ago. The first specimen, a partial adult skeleton, was discovered in Speeto ...
'', '' Sveltonectes'', '' Caypullisaurus'', and '' Maiaspondylus''. In 2013, a Cretaceous basal thunnosaurian was revealed: '' Malawania''. Indeed, likely a radiation during the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous ( chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pr ...
occurred due to an increase of coastlines when the continents further broke up. The demise of the ichthyosaurs has recently been described as a two-step process. A first extinction event in the beginning of the
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in ...
eliminated two of the three ichthyosaur feeding guilds then present, the 'soft-prey specialists' and the 'generalists', leaving only an apex predator group, which probably was not particularly specialized. The second major ichthyosaur extinction took place during the
Cenomanian-Turonian boundary event The Cenomanian-Turonian boundary event, also known as the Cenomanian-Turonian extinction, Cenomanian-Turonian oceanic anoxic event ( OAE 2), and referred to also as the Bonarelli event, was one of two anoxic extinction events in the Cretaceous pe ...
, an '
anoxic event Oceanic anoxic events or anoxic events ( anoxia conditions) describe periods wherein large expanses of Earth's oceans were depleted of dissolved oxygen (O2), creating toxic, euxinic (anoxic and sulfidic) waters. Although anoxic events have not ...
', eliminating the apex predators, after which just a single lineage survived, ''Platypterygius hercynicus'', which disappeared shortly afterwards, about 93 million years ago. Ichthyosaur extinction was thus a pair of abrupt events rather than a long decline, probably related to the environmental upheavals and climatic changes in the Cenomanian and
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded ...
. If so, it was not caused by mosasaurid competition; large mosasaurs did not appear until 3 million years after the ichthyosaur extinction, likely to fill the resulting ecological void. Plesiosaurian polycoltylids perhaps also filled some of the niches previously occupied by ichthyosaurs, although they had coexisted for 19 million years. The extinction was most likely the result of ecological change and volatility that caused changes in migration, food availability, and birthing grounds. This part of the Cretaceous was one in which many other marine extinctions occurred, including those of some types of microplankton, ammonites,
belemnite Belemnitida (or the belemnite) is an extinct order of squid-like cephalopods that existed from the Late Triassic to Late Cretaceous. Unlike squid, belemnites had an internal skeleton that made up the cone. The parts are, from the arms-most to ...
s, and reef-building
bivalves Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, biv ...
.


Taxonomy

This
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
was presented by Michael Werner Maisch in his 2010 review of ichthyosaur classification. * Clade Ichthyopterygia (''sensu'' Motani (1999)) ** Genus '' Chaohusaurus'' ** Family Grippiidae ** Family Parvinatatoridae ** Family Thaisauridae ** Family Utatsusauridae ** ? Family Omphalosauridae ** ? Genus '' Isfjordosaurus'' ** Order Ichthyosauria (''sensu'' Motani (1999)) *** Family Quasianosteosauridae *** Parvorder Hueneosauria **** Nanorder Mixosauria ***** Family Wimaniidae ***** Family Mixosauridae **** Nanorder Longipinnati ***** Family Toretocnemidae ***** Family Cymbospondylidae ***** Hyporder Merriamosauria ****** Family Merriamosauridae ****** Family Besanosauridae ****** Family Shastasauridae ****** Family Shonisauridae ****** Family Californosauridae ****** Minorder Parvipelvia ******* Family Hudsonelpidiidae ******* Family Macgowaniidae ******* Suborder Neoichthyosauria ******** Family Temnodontosauridae ******** Family Leptonectidae ******** Family Suevoleviathanidae ******** Infraorder Thunnosauria ********* Family Ichthyosauridae ********* Family Stenopterygiidae ********* Family Ophthalmosauridae


Phylogeny

In modern
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological sp ...
, clades are defined that contain all species forming a certain branch of the evolutionary tree. This also allows one to clearly indicate all relationships between the several subgroups in a
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 ...
. In 1999, a node clade Ichthyopterygia was defined by Motani as the group consisting of the
last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of ''Ichthyosaurus communis'', ''Utatsusaurus hataii'' and ''Parvinatator wapitiensis''; and all its descendants. Within Motani's phylogeny, the Ichthyopterygia were the larger parent clade of a smaller stem clade Ichthyosauria that was defined as the group consisting of ''Ichthyosaurus communis'' and all species more closely related to ''Ichthyosaurus'' than to ''Grippia longirostris''. Motani's concept of the Ichthyosauria was thus more limited than the traditional one that also contained basal forms, such as ''Grippia'', ''Utatsusaurus'', and ''Parvinatator''. The following
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 ...
is based on Motani (1999): An alternative terminology was proposed by Maisch & Matzke in 2000, trying to preserve the traditional, more encompassing content of the concept Ichthyosauria. They defined a node clade Ichthyosauria as the group consisting of the last common ancestor of '' Thaisaurus chonglakmanii'', ''Utatsusaurus hataii'', and ''Ophthalmosaurus icenicus'', and all its descendants. Ichthyosauria ''sensu'' Motani might materially be identical to a clade that Maisch & Matzke in 2000 called Hueneosauria, depending on the actual relationships. Cladogram based on Maisch and Matzke (2000) and Maisch and Matzke (2003) with clade names following Maisch (2010):


Description


Size

Ichthyosaurs averaged about in length. Some individual specimens were as short as ; some species were much larger: the Triassic '' Shonisaurus popularis'' was about long and in 2004 '' Shonisaurus sikanniensis'' (classified as a shastasaurus between 2011 and 2013) was estimated to have been in length. Fragmentary finds suggest the presence of a form in the early Jurassic. In 2018, lower jaw fragments from England were reported indicating a length of between 20 and 25 m (66 to 82 ft). According to weight estimates by Ryosuke Motani a ''Stenopterygius'' weighed around , whilst a ''
Ophthalmosaurus ''Ophthalmosaurus'' (meaning "eye lizard" in Greek) is an ichthyosaur of the Jurassic period (165–150 million years ago). Possible remains from the Cretaceous, around 145 million years ago, are also known. It was a relatively medium-sized ichth ...
icenicus'' weighed .


General build

While the earliest known members of the ichthyosaur lineage were more eel-like in build, later ichthyosaurs resembled more typical fishes or dolphins, having a
porpoise Porpoises are a group of fully aquatic marine mammals, all of which are classified under the family Phocoenidae, parvorder Odontoceti (toothed whales). Although similar in appearance to dolphins, they are more closely related to narwhals a ...
-like head with a short neck and a long snout. Ichthyosaur fore and hind limbs had been fully transformed into flippers. Some species had a fin on their backs and a more or less vertical fin at the rear of a rather short tail. Although ichthyosaurs looked like fish, they were not.Evolutionary biologist
Stephen Jay Gould Stephen Jay Gould (; September 10, 1941 – May 20, 2002) was an American paleontologist, evolutionary biologist, and historian of science. He was one of the most influential and widely read authors of popular science of his generation. Gould ...
said that the ichthyosaur was his favourite example of convergent evolution, where similarities of structure are
analogous Analogy (from Greek ''analogia'', "proportion", from ''ana-'' "upon, according to" lso "against", "anew"+ ''logos'' "ratio" lso "word, speech, reckoning" is a cognitive process of transferring information or meaning from a particular subject (t ...
, not homologous, thus not caused by a common descent, but by a similar adaptation to an identical environment:
This sea-going reptile with terrestrial ancestors converged so strongly on fishes that it actually evolved a dorsal fin and tail in just the right place and with just the right hydrological design. These structures are all the more remarkable because they evolved from nothing—the ancestral terrestrial reptile had no hump on its back or blade on its tail to serve as a precursor.


Diagnostic traits

Derived ichthyosaurs in the narrow sense, as defined by Motani in 1999, differ from their closest basal ichthyopterygian relatives in certain traits. Motani listed a number of these. The external nostril is located on the side of the skull, and is hardly visible from above. The upper rim of the eye socket consists of a bone bar formed by the prefrontal and the postfrontal bones. The postorbital in side view is excluded from the supratemporal fenestra. The opening for the parietal eye is located on the border of the parietal and the frontal bone. The lateral wing of the pterygoid is incompletely and variably ossified. The ulna lacks the part behind the original shaft axis. The rear dorsal vertebrae are disc-shaped.


Skeleton


Skull

Basal Ichthyopterygia already had elongated, triangular skulls. With ichthyosaurs in the narrow sense, their snouts became very pointy. The snout is formed by the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
. The
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The ...
behind it is usually shorter and sometimes excluded from the external nostril by the rear branch of the premaxilla. Accordingly, the number of premaxillary teeth is high, while the maxillary teeth are fewer in number or even completely absent. The rear top of the snout is formed by the nasal bones. Derived species have a foramen internasale, a midline opening separating the rear of the nasal bones. The nasal bone usually forms the top and front rim of the bony nostril, itself often placed just in front of the eye socket. However, with some Triassic species, the premaxilla is so strongly extended at its back that it even excludes the nasal from the nostril. The rear of the skull is dominated by a large
eye socket In anatomy, the orbit is the cavity or socket of the skull in which the eye and its appendages are situated. "Orbit" can refer to the bony socket, or it can also be used to imply the contents. In the adult human, the volume of the orbit is , of ...
, often covering the major part of the rear side surface. In the socket, a large scleral ring is present; this is a circular structure of small, overlapping bone segments protecting the eye against the water pressure. Both in the relative and absolute senses, ichthyosaurs have the largest eye sockets of all known vertebrates. The other rear skull elements are typically so compressed and fused that they are difficult to identify. The top rear element of the skull was usually assumed to be the supratemporal bone, while the
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestral co ...
and
quadratojugal The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians. Anatomy and function In animals with a quadratojugal bone, it is typically found connected to the jugal (cheek) bone from the front and ...
were sometimes fused. However, in 1968,
Alfred Sherwood Romer Alfred Sherwood Romer (December 28, 1894 – November 5, 1973) was an American paleontologist and biologist and a specialist in vertebrate evolution. Biography Alfred Romer was born in White Plains, New York, the son of Harry Houston Romer an ...
stated that the presumed supratemporal was in fact the squamosal, an interpretation which was supported by McGowan in 1973. In 1990, though, John Steve Massare convinced most researchers that the original identification had been the correct one after all. The supratemporal forms the rear rim of the supratemporal opening; a lower temporal opening at the side is lacking. The front rim of the supratemporal opening is typically formed by the
postfrontal The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
; only with the very basal '' Utatsusaurus'' the
postorbital The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...
and the squamosal still reach the edge. Between the paired supratemporal openings, the skull roof is narrow; some species have a longitudinal crest on it as an attachment for the jaw muscles. Basal Ichthyopterygia have a
parietal eye A parietal eye, also known as a third eye or pineal eye, is a part of the epithalamus present in some vertebrates. The eye is located at the top of the head, is photoreceptive and is associated with the pineal gland, regulating circadian rhyth ...
opening between the paired
parietal bone The parietal bones () are two bones in the skull which, when joined at a fibrous joint, form the sides and roof of the cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, and four angles. It is named ...
s. With ichthyosaurs proper, this opening moves to the front, first to the border between the parietals and the frontals and ultimately between the frontals, a condition shown by derived species. Postparietal and
tabular bone The tabular bones are a pair of triangular flat bones along the rear edge of the skull which form pointed structures known as tabular horns in primitive Teleostomi. References Fish anatomy Amphibian anatomy {{vertebrate-anatomy-stub ...
s are lacking. Often, the bones of the back of the skull and the palate are incompletely ossified, apparently having partly remained
cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck a ...
. The
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
is typically very convex. The
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foo ...
, the bone transmitting sound waves from the
eardrum In the anatomy of humans and various other tetrapods, the eardrum, also called the tympanic membrane or myringa, is a thin, cone-shaped membrane that separates the external ear from the middle ear. Its function is to transmit sound from the ai ...
to the
middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
, is elongated and not pierced by a foramen. Pterygoid teeth are typically lacking.


Lower jaws

Like the snout, the lower jaws are elongated. However, in some species, such as '' Eurhinosaurus'' and '' Excalibosaurus'', the front of the snout far protrudes beyond the lower jaws. While the front of the lower jaw is typically low, its rear depth is very variable. The greater part of the lower jaw is formed by the front
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
, the tooth-bearing bone. At its inner side the dentary is covered by a
splenial The splenial is a small bone in the lower jaw of reptiles, amphibians and birds, usually located on the lingual side (closest to the tongue) between the angular Angular may refer to: Anatomy * Angular artery, the terminal part of the facial arte ...
that extends forwards until the symphysis, the common contact surface where both lower jaws are grown together. The jaw joints do not allow a horizontal chewing movement: they function as simple hinges to vertically open or close the jaws.


Teeth

Ichthyosaur teeth are typically conical. Fish-eating species have long and slender tooth crowns that are slightly recurved. Forms specialised in catching larger prey have shorter, broader, and straighter teeth; sometimes, cutting edges are present. ''
Thalattoarchon ''Thalattoarchon'' is an extinct genus of ichthyosaur from the Middle Triassic of the western United States. The type species ''Thalattoarchon saurophagis'' (meaning "lizard-eating sovereign of the sea" in Greek) was discovered in Nevada, USA, ...
'', an apex predator, had larger teeth formed like flattened blades. Durophagous species that ate
shellfish Shellfish is a colloquial and fisheries term for exoskeleton-bearing aquatic invertebrates used as food, including various species of molluscs, crustaceans, and echinoderms. Although most kinds of shellfish are harvested from saltwater enviro ...
have low, convex teeth that are closely packed. Many ichthyosaur dentitions are heterodont, combining several tooth shapes, e.g. small teeth in the front and larger teeth at the rear. The teeth are usually placed in tooth sockets; derived species possess a common tooth groove. In the latter case, adult individuals sometimes become toothless. Teeth in tooth sockets sometimes fuse with the jawbone. With ichthyosaur teeth, the
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by en ...
shows prominent vertical wrinkles. Durophagous forms have teeth with deep vertical grooves and wrinkles in the enamel.


Postcrania


=Vertebral column

= Basal Ichthyopterygia, like their land-dwelling ancestors, still had
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e that possessed a full set of processes that allowed them to interlock and articulate, forming a vertebral column supporting the weight of the body. As ichthyosaurs were fully aquatic, their bodies were supported by the Archimedes force exerted by the water; in other words, they were
buoyant Buoyancy (), or upthrust, is an upward force exerted by a fluid that opposes the weight of a partially or fully immersed object. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus the pr ...
. Therefore, the vertebral processes had lost much of their function. Early ichthyosaurs proper had rear dorsal vertebrae that had become disc-shaped, like those of typical fishes. With more derived species, the front dorsals also became discs. Gradually, most processes were lost, including those for rib attachment. The vertebral bodies became much shorter. The front and rear sides of the discs were hollowed out, resulting in a so-called amphicoelous condition. A transverse cross-section of such a vertebra has an hourglass shape. This morphology is unique within the
Amniota Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are distingu ...
and makes discerning ichthyosaur vertebrae from those of other marine reptiles easy. The only process that kept its function was the spine at the top, serving as an attachment for the dorsal muscles. However, even the spine became a simple structure. The neural arch, of which it was an outgrowth, typically no longer fused to the vertebral centre. The neck is short, and derived species show a reduction in the number of
cervical vertebra In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In sau ...
e. The short neck positions the skull close to the trunk, usually in a slight oblique elevation to it. Derived species usually also have a reduced number of dorsals, the total of presacral vertebrae totalling about forty to fifty. The vertebral column is little differentiated. Basal Ichthyopterygia still have two sacral vertebrae, but these are not fused. Early Triassic forms have a transversely flattened tail base with high spines for an undulating tail movement. Derived forms have a shorter tail with the characteristic kink at the end; a section of wedge-shaped vertebrae, itself supporting the fleshy upper tail fin lobe, forced the tail end into the lower fin lobe. As derived species no longer have transversal processes on their vertebrae—again a condition unique in the Amniota—the parapophyseal and diapophysael rib joints have been reduced to flat facets, at least one of which is located on the vertebral body. The number of facets can be one or two; their profile can be circular or oval. Their shape often differs according to the position of the vertebra within the column. The presence of two facets per side does not imply that the rib itself is double-headed: often, even in that case, it has a single head. The ribs typically are very thin and possess a longitudinal groove on both the inner and the outer sides. The lower side of the chest is formed by
gastralia Gastralia (singular gastralium) are dermal bones found in the ventral body wall of modern crocodilians and tuatara, and many prehistoric tetrapods. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In ...
. These belly ribs have a single centre segment and one or two outer segments per side. They are not fused into a real plastron. Usually two gastralia are present per dorsal rib.


=Appendicular skeleton

= The
shoulder girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of ...
of ichthyosaurs is not much modified from its original condition. Some basal forms show a hatchet- or crescent-shaped shoulder blade or
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...
; derived forms have an elongated blade positioned on a broader base. The scapula is not fused with the
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
into a
scapulocoracoid The scapulocoracoid is the unit of the pectoral girdle that contains the coracoid and scapula. The coracoid itself is a beak-shaped bone that is commonly found in most vertebrates with a few exceptions. The scapula is commonly known as the ''shoulde ...
, indicating that the forces exerted on the shoulder girdle were moderate. The
shoulder joint The shoulder joint (or glenohumeral joint from Greek ''glene'', eyeball, + -''oid'', 'form of', + Latin ''humerus'', shoulder) is structurally classified as a synovial ball-and-socket joint and functionally as a diarthrosis and multiaxial joint ...
is positioned on the border between the scapula and the coracoid. Both coracoids are fused on their common midline. The coracoid shape is very variable, but usually it is rather low. The upper part of the shoulder girdle is formed by two long and slender
clavicle The clavicle, or collarbone, is a slender, S-shaped long bone approximately 6 inches (15 cm) long that serves as a strut between the shoulder blade and the sternum (breastbone). There are two clavicles, one on the left and one on the righ ...
s, crowned by a central interclavicular bone that is large and triangular with basal forms, small and T-shaped in Jurassic species. Breast bones or sterna are absent. Basal forms have a forelimb that is still functionally differentiated, in some details resembling the arm of their land-dwelling forebears; the
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
and
radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...
are elongated and somewhat separated; the
carpal The carpal bones are the eight small bones that make up the wrist (or carpus) that connects the hand to the forearm. The term "carpus" is derived from the Latin carpus and the Greek καρπός (karpós), meaning "wrist". In human anatomy, th ...
s are rounded, allowing the wrist to rotate; the number of phalanges is within the range shown by land animals. Ichthyosaurs proper, to the contrary, have a forelimb that is fully adapted to its function as a flipper. However, the adaptations are very variable. Triassic species typically have a very derived
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a round ...
, changed into a disc. Jurassic species tend to have a more elongated humeral form with a rounded head, narrow shaft, and expanded lower end. The radius and ulna are always strongly flattened, but can be circular, with or without notch, or have a waist. Notches can be homologous to the original shafts, but also be newly formed. Jurassic forms no longer have a space, the ''spatium interosseum'', between the radius and ulna. Often, the latter bones gradually merge into lower, disc-shaped elements - the up to four carpals which again differ little in form from the up to five
metacarpal In human anatomy, the metacarpal bones or metacarpus form the intermediate part of the skeletal hand located between the phalanges of the fingers and the carpal bones of the wrist, which forms the connection to the forearm. The metacarpal bones ar ...
s. A strongly derived condition show the
phalanges The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones ...
, small, disc-shaped elements positioned in long rows. Sometimes, the number of fingers is reduced, to as low as two. This is a rather common phenomenon within the Tetrapoda. Unique, however, for derived tetrapods, is the fact that some species show nonpathological
polydactyly Polydactyly or polydactylism (), also known as hyperdactyly, is an anomaly in humans and animals resulting in supernumerary fingers and/or toes. Polydactyly is the opposite of oligodactyly (fewer fingers or toes). Signs and symptoms In human ...
, the number of fingers being higher than five. Some species have ten fingers per hand. These fingers, again, can have an increased number of phalanges, up to thirty, a phenomenon called hyperphalangy, also known from the Plesiosauria,
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the ' Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht o ...
s, and the
Cetacea Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel the ...
. The high number of elements allows the flipper to be shaped as a
hydrofoil A hydrofoil is a lifting surface, or foil, that operates in water. They are similar in appearance and purpose to aerofoils used by aeroplanes. Boats that use hydrofoil technology are also simply termed hydrofoils. As a hydrofoil craft gains sp ...
. When a high number of fingers is present, their identity is difficult to determine. It is usually assumed that fingers were added at both the front and at the rear, perhaps to a core of four original fingers. If fingers are added, often the number of metacarpals and carpals is also increased; sometimes even an extra lower arm element is present. Earlier, ichthyosaurs were commonly divided into "longipinnate" and "latipinnate" forms, according to the long or wide shape of the front flippers, but recent research has shown that these are not natural groups; ichthyosaur clades often contain species with and without elongated forelimbs. The ichthyosaur
pelvis The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
is typically rather reduced. The three pelvic bones: the ilium, the
ischium The ischium () form ...
, and the
pubic bone In vertebrates, the pubic region ( la, pubis) is the most forward-facing (ventral and anterior) of the three main regions making up the coxal bone. The left and right pubic regions are each made up of three sections, a superior ramus, inferior r ...
, are not fused and often do not even touch each other. Also, the left and right pelvic sides no longer touch; only basal forms still have sacral ribs connecting the ilia to the vertebral column. The hip joint is not closed on the inside. The pubic bone typically does not connect to the ischium behind it; the space in between is by some workers identified as the ''fenestra thyreoidea''; other researchers deny that the term is applicable given the general loose structure of the pelvis. Some later species have a connected pubic bone and ischium, but in this case, the femoral head no longer articulates with the hip joint. Triassic species have plate-like pubic bones and ischia; in later species these elements become elongated with a narrow shaft and can form a single rod. Typically, the hindlimbs are shorter than the forelimbs, possessing a lesser number of elements. Often, the rear flipper is only half the length of the front flipper. The thighbone is short and broad, often with a narrow waist and an expanded lower end. The tibia, fibula and
metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the me ...
s are merged into a mosaic of bone discs supporting the hydrofoil. Three to six toes are present. The toe phalanges also show hyperphalangy; exceptionally, ''
Ophthalmosaurus ''Ophthalmosaurus'' (meaning "eye lizard" in Greek) is an ichthyosaur of the Jurassic period (165–150 million years ago). Possible remains from the Cretaceous, around 145 million years ago, are also known. It was a relatively medium-sized ichth ...
'' shows a reduced number of phalanges.


Soft tissue

The earliest reconstructions of ichthyosaurs all omitted dorsal fins and caudal (tail) flukes, which were not supported by any hard skeletal structure, so were not preserved in many fossils. Only the lower tail lobe is supported by the vertebral column. In the early 1880s, the first body outlines of ichthyosaurs were discovered. In 1881, Richard Owen reported ichthyosaur body outlines showing tail flukes from Lower Jurassic rocks in Barrow-upon-Soar, England. Other well-preserved specimens have since shown that in some more primitive ichthyosaurs, like a specimen of '' Chaohusaurus geishanensis'', the tail fluke was weakly developed and only had a dorsal tail lobe, making the tail more paddle-like. Over the years, the visibility of the tail lobe has faded away in this specimen. The presence of dorsal fins in ichthyosaurs has been controversial. Finely preserved specimens from the Holzmaden ''
Lagerstätte A Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These for ...
n'' in Germany found in the late 19th century revealed additional traces, usually preserved in black, of the outline of the entire body, including the first evidence of dorsal fins in ichthyosaurs. Unique conditions permitted the preservation of these outlines, which probably consist of bacterial mats, not the remains of the original tissues themselves. In 1987,
David Martill {{Short pages monitor