Cymbospondylus

''Cymbospondylus'' (meaning "cupped vertebrae") is an extinct genus of large ichthyosaurs, of which it is among the oldest representatives, that lived from the Lower to Middle Triassic in what are now North America and Europe. The first known fossils of this taxon are a set of more or less complete vertebrae which were discovered in the 19th century in various mountain ranges of Nevada, in the United States, before being named and described by Joseph Leidy in 1868. It is in the beginning of the 20th century that more complete fossils were discovered through several expeditions launched by the University of California, and described in more detail by John Campbell Merriam in 1908, thus visualizing the overall anatomy of the animal. While many species have been assigned to the genus, only five are recognized as valid, the others being considered synonymous, doubtful or belonging to other genera. ''Cymbospondylus'' was formerly classified as a representative of the Shastasauridae, but more recent studies consider it to be more basal, view as the type genus of the Cymbospondylidae.

As an ichthyosaur, ''Cymbospondylus'' had flippers for limbs and a fin on the tail. Like other non-parvipelvian ichthyosaurs, ''Cymbospondylus'' has a very slender profile, unlike later ichthyosaurs which have a morphology similar to those of dolphins. The different species of ''Cymbospondylus'' vary greatly in size, with the smallest reaching around in length. The largest known species, ''C. youngorum'', is estimated over long, making ''Cymbospondylus'' one of the largest ichthyosaurs identified to date, but also one of the largest animals known of its time. The animal has a skull with a long, thin snout, proportionally small eye sockets, an elongated trunk, and a less pronounced tail than in later ichthyosaurs. The teeth are conical and pointed, having longitudinal ridges, indicating a diet of fishes and cephalopods, and possibly other marine reptiles for larger species.

Unlike cetaceans, Triassic ichthyosaurs like ''Cymbospondylus'' show that they reached large sizes very quickly after their appearance, probably because of the adaptive radiation of their prey, conodonts and ammonites, after the Permian–Triassic extinction. The size of ichthyosaurs began to decrease later, notably due to the increase in the size of their eyes, which were very useful for spotting prey. All established species of ''Cymbospondylus'' are known from the fossil records of Nevada and Switzerland, with referred specimens without specific affiliations having nevertheless been discovered in Idaho, the rest of the Alps and Spitsbergen, an island in Norway. The formations where the recognized species were discovered show that ''Cymbospondylus'' lived in marine ecosystems alongside molluscs like bivalves and ammonites, bony fishes like ''Saurichthys'' and coelacanths, cartilaginous fishes like hybodonts, and marine reptiles like sauropterygians and other ichthyosaurs. The different ichthyosaurs from these localities would have used different feeding strategies to avoid competition.

Research history

Discovery and identification

In 1868, American paleontologist Joseph Leidy described two new genera of ichthyosaurs dating from the Middle Triassic on the basis of fossil vertebrae discovered in several localities in Nevada, United States, all of which were transmitted through the geologist Josiah Dwight Whitney. One of the two genera he named is ''Cymbospondylus'', to which he assigned two species. The first one is ''C. piscosus'', which Leidy named on the basis of several more or less complete vertebrae having been discovered in different mountain ranges of the state. The holotype of ''C. piscosus'' is a block containing five incomplete dorsal vertebrae that was discovered in the New Pass Range, northwest of the city of Austin. Leidy attributes two other specimens to the species, one being a series of eight caudal vertebrae discovered at Star Canyon in the Humboldt Range, and the other being a single vertebra, probably also caudal, discovered in the Toiyabe Range, in the Reese River, northeast of Austin. The second species is ''C. petrinus'', named from five large dorsal vertebrae discovered in the Humboldt Range. The genus name ''Cymbospondylus'' derives from the Ancient Greek words κύμβη (''kymbē'', "cup") and σπόνδυλος (''spondylos'', "vertebra"), all taken together literally meaning ''cupped vertebrae'', in reference to the rather obvious shape of this part of the skeleton. As no type species was designated in the 1868 article, it was not until 1902 that John Campbell Merriam assigned ''C. piscosus'' to this title, the latter being the first named in Leidy's official description of the genus.

Between 1901 and 1907, the University of California sent ten expeditions across different corners of the United States to recover as many ichthyosaur fossils as possible dating from the Triassic, in order to be analyzed in more detail. These same expeditions were led by Merriam and were almost all financed by Annie Montague Alexander. The research finally collected more than fifty specimens, each preserving a significant portion of their skeletons. Among these fossils are several specimens of excellent quality from ''C. petrinus'', including an almost complete skeleton, cataloged as UCMP 9950, all discovered in the origin locality mentioned by Leidy. Like the other ichthyosaurs mentioned in the work, Merriam describes the taxon in more depth based on new known fossil material. Merriam's anatomical descriptions of ''C. petrinus'' are still recognized as viable, and are even used in comparative anatomy in later studies of the genus ''Cymbospondylus'', thus lending validity to the species.

Unlike ''C. petrinus'', no additional fossils of the type species ''C. piscosus'' have been discovered. In Merriam's works, the latter recognized ''C. piscosus'' as distinct on the basis of its smaller size and by the regularly concave faces of the vertebrae of the holotype specimen. Only later did the validity of ''C. piscosus'' begin to be questioned, with authors mentioning the questionable nature of the fossils. In their work published in 2003, Christopher McGowan and Ryosuke Motani assert that the fossil vertebrae attributed to ''C. piscosus'' do not present distinctive characteristics to prove its validity. This would therefore pose a taxonomic problem, because if the type species turns out to be a ''nomen dubium'', the genus to which it is classified will also be. To try to resolve this problem, the two authors suggest that the most complete known skeleton of ''C. petrinus'' (UCMP 9950) could be designated as a neotype of ''C. piscosus''. However, as no formal ICZN appeal has been established to date, the name ''C. petrinus'' for the proposed neotype should be retained until further notice. Therefore, ''C. piscosus'' is not included in most descriptions of the genus, although it is still recognized as the type species.

Recognized species

In 1927, a partial skeleton of a large ichthyosaur was discovered in the Grenzbitumenzone Member at the Monte San Giorgio fossil site in Switzerland and was mentioned by Bernhard Peyer in 1944. Twenty years later, in 1964, published a photo of this same specimen and suggested that it shared affinities with ''Cymbospondylus'', then only known from North America at that time. The specimen in question, cataloged as PIMUZ T 4351, was formally described for the first time in 1989 under the name ''C. buchseri'' by , thus confirming the presence of the genus in Europe. The specific epithet ''buchseri'' is named in honor of Fritz Buchser, a member of the Museum of Paleontology at the University of Zurich, the latter having prepared the holotype skeleton in 1931 as his first major professional achievement.

While ''C. petrinus'' was for a time seen as the only valid species of the genus known from Nevada, it was not until the early 21th century that later discoveries contradicted this assertion. In 2006, Nadia Fröbisch and colleagues described the species ''C. nichollsi'' based on an incomplete skeleton, cataloged as FMNH PR 2251, which was discovered in the Augusta Mountains. The fossil was originally exhumed in the hope of finding a new specimen of ''C. petrinus'', but the number of significant anatomical differences led researchers to establish a separate species. The species in question is named in honor to Elizabeth Nicholls, an American-Canadian paleontologist specializing in Triassic marine reptiles, who made a major contribution to the ichthyosaurs that lived during this period. In their description, Fröbisch and his colleagues consider that an almost complete skull attributed to ''C. petrinus'', cataloged as UCMP 9913, could in fact belong to ''C. nichollsi'', because it presents similar characteristics. However, as the specimen has been relatively little described in the scientific literature, the authors do not know whether it would show intraspecific variations within ''C. petrinus'', judging therefore that a more in-depth description is necessary. Subsequent studies carried out on the genus ''Cymbospondylus'' nevertheless always refer specimen UCMP 9913 to ''C. petrinus'', but still mentioning some notable differences.

In 2011, a notable new ''Cymbospondylus'' fossil was also discovered in the Augusta Mountains, then exhumed three years later in 2014. This discovery was a partial skeleton that was briefly mentioned in a 2013 in a secondary article describing the large contemporary ichthyosaur ''Thalattoarchon'', as well as in a 2018 histological study, where it is among the specimens analyzed. However, it was only later that the specimen, cataloged as LACM DI 158109, was formally designated a holotype for the species ''C. duelferi'' by Nicole Klein and colleagues in 2020. The species name ''duelferi'' was chosen in honor of Olaf Dülfer, fossil preparer who made many practical contributions to research on Mesozoic marine reptiles.

In 1998, still in the Augusta Mountains, Sander discovered another notable specimen of ''Cymbospondylus'', and exhumed it with his colleagues between 2014 and 2015. After preparation of the fossils, the specimen, cataloged as LACM DI 157871, consists of a large complete skull, some cervical vertebrae, the right humerus as well as fragments of the shoulder girdle. It was in 2021, one year after the identification of ''C. duelferi'', that a new species of the genus was named from this specimen in ''Science'' by Sander and his colleagues. This species, ''C. youngorum'', is named in honor of Tom and Bonda Young, these latter having financially supported the fossil exhumation project.

Specimens that may belong to the genus

Many other more partial specimens of ''Cymbospodylus'' have been discovered in various geological formations in Europe, but their specific attribution cannot be determined, the latter are then referred to under the name ''Cymbospondylus'' sp. in the scientific literature. Three of these specimens, including one from Idaho, and two from the Norwegian archipelago of Svalbard, are dated to the Olenekian stage of the Lower Triassic, making them the oldest known representatives of the genus. Below, the list of specimens that could potentially belong to ''Cymbospondylus'':

* In 1980, Kuhn-Schnyder described an anterior partial skeleton of an ichthyosaur discovered in , Italy. The described specimen was first referred by the author to '' Shastasaurus'' sp., before being referred to ''Cymbospondylus'' in the official description of ''C. buchseri'' by Sander in 1989, an attribution which seems to still be recognized today.

* In 1992, Sander described two specimens attributed to ''Cymbospondylus'' having been discovered in different localities of Spitsbergen, located in the Norwegian archipelago of Svalbard. The first and the better preserved of the two specimens described in the article, consists of a series of 17 vertebrae associated with ribs which was discovered in 1961 in the Botneheia mountain, being cataloged as PIMUZ A/III 496. The second consists of two isolated vertebrae, cataloged as PIMUZ A/III 554 and 555, which were both discovered in the bay of Wichebukta.

* In 1994, Judy A. Massare and Jack M. Callaway referred to a number of ''Cymbospondylus'' fossils discovered in 1985 by H. Gregory McDonald in the Platy Siltstone Member of the Thaynes Formation, in Idaho, United States.

* In 2001, Olivier Rieppel and Fabio Marco Dalla Vecchia listed a set of fossils of marine reptiles from the Triassic and having been discovered in the ''comune'' of Forni di Sotto, in Italy, including two that they attributed with doubt to ''Cymbospondylus''. The first of these collections consists of a single vertebra, a neural spine and three rib fragments, while the second consists of two isolated vertebrae.

* In 2012, Balini and Renesto described four more or less partial vertebrae attributed to ''Cymbospondylus'' which were discovered in the ''comune'' of Piazza Brembana, in Italy, being cataloged MCSNB 11689 A, B, C, and D.

* In 2013 and 2018, numerous genus-assigned vertebrae were identified in the Vendomdalen Member of the Vikinghøgda Formation, Svalbard.

Formerly assigned species

Although many valid and distinct species have been assigned to ''Cymbospondylus'' throughout its taxonomic history, some of these have been reassigned to different genera or are considered synonymous or even doubtful. In his 1868 paper describing ''Cymbospondylus'', Leidy also named another ichthyosaur as ''Chonespondylus grandis'', based on a fragment of a caudal vertebra found at Star Canyon in the Humboldt Range. The genus name ''Chonespondylus'' derives from the Ancient Greek words χοάνη (''khoánē'', "funnel") and σπόνδυλος (''spondylos'', "vertebra") named in the same way as for ''Cymbospondylus''. The specific epithet comes from the Latin ''grandis'', meaning "large, wide". In 1902, Merriam listed Leidy's discoveries, but having found no features distinguishing ''Chonespondylus'' from ''Cymbospondylus'', he decided to synonymize the first name with the second, under the name ''C. (?) grandis''. In 1908, after the discovery of new very complete fossils from ''C. petrinus'', Merriam decided to definitively synonymize ''C. (?) grandis'' with the latter.

In 1873, John Whitaker Hulke described a species of ''Ichthyosaurus'', ''I. polaris'', named after two sets of vertebrae associated with rib fragments that were discovered on Isfjorden, Spitsbergen, an island in Norway. In 1902, the Russian paleontologist Nikolai Nikolajewitch Yakowlew moved the species within the genus '' Shastasaurus'', referring an isolated vertebra to this taxon. In 1908, Merriam in turn moved this species into the genus ''Cymbospondylus'', under the name ''C. (?) polaris''. Merriam still expresses some hesitation about this attribution, asserting that the true generic identity cannot be determined for this species due to the few known fossils. In 1910, the species was moved to the newly erected genus '' Pessosaurus'' by Carl Wiman, as ''P. polaris'', to which it has always been referred by this name ever since. Although this taxon is declared as a ''nomen dubium'' according to studies published at the end of the 20th century, it is seen as a ''species inquirenda'' according to McGowan and Motani in 2003, i.e. a taxon under investigation, as numerous fossils that have since been referred to ''P. polaris'' could make it once again as valid.

Still in his 1908 work, Merriam erects two new species of the genus, coming from the same locality from which ''C. petrinus'' is known. The first is ''C. nevadanus'', named from fossils constituting a hind limb. Merriam distinguishes this species from ''C. petrinus'' on the basis of its larger size and the different proportions of some bones. However, the ''C. nevadanus'' material is not sufficiently diagnostic to support the validity of this species, and is considered a ''species inquireda'' according to McGowan and Motani in 2003. The second species erected by Merriam is ''C. natans'', which he names from an isolated humerus, to which he attributes a radius, an ulna, carpals and a series of caudal vertebrae. In his article, he notes the similarity of these bones with those of '' Mixosaurus'', leading the author to rename the species to ''M (?) natans'' in 1911. For much of the 20th century, ''M (?) natans'' was recognized as a valid species of ''Mixosaurus'' until 1999, when it was synonymized with ''M. nordenskioeldii''. Although ''M. nordenskioeldii'' itself has been considered a ''nomen dubium'' since 2005, the fossil material concerned remains attributed to the family Mixosauridae and is no longer attributable to ''Cymbospondylus''.

In a review of German ichthyosaurs published in 1916, Friedrich von Huene described two other species of ''Cymbospondylus'' whose fossils were discovered in the Muschelkalk of the town of Laufenburg in the state of Baden-Württemberg. The first is ''C. germanicus'', which Huene names from a single vertebra associated with other vertebrae as well as a basioccipital. Immediately afterwards, the validity of ''C. germanicus'' was questioned the same year by Ferdinand Broili, the latter citing that the fossils concerned did not present notable features to be recognized as distinct. Additionally, the fossils appear to be very poorly preserved to be distinguished as a valid species, and is therefore a ''nomen dubium''.

In 2002, paleontologists Chun Li and Hai-Lu You named a new species as ''C. asiaticus'' based on a complete skull discovered in the Xiaowa Formation, located in Guizhou Province, China, and it was considered as the most recent known representative of the genus. In the official description of ''C. nichollsi'' published in 2006, the authors are doubtful regarding the attribution of this species to ''Cymbospondylus''. They mention that the latter does not share any notable commonalities with the three then recognized species of the genus at the time, namely ''C. petrinus'', ''C. buchseri'' and ''C. nichollsi'', and suggest that it would in fact be a junior synonym of '' Guizhouichthyosaurus tangae''. The synonymy proposed by Fröbisch and colleagues was accepted in 2009 by Qing-Hua Shang and Li after the discovery of an almost complete skeleton of ''Guizhouichthyosaurus'' from the same formation. However, considering that ''Guizhouichthyosaurus'' is similar to ''Shastasaurus'', they moved the species as ''S. tangae''. This synonymy was contested the following year, in 2010, in which Michael W. Maisch provisionally reclassified ''Guizhouichthyosaurus'' as a distinct genus. In 2011, Sander and his colleagues considered that ''Guizhouichthyosaurus'' was distinct, while a 2013 study by Shang and Li still synonymizes it with ''Shastasaurus''. However, numerous phylogenetic and morphological analyzes subsequently published consider ''Guizhouichthyosaurus'' to be distinct genus from ''Shastasaurus''.

Description

''Cymbospondylus'', like other ichthyosaurs, had a long, thin snout, large eye sockets, and a tail fluke that was supported by vertebrae in the lower half. Ichthyosaurs were superficially similar to dolphins and had flippers rather than legs, but the oldest representatives (with the exception of the parvipelvians, more advanced) do not have dorsal fins, or would have one but relatively poorly developed. Like most Triassic ichthyosaurs, ''Cymbospondylus'' has a more slender anatomy, possessing an elongated trunk and a long, poorly pronounced tail. Although the colour of ''Temnodontosaurus'' is unknown, at least some ichthyosaurs may have been uniformly dark-coloured in life, which is evidenced by the discovery of high concentrations of eumelanin pigments in the preserved skin of an early ichthyosaur fossil.

Size

The size and weight of ''Cymbospondylus'' varies greatly between recognized species. Estimates of the size of ''C. petrinus'' have changed relatively little since 1908, mainly due to the almost complete skeleton of specimen UCMP 9950, considered the best known specimen of ''Cymbospondylus''. Merriam suggests that ''C. petrinus'' would reach a size exceeding in length based on specimens UCMP 9947 and 9950. In 2020, Klein and colleagues increased the size estimate of ''C. petrinus'' a little further, seeing them as reaching for a skull. In 2021, Sander and is colleagues in 2021 estimates ''C. petrinus'' to long for about , while keeping the same cranial measurements. The explanation for the origin of this size is described in a paper published in 2024. The study explains that Sander made a further revision regarding the size of ''C. petrinus'' based on specimens UCMP 9947 and 9950, and suggested that the combination of the two gives a total of approximately . Specimen UCMP 9947 is missing several posterior caudal vertebrae, so the increase in size to over long is not seen as unreasonable. The holotype specimen of ''C. youngorum'' having a skull measuring almost long and its humerus being the second largest bone of this type recorded in an ichthyosaur, the maximum size of the animal is therefore estimated at for a weight of . This estimate not only makes ''C. youngourum'' one of the largest ichthyosaurians identified to date behind ''Ichthyotitan'', but also makes it one of the largest animals known of its time. The antiquity as well as such an imposing size for an animal dating from the beginning of the Middle Triassic makes ''C. youngorum'' qualify as "the first aquatic giant" according to Lene Liebe Delsett and Nicholas David Pyenson.

Estimating the size of ''C. nichollsi'' is more complex, because although the holotype specimen is known from a significant portion of the skeleton, the latter preserves only half of the skull. However, based on the comparison with ''C. buchseri'' and ''C. petrinus'', the total length of the skull is estimated at for an animal measuring long, all for a body mass of . The estimated size of the ''C. buchseri'' holotype is slightly shorter, reaching a length of with a skull long, although no estimate of its weight has yet been published. Possessing a skull which would measure a total of , ''C. duelferi'' is the smallest known species of the genus, having a size estimated at long in the study officially describing it. The measurements of the species are estimated again at around for a body mass of by Sander and colleagues in 2021.

Individuals with undetermined specific attributions have also been given estimates regarding their size, although known from thinner fossil remains. Using the same measurement technique as those done for ''C. youngorum'', a specimen cataloged as IGPB R660, known from the Vikinghøgda Formation, has an estimated size between long, making it the largest known ichthyosaur specimen from the Lower Triassic.

Skull

Like other ichthyosaurs, the snout of ''Cymbospondylus'' is elongated into a long, conical rostrum, the largest bones of which are the premaxillae and nasal bones. The nasal bones also extend far back, and, with the frontal bones, reach the anterior edge of the temporal fenestrae. The eye sockets are oval to ovoid in shape and are proportionally small in relation to the size of the skull. The shape of the superior temporal fenestra differs between some species, being oval in ''C. nichollsi'' and ''C. duelferi'' but triangular in ''C. petrinus''. The number of bony elements constituting the sclerotic rings varies between species :
12 ossicles in ''C. duelferi''; 13 ossicles in ''C. buchseri''; 14 ossicles in ''C. nichollsi''; and between 14 and 18 ossicles in ''C. petrinus''. Although a sclerotic ring is preserved in ''C. youngorum'', nothing has been said about the number of ossicles constituting it in the latter. The sagittal crest is more or less pronounced in the different species, being low in ''C. duelferi'' and ''C. nichollsi'', clearly high in ''C. petrinus'', and totally absent in ''C. youngorum''. The occipital condyle of ''Cymbospondylus'' is concave in shape. Like other ichthyosaurs, ''Cymbospondylus'' has an elongated and thin lower jaw, extending backwards to beyond the back of the skull. The dentary, the main bone making up the lower jaw, extends almost to the level of the middle of the eye socket. The surangular also represents an important part of the mandible, and thins down to the retro-articular process. The angular bone forms the ventral part of the lateral side of the lower jaw and contacts the surangular via a long suture.

The dentition of ''Cymbospondylus'' is generally thecodont, meaning that the tooth roots are deeply cemented into the jawbone. However, not all species share the same robustness in terms of their dental implantation. ''C. petrinus'' has a particular form of thecodont dentition, its teeth appearing to be fused at the bottom of the alveoli. ''C. duelferi'' has a subthecodont type of dentition, meaning that its teeth are implanted in shallow sockets. ''C. youngorum'' has a thick base of the bone attaching to the teeth, a trait not seen in other ichthyosaurs. The teeth are homodont, that is to say they share an identical shape, being conical, ridged and pointed. The teeth also have longitudinal ridges that extend from the base of the crown to the apical third. The total number of teeth in the different species of ''Cymbospondylus'' is difficult to determine, because the state of preservation of certain fossils prevents formal evaluations from being obtained, being poorly preserved in ''C. buchseri'', and totally unknown in ''C. nichollsi''. Only three species were able to have a more or less clear estimate of their number of teeth : ''C. duelferi'' having a number greater than 21 teeth known in the upper jaw, but unknown in the lower jaw; ''C. petrinus'' with between 30 and 35 teeth in the upper and lower jaws; and ''C. youngorum'' having 43 teeth in the upper jaw and more than 31 teeth in the lower jaw.

Classification

Phylogeny

The exact placement of ''Cymbospondylus'' within Ichthyosauria is poorly understood with its position varying between different studies, sometimes being recovered as more and sometimes as less derived than mixosaurids. However it is agreed upon that ''Cymbospondylus'' is a rather basal member of the clade. Early phylogenies placed ''Cymbospondylus'' within Shastasauridae. In the analysis of Bindellini ''et al.'' (2021)'', Cymbospondylus'' is placed at the very base of Ichthyosauria, outside the more derived members of Hueneosauria (including Mixosauridae and Shastasauridae). In the publication describing ''C. duelferi'', Klein and colleagues recovered that all species from the Fossil Hill Member in Nevada form a clade with one another. The description of ''C. youngorum'' further supports this Nevadan clade, recovering ''C. youngorum'' as its most derived member while ''C. buchseri'' from Europe sits at the base of the genus. Much like in the analysis by Bindellini and colleagues, shastasaurids and mixosaurids were recovered as more derived ichthyosaurs.

Like in many analyses prior, the type species was not included in the dataset due to its questionable and fragmentary nature. This causes ''Cymbospondylus'' to have a very convoluted taxonomy, with it being suggested that the type species should be neglected. The 2020 study reviewed the skull morphology of ''C. nichollsi'' and found the species to be valid, as the skull morphology accords with that of ''C. petrinus'' but is distinct enough to be separate, such as the upper temporal fenestra shape being oval in ''C. nichollsi'' but triangular in ''C. petrinus''. In their phylogenetic analysis the authors did not recover a definite placement for ''C. buchseri'', leading them to state that further study was needed to determine whether the Swiss species belonged to the genus.

The following cladogram shows the position of ''Cymbospondylus'' within the Ichthyosauria after Sander ''et al.'', (2021):

Evolution

Ichthyosaurs form one of the major groups of marine reptiles that flourished during a large part of the Mesozoic, between approximately 248 and 90 million years ago, i.e. during the end of the Lower Triassic until approximately the beginning of the Upper Cretaceous. Appearing in the early temporal stages of this group, ''Cymbospondylus'' is therefore one of the oldest representatives to have been identified to date. However, despite its age, the genus shows that certain ichthyosaurs adopted a rapid increase in size throughout their evolution. Indeed, the oldest known representatives of ichthyosauriforms (a group that includes ichthyosaurs, their ancestors and related lineages), such as ''Cartorhynchus'', have a skull measuring long, while the largest known species of ''Cymbospondylus'', ''C. youngorum'', has a skull up to about long, and yet these two taxa are only separated by about 2.5 million years. To compare with the evolution of a group of similar tetrapods, namely the cetaceans, between ''Pakicetus'', which has a skull width of , and ''Basilosaurus'', to which the latter has a skull width of , between 10 and 14 million years ago. A similar case is also observed in the subgroup of odontocetes, because between '' Simocetus'', which has a skull wide of , and ''Livyatan'', which has a skull wide of approximately , approximately more than 25 million years ago. This rapid increase in size among ichthyosaurs could have been favored by the rapid diversification of conodonts and ammonites after the Permian–Triassic extinction event. The evolution of large eyes would subsequently have considerably reduced the large measurements in ichthyosaurs, because they helped better identify their source of food.

Palaeobiology

Massare & Callaway (1990) propose that many Triassic ichthyosaurs including ''Cymbospondylus'' may have been ambush predators. They argue that the long neck and torso would create drag in water while the laterally-flattened tail lacking the lunate fluke of later ichthyosaur taxa was more suited for an undulating swimming style. In their research they suggest that the elongated flexible bodies of early ichthyosaurs were built to support an undulating swimming style while the powerful tail would provide bursts of speed, both of which they cite as being possible adaptations to ambush prey. Massare & Callaway put this in contrast with Jurassic taxa, known for their compact, dolphin-like bodies adapted for more continuous swimming favorable to pursuit predators. A strikingly similar bauplan was later obtained by two other large bodied marine amniote groups, mosasaurs and archaeocete whales.

Direct evidence for its diet exists for the medium-sized ''Cymbospondylus buchseri'' from Switzerland, which was found with its stomach contents exclusively consisting of hooks belonging to soft-bodied coleoid cephalopods. However, this does not exclude the possibility that ''C. buchseri'' could have taken larger prey, as its last meal may not reflect its typical diet accurately. Bindellini and colleagues suggest that ''C. buchseri'' may have employed a more forceful feeding strategy with a slower feeding cycle and a higher biteforce, supported by the animal's robust rostrum. In the Besano Formation, ''Cymbospondylus'' would have coexisted with two other smaller ichthyosaurs, the more gracile skulled ''Besanosaurus'' and small mixosaurs. Whether or not ''C. buchseri'' would have gone after large vertebrate prey, all three taxa display clear adaptations for different hunting strategies and prey preferences, however the details of their ecologies are not yet fully understood.

For ''C. youngorum'' a generalist diet of squid and fish is inferred based on the blunt and conical teeth in combination with the elongated rostrum. However, as with ''C. buchseri'', Sander ''et al.'' entertain the possibility that ''C. youngorum'' could have fed on large-bodied vertebrates as well, including the other ''Cymbospondylus'' species of the region.

Bindellini and colleagues notes that shastasaurid diversity may have profited from the extinction of ''Cymbospondylus'', such as the Carnian of China, known to have supported three ecologically different shastasaurids but no examples of cymbospondylids, which had gone extinct by that time.

Reproduction

The holotype of ''C. duelferi'' preserves three small strings of articulated vertebrae located within the trunk region of the specimen. These vertebrae, which are only a third the size of the adult specimen, have been interpreted to represent the remains of three fetuses, with one specimen specifically facing towards the rear end of the putative mother. Following this interpretation, ''Cymbospondylus'' would have given live birth to a minimum of three offspring.

Paleoecology

North America

All North American species of ''Cymbospondylus'' are known from fossils found in two geologic formations in the Star Peak Group, located in Nevada. ''C. petrinus'', ''C. nichollsi'', ''C. duelferi'' and ''C. youngorum'' are known from the Favret Formation, but the first named species is the only known representative of the genus who have been identified in the Prida Formation. These two formations are linked by a single member, known as the Fossil Hill Member. In the Prida Formation, this member outcrops west of the Humboldt Range, and extends to the Favret Formation, outcropping the Augusta Mountains, where it reaches up to more than 300 m wide. Although they are neighbors, the two formations do not share precisely the same age, the Prida one dating from the Middle Anisian, while Favret dates from the Late Anisian, between approximately 244 and 242 million years ago.

The significant presence of marine reptiles, ammonites and other invertebrates in the Fossil Hill Member indicates that the surface waters were well aerated, but there is however little animal presence in the benthic zones, with the notable exception of bivalves of the Halobiidae family. The fossils found show that the rock unit was once a pelagic ecosystem with a stable food web. Bony fish are little known and have currently only been discovered in the Favret Formation. Among the fish discovered, we find the actinopterygians ''Saurichthys'' and an undetermined representative, while among the sarcopterygians, numerous specimens of indeterminate coelacanthids are known. In the Prida Formation, a significant number of cartilaginous fishes have been identified. These include the hybodontiforms, one synechodontiform, and problematically positioned elasmobranchians. The most abundant marine reptiles of the Fossil Hill Member are the ichthyosaurs, including ''Cymbospondylus'' itself, '' Omphalosaurus'', '' Phalarodon'' and the large ''Thalattoarchon''. Few other marine reptiles are known from the Fossil Hill Member, the only clearly identified being the sauropterygian ''Augustasaurus''.

Europe

The only currently known specimen of ''C. buschseri'' is recorded from the Besano Formation, also known as the Grenzbitumenzone in Switzerland. This formation is located in the Alps and extends from southern Switzerland to northern Italy, containing numerous fossils dating from between the end of the Anisian and the beginning of the Ladinian. This formation is one of a series of Middle Triassic units atop a carbonate platform at Monte San Giorgio, and measures thick. During the time when the animal lived, when the Besano Formation was being deposited, the region where Monte San Giorgio is would have been a marine lagoon, located in a basin on the western side of the Tethys Ocean. Researchers estimate that this same lagoon would have reached between deep. The near-surface waters would have been well oxygenated and were inhabited by a wide range of plankton and free-swimming organisms. However, water circulation within the lagoon was poor, resulting in typically anoxic water at the bottom, deprived of oxygen. The lagoon bottom would have been quite calm, as evidenced by the fine lamination of the rocks, and there is little evidence of bottom-dwelling organisms modifying the sediment. The presence of terrestrial fossils, such as conifers and land-dwelling reptiles indicates that the region would have been near land.

Among the most common of the invertebrates from the Besano Formation is the bivalve '' Daonella''. Many gastropods are known from the Besano Formation; predominantly those that could have lived as plankton or on algae. Cephalopods present include nautiloids, coleoids, and the especially common ammonites. The coleoids from the Besano Formation are not particularly diverse, but this may be due to their remains not readily fossilizing, with many of their known remains being preserved as stomach contents within the bodies of ichthyosaurs. Arthropods known from the formation include ostracods, thylacocephalans, and shrimp. Other, rarer invertebrate groups known from the formation include brachiopods and echinoids, which lived on the seabed. Radiolarians and macroalgae are also known in the formation, though the latter may have been washed in from elsewhere, as with many other bottom-dwelling organisms. A very large number of bony fish have been recorded in this formation. Many bony fish have been recorded in this formation, with actinopterygians being quite diverse, including abundant small species as well as larger representatives like ''Saurichthys''. Among the sarcopterygians, the number is more limited with in particular '' Rieppelia'', '' Ticinepomis'' and possibly '' Holophagus'', which are all coelacanthiforms. cartilaginous fishes of the Besano Formation are uncommon as well and mainly consist of hybodonts.

Unlike the Fossil Hill Member in Nevada, ichthyosaurs do not represent the most diverse marine reptiles in the Besano Formation, the latter being limited only to ''Besanosaurus'', ''C. buchseri'', ''Phalarodon'' and ''Mixosaurus'', their abundance in the middle part of this zone correlating with the time when the lagoon was deepest. Conversely, the sauropterygians represent the largest part of the assemblage of marine reptiles of this formation. Among these are the shell-crushing placodonts ''Paraplacodus'' and '' Cyamodus''. as well as pachypleurosaurs and nothosaurids. The pachypleurosaur '' Odoiporosaurus'' is known from the middle Besano Formation, while the particularly abundant '' Serpianosaurus'' did not appear until the upper portion of the formation, where ichthyosaurs are becoming rarer. Nothosaurids include the genera '' Silvestrosaurus'' and ''Nothosaurus'', the latter notably including ''N. giganteus'' and possibly ''N. juvenilis''. While rare, ''N. giganteus'' may have been an apex predator like ''C. buschseri''. Apart from ichthyosaurs and sauropterygians, other marine reptiles include the long-necked ''Tanystropheus'' and '' Macrocnemus'', and the thalattosaurians '' Askeptosaurus'', '' Clarazia'' and '' Hescheleria''.

Niche partitioning

In both the Fossil Hill Member and the Besano Formation, ''Cymbospondylus'' is one of a variety of ichthyosaurs. The different species known would have had different feeding strategies to avoid competition. Due to its large and sharp teeth, ''Thalattoarchon'' would probably have been the only apex predator with which ''Cymbospondylus'' was contemporary, probably attacking smaller marine reptiles, or even juveniles. ''Besanosaurus'' would likely have specialized in feeding on coleoids, based on the shape and small size of its teeth. The stomach contents of ''Mixosaurus cornalianus'' show the remains of small coleoids and fish, suggesting that it would have gone after smaller prey than its larger relatives. The rarer mixosaurids ''Mixosaurus kuhnschnyderi'' and ''Phalarodon'' both possess broad crushing teeth. ''M. kuhnschnyderi'' is understood to have consumed coleoids, while the larger teeth of ''Phalarodon'' may have been suited for crushing prey items with external shells. ''Omphalosaurus'' was probably a bulk feeder specialized in grinding up ammonites.

Extinction

In 2021, Gabriele Bindellini and colleagues note that shastasaurid diversity may have benefited from the extinction of ''Cymbospondylus'', as shown in the Carnian fossil record of China, known to have three ecologically different shastasaurids, but no examples of cymbospondylids, being extinct at this time.

Notable appearances in media

A ''Cymbospondylus'' is present in the 2003 BBC docufiction ''Sea Monsters'', and more precisely in a sequence featuring various marine reptiles of the Triassic. In the only scene in which it appears, the latter grabs by surprise a torn tail of a ''Tanystropheus'', until then held by Nigel Marven, before the animal appears threatening towards the presenter.

See also

* List of ichthyosaurs
* Timeline of ichthyosaur research

Notes

References

Bibliography

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External links

; Paleontological videos
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{{Taxonbar, from=Q133027
Middle Triassic ichthyosaurs
Early Triassic ichthyosaurs
Middle Triassic reptiles of Europe
Middle Triassic reptiles of North America
Early Triassic reptiles of Europe
Ichthyosauromorph genera
Taxa named by Joseph Leidy
Apex predators