Besanosaurus - Knowledge

1142:(vertebral bodies, some of which were subsequently lost) making up its holotype were nondiagnostic. While they acknowledged the distinctiveness of the forelimb and shoulder material, they argued it was too poorly known to make any definite taxonomic statement on it, and that there was insufficient evidence to assume only one was present in the Upper Saurian Niveau. Due to the Svalbard specimens not matching those from Monte San Giorgio in age, Maxwell and Kear argued against the assignment of the former to species from the latter location. In 2024, Bindellini and colleagues determined that the similar femur anatomy of PIMUZ T 4376 and 753:

complete, and the preservation of the bones is good. This specimen was recovered from layer 71 in the Valle Stelle mine. The other specimen, hailing from layer 116 of the Cava Tre Fontane mine, is cataloged as PIMUZ T 4847 and is very large, and poorly preserved and disarticulated, missing the tail and limbs. Sander and Jean Michel Mazin considered both of these specimens to represent a distinct genera in 1993. The medium-sized specimen was studied by David Cook, who had an abstract published in 1994, in which this specimen was interpreted as pertaining to a new genus. Dal Sasso and Pinna considered this specimen similar to

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merriamosaurs. Historically, this orientation was thought to have evolved in ichthyosaurs to reduce the likelihood of the newborns drowning during birth; however, after surveying different vivaporous amniotes, Miedema and colleagues argued that the evidence for such an adaptation was lacking, and instead proposed that birth orientation was related to which way was easier to push the fetus through the birth canal or which way made affected the mobility of the pregnant adult less. However, embryos are not the only interpretation of these remains. Jiang and colleagues in 2020 were doubtful about the inability of

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4 metres (11 by 13 ft). The skeleton is flattened from top to bottom, with the exception of the skull, which was instead crushed flat from side to side to a thickness of under 1 centimetre (0.39 in). Most disarticulation of the skeleton is fairly minimal, though the bones of the fingers are scattered. The slabs contain an additional small mixosaurid as well as the large ichthyosaur.

3301: 6954: 6432: 3317: 1289: 6444: 952:, PIMUZ T 4376, PIMUZ T 4847, PIMUZ T 1895, and BES SC 1016, most of which had not previously had their skull anatomy described in detail. BES SC 1016 had not been previously studied at all, and is an incomplete, somewhat flattened, partially articulated skull preserved in dolomite, which was analyzed with 1058:

from other shastasaurids, and that it was too fragmentary to establish a species on. Additionally, they questioned Wiman's assumption that only one large ichthyosaur was present in the Upper Saurian Niveau, noting that multiple species of large ichthyosaurs were known to coexist elsewhere. Therefore,

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rather than its own genus, though he also noted that it may instead belong to the same species as the unnamed medium-sized shastasaurid. However, Sander argued that the better-preserved material would need to be studied before definite assignments could be made. Later the same year, Maisch and Matzke

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covered an area of 35 by 45 centimetres (14 by 18 in), and 145 were needed to show the entire specimen, which was ultimately revealed to be spread across a total of 33 slabs. While the slabs are only about 2.7 centimetres (1.1 in) thick, when put together, they cover an area of about 3.5 by

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and some other shastasaurids in the analysis of Ji and colleagues, preferring this study over Moon's larger one as it was constructed with more direct observation of the specimens. They found multiple different configurations of shastasaurids, with the group either being a grade or a clade, with the

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would have been fairly straight. However, Bindellini and colleagues reported a tailbend in PIMUZ T 1895. The articular faces of the centra of the tail vertebrae vary in shape; the front ones are roughly triangular, the middle are hexagonal, and the rear are elliptical. Beneath the tail vertebrae are

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bear even stronger ridging. The teeth are separated from each other by a rather wide amount of space and interlock when the jaw is closed. The frontmost teeth in the upper jaw are the longest, and somewhat curved, while the back teeth are shorter and thicker, with a similar pattern of size and shape

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in the rear part of the palate was interpretted as wide by Maisch and Matzke in their description and reconstruction of GPIT 1793/1; however, Bindellini and colleagues in their redescription argued that this was not necessarily the case due to the specimen's disarticulation, and noted that the width

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As ichthyosaurs grow, their skulls typically become proportionately smaller. In PIMUZ T 4376, the skull is roughly half the length of the trunk; whereas it is only about a third as long as the trunk in the holotype. This difference was considered too extreme to be the result of ontogeney by Maisch

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also runs along these two pairs of bones. The frontals are long, broad, and flat bones involved in the supratemporal terraces, though they do not form any part of the borders of the supratemporal fenestrae. The parietals bear a variety of prominent surfaces for jaw muscle attachment, including the

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from the layer below broke loose. Preserved within this piece of shale from bituminous level n. 65 were the parts of the jaws of a large ichthyosaur. The rest of the skeleton of this ichthyosaur was extracted through the following summer by collecting the entire rock layer it was preserved in. The

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are about 15% longer than its hindlimbs. Each flipper bears a total of four digits, three of them well-developed and a fourth being an accessory digit (a digit not originating from the wrist or ankle). The humeri are very short, being wider than they are long, and have a rounded profile. Both the

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make up the species name. The authors stated that a more detailed study could be produced once preparation was finished, a task they predicted could take as long as 8000 more hours. It ultimately took 16500 hours spread over 5 years in total to fully reveal the skeleton, which was concealed under

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A third major analysis of ichthyosaur relationships was done by Maisch and Matzke later in 2000, featuring many similarities to Motani's analysis but based on a larger data set. Like Motani, Maisch and Matzke found the traditional shastasaurids to not be a natural group, however, they also found

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would have been able to take on mixosaurid-sized prey. In a 2021 abstract, Feiko Miedema and colleagues regarded the embryonic remains as pertaining to one embryo. In 2023, Miedema and colleagues interpretted the embryo as being oriented so that it would have been born tail first, as typical in

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as the earliest diverging member. Another series of very large analyses were run by Benjamin Moon in 2017, created by revising and compiling previous analyses. The analyses, however, were unable to clearly elucidate the relationships of many early merriamosaurs, making it unclear whether or not

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was named for the group including the shastasaurids and later ichthyosaurs. Motani noted that some parts of the phylogeny were unstable, and that many traditional shastasaurids were "problematic" due to their incompleteness. Another large phylogenetic study conducted by Sander was published the

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in the 1920s, both of them being mentioned in passing in the literature of the century. The smaller of the two, numbered PIMUZ T 4376, is a skeleton with a somewhat articulated skull and trunk but disarticulated limbs and tail. While the tail is missing its end the specimen is otherwise nearly

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would not have been able to support a powerful bite, Bindellini and colleagues noted that it could instead have allowed the jaws to rapidly close. The length of the jaws would have caused their tips to move quite quickly during biting. Additionally, the long, thin snout could have been swung

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is formed by its very long tail. The holotype specimen measures 5.065 meters (16.62 ft) long from its snout tip to tail tip. The specimen PIMUZ T 4847 was estimated by Bindellini and colleagues in 2021 to have a total length of 8 meters (26 ft), and is the largest known specimen of

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in overall body shape, they found it to compare more favorably to the shastasaurines in the fine points of its anatomy, and thus tentatively assigned it to Shastasaurinae. The phylogenetic analysis of Dal Sasso and Pinna was one of the earlier of such studies on ichthyosaurs, and subsequent

3272:. Bindellini and colleagues, however, did not find ontogeny to be an unreasonable explanation for this proportional difference in their 2021 study, instead finding the six specimens they studied to fit into a growth series when ordered by size. Unlike the skull length-body length ratio of 789:

by Fritz Lörcher. While the individual bones are generally fairly well preserved, the skull overall is crushed and strongly disarticulated. Determining it distinct after exhaustive comparisons with other ichthyosaurs, Maisch and Matzke named it the holotype of a new genus and species,

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being basal to the two. This analysis was criticized by subsequent work for only using a small sample of ichthyosaurs and encoding proportional characters based on arbitrary divisions. Concerns were also raised about the reproducibility of his results and the extremely high amount of

909:, McGowan and Motani suggested that due to its wide historical recognition the name could be revived for some distinctive referred material, noting its similarity to that of the Swiss medium-sized shastasaurid. However, they argued that the medium-sized skeleton, as well as similar 605:

and Giovanni Pinna was published later that year, the authors considering it warranted by the skeleton's distinctiveness and completeness, with the unprepared portions of the skeleton studied through radiography. Dal Sasso and Pinna found the specimen to differ from other

1391:. Extending forwards from each supratemporal fenestra is a depression, forming roughly elliptical terraces in front of the fenestrae. The region behind the orbits is short, occupying 13% of the skull's length, though it is still long compared to some parvipelvians. 3259:

to have consumed mixosaurids, and argued that it could not be ruled out that the supposed embryos may be stomach contents instead, noting that the vertebrae were the right size for an immature mixosaurid, and were more ossified than would be expected in an embryo.

1115:, he took a different stance in his 2003 book with McGowan, noting that the morphology of the humerus was unique, as was that of the bones associated with it. Like Maisch and Matzke, he recognized its similarity with the specimen in Zürich, but instead argued that 1248:, specifically various limb bones in the case of the former, considering the differences observed to be growth-related. Maisch, however, in 2010, argued against this synonymy, noting that his previous publications with Matzke had found various differences between 3179:

in its stomach region, Da-Yong Jiang and colleagues speculated in 2020 that other large ichthyosaurs adapted for consuming cephalopods may have been able to take on large prey. The researchers noted that despite the rather delicate construction of the skull of

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are also constricted at their middles and narrower than the radii and ulnae. The tibiae are the longer of the two pairs of lower leg bones, and the fibulae have heavily expanded lower ends. Compared with the corresponding elements of the forelimb, the tarsals,

1093:, was diagnostic. They found it to be very similar to PIMUZ T 4376, the medium-sized shastasaurid from Monte San Giorgio. They observed some differences between the two, but argued that they could be growth-related. As they considered PMIUZ T 4376 to belong to 1989:(natural group). The results of another comprehensive analysis by Cheng Ji and colleagues were published in 2016, and was based on a much larger data set than previous studies. A large, well-supported clade of shastasaurids was recovered by their study, with 580:

The large ichthyosaur skeleton was deposited in the Milan Natural History Museum, where it was given the specimen number BES SC 999. To determine the extent of the skeleton, the museum collaborated with a hospital in the city to scan the slabs using

1824:, Dal Sasso and Pinna coded it into a data matrix created that was created by Jack Callaway in 1989. They found shastasaurids to be divisible into two main groups, termed Cymbospondylinae and Shastasaurinae by Callaway, with the former being a 1537:

form the top portion of the rear part of the lower jaw. The upper margin of each surangular bears two eminences, a taller, pointed one immediately in front of the jaw joint and another, lower one further forwards. Based on comparisons with

3437:. 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. 693:

21,000 square centimetres (3,300 sq in) of rock. The holotype subsequently entered storage as a total of 25 slabs, while a cast of the entire specimen put on display in 1999. BES SC 999 remains the most complete specimen of

1398:. The front ends of these bones are blunt, while their rear ends are forked, with processes extending both above and below the external nares. The back edges of the external nares are formed by the upwards-directed processes of the 3055:-like) locomotion, swimming by undulating their entire body from side to side, owing to their long tails without strong bends and their long, flexible trunks. This differs from the more tail-driven locomotion presumed for the more 1931:

to be more closely related to the post-Triassic ichthyosaurs. However, the phylogeny of Maisch and Matzke (2000) has subsequently been criticized for using a hypothetical ancestor, rather than actual non-ichthyosaurian taxa, as an

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with roughly round cross-sections. The roughly rectangular neural spines of the trunk vertebrae are also tall, and are more than half as wide from side to side as they are from front to back. The vertebral bodies, known as

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was subsequently recognized by many authors, but its validity was questioned by Sander and Faber in 1998. Specifically, these authors noted that there were not enough differences preserved in the material to distinguish

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parvipelvians. While not used in propulsion, ichthyosaur limbs would have been used to steer and keep the body upright. The large eyes of ichthyosaurs indicate that vision would have been an important sense for them.

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The first comprehensive, in-depth study of overall ichthyosaur relationships was done by Motani in 1999. The traditional concept of Shastasauridae was not recovered by his analysis, with some "shastasaurids" such as

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are thin, strip-like bones bent at right angles. The articulations with other bones holding it in place were rather weak, and resultingly the jugals were often disarticulated from the skull during fossilization. The

1451:, which also reach upwards and enter the supratemporal fenestrae. On the lower regions of the sides of the skull, wide portions of the postorbitals are visible, which Bindellini and colleagues considered potentially 1406:, which run along the top of the snout, do participate in the upper back corners of the external nares. Along the midline of the skull, between the front ends of the temporal fenestrae, is a small opening called the 1607:, of the trunk vertebrae are at most half as long as tall. The front and back faces of all the centra are concave, being bowed inwards to a very thin layer of bone if not an opening. The articular surfaces for the 3230:, with the head being able to have broad movement while the body stayed in place. The authors considered the animal's large size to have potentially resulted from its foraging method being particularly efficient. 868:, PIMUZ T 1895. This specimen, coming from the Cava Tre Fontane mine, consists of a skull associated with postcranial remains, with little limb and tail material. Preparation of this specimen remains incomplete. 1579:, with the exception of the rear 30% of the maxillary teeth, which are implanted in a groove. Conversely, in the lower jaw, the front teeth are located in a groove while the back teeth are set into sockets. 1446:

make up the top of the orbital rim, where they are thickened. The prefrontals also extend forwards, where they contact the ascending processes of the maxillae. The back edges of the orbits are formed by the

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had any wedge-shaped caudal centra due to how the tail was preserved. Such vertebrae would form a downward kink known as a tailbend. However, Dal Sasso and Pinna compared the vertebral shape with that of

785:, cataloged as GPIT 1793/1. This specimen comes from an unknown position in the Besano Formation in Switzerland. This skull is preserved on three shale slabs, and detailed preparation work was done using 3202:

efficiently through the water, allowing quick up and down or side to side movements. Altogether, this configuration is ideal for feeding on prey items that are small and agile. Long, thin snouts have

3161:, noting that the slender teeth in front would have been suited for piercing small prey while the slightly blunter ones further back suited for catching cephalopods by grasping. Direct evidence of 1820:

has generally been considered a shastasaurid, how it is related to other shastasaurids is unclear due to the poorly understood phylogeny of the group. To determine the phylogenetic affinities of

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of the central European Germanic Basin: Its evolution, paleobiogeography and paleoecology' by C.G. Diedrich (Historical Biology, iFirst article, 2011, 1 – 19, doi:10.1080/08912963.2011.575938)"

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specimens all were likely recovered from the middle portion of the Besano Formation, though the exact localities of some specimens are unknown. The middle Besano Formation is also known as the

4376:(Reptilia: Ichthyosauria) from the Middle Triassic Besano Formation of Monte San Giorgio (Italy/Switzerland), with implications for reconstructing the swimming styles of Triassic ichthyosaurs" 1615:(those attached to the neck vertebrae) are double-headed, unlike the other, single-headed cervical ribs behind them. The dorsal (trunk) ribs bear grooves along their shafts. The heavily built 483:

populated by a wide variety of marine life, including a variety of other ichthyosaurs. These different ichthyosaurs are thought to have used different feeding strategies to avoid competition.

1206:, was named based on various specimens by Wiman in 1910. However, the material Wiman named this species based on is chimaeric, comprising both ichthyopterygian remains and jaw material of 405:

is a large ichthyosaur, with the largest known specimen estimated to measure about 8 metres (26 ft) long. It has a long, slender body with a small head and long tail. The snout of

1489:, and therefore suggested that it may be a fairly widespread feature of shastasaurids. The vertebral column is articulated with a convex eminence on the back of the skull known as the 841:

would have lived in the same environment to be ecologically implausible given how many other ichthyosaurs had been reported from Monte San Giorgio. He also considered it possible that

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Dal Sasso & Pinna, 1996 (Reptilia: Ichthyosauria) from the Middle Triassic Besano Formation of Monte San Giorgio, Italy/Switzerland: taxonomic and palaeobiological implications"

4899: 893:, and also considered the high diversity of ichthyosaurs reported from Monte San Giorgio suspicious. They considered both genera potentially synonymous with the much earlier-named 1622:

Most of the neural spines of the tail vertebrae slope backwards, except near its end, where they lean in the opposite direction. Dal Sasso and Pinna were not able to determine if

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following year, focusing on the more completely known ichthyosaurs. Unlike Motani, Sander found the three shastasaurids he included in his analysis to form a natural group, with

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is occupied by its very long, thin snout lined with small teeth. This distinctive snout is strongly demarcated from the rear part of the skull by a constriction. Well-preserved

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is proportionately quite small, making up less than 10% of the animal's total length in the holotype. In addition to having an elongate trunk, about half of the total length of

4955: 1744:(front lower hip bones) are vaguely circular in shape. They each bear a well-developed but narrow notch extending to the edge of the bone, rather than an enclosed foramen. The 1402:, gracile, triradiate bones behind the premaxillae. While initially interpreted as not forming part of the narial border in the holotype, further specimens revealed that the 3558:

may have been suited for crushing prey items with external shells. The abundance of ichthyosaurs in the middle Besano Formation correlates with when the lagoon was deepest.

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front and back edges of the humeri are bowed inwards, though this emargination is more notch-like on the front edges. The lower arm bones as wide as they are long, with the

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Miedema, F.; Bindellini, G.; Dal Sasso, C.; Scheyer, T.M.; Maxwell, E.E. (2021). "Conserved cranial development and early ontogeny in Triassic and Jurassic ichthyosaurs".

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is the oldest definite shastasaurid genus. Noting that other Middle Triassic ichthyosaur genera were very widespread, Bindellini and colleagues considered it likely that

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are longer than wide, though only by a by a factor of 1.22 and are resultingly still quite stout. The femora are narrowest midshaft, broadening at their upper ends. The

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Dal Sasso and Pinna discovered three or four small series of vertebrae within the holotype's chest through radiography. They interpretted the remains as pertaining to

577:

to clear out the dolomite above it, then divided into slabs and extracted with less forceful tools. Being rather fragile, the shale often broke during the excavation.

5435:"The Kalkschieferzone (Upper Meride Limestone, Ladinian) near Meride (Canton Ticino, Southern Switzerland) and the evolution of a Middle Triassic intraplatform basin" 7877: 7797: 4850:

Ji, C.; Jiang, D. Y.; Motani, R.; Rieppel, O.; Hao, W. C.; Sun, Z. Y. (2016). "Phylogeny of the Ichthyopterygia incorporating recent discoveries from South China".

1594:, located in front of the shoulders. Behind the presacral vertebrae, there are 2 vertebrae in the hip region and 139 in the tail. The neck vertebrae have prominent 1475:

in the cranium. Each quadrate has a triangular eminence on the lower part of its inner margin. In 1997, Maisch and Matzke considered this to be a unique feature of

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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

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were found to be more closely related to the later, post-Triassic ichthyosaurs. Some shastasaurids, however, were found to form a natural group in his analysis,

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of the interpterygoid vacuity was of typical width for a shastasaurid in BES SC 1016. The pterygoids possess long processes for articulation with the quadrates.

1375:, circles of bony, quadrangular plates that supported the eyeballs. Bindellini and colleagues estimated a total of between 15 and 17 of these plates per ring in 1841:

publications have criticized it for a variety of reasons. Key among these issues is that their analysis did not test their assumption that Shastasauridae was a

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specimen. This skeleton is preserved across multiple thin rock slabs spanning 3.5 by 4 metres (11 by 13 ft) when assembled and took thousands of hours to

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is one of a variety of ichthyosaurs from the Besano Formation. These different species would have had different feeding strategies to avoid competition.

549:. The excavation at Sasso Caldo is the longest-lasting excavation that was conducted at Monte San Giorgio, and was part of an operation organized by the 3532:

is known to have fed on coleoids, but its strong snout indicates that it could have taken larger prey with powerful bites, meaning it may have been an

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that attained a large body size. Bindellini listed many potential advantages of large size, including it allowing various feeding techniques, being an

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did not appear until the upper portion of the formation, when ichthyosaurs were much less common. Nothosaurids from the Besano Formation consist of

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show the remains of small coleoids and fish, suggesting that it would have gone after smaller prey than its larger relatives. The rarer mixosaurids

1238:, and that the ichthyopterygian remains referred to it also represented multiple species. They noted that some elements were comparable to those of 1130:

In their 2013 reassessment of Triassic ichthyosaurs of Svalbard, Erin Maxwell and Benjamin Kear argued in favor of Sander and Faber's assessment of

533:. This site is located about 800 metres (2,600 ft) above sea level, and was fairly easy to reach. The rocks of this quarry were unmodified by 3197:

The large coronoid processes and rough, concave outer faces of the surangulars would have anchored strong jaw musculature. While the thin jaws of

1085:

Maisch and Matzke in 2000 agreed with the taxonomic position of Sander and Faber, but argued that PMU 24584 (formerly cataloged as PMU R176), a "

5667:"Mixosaurier (Reptilia, Ichthyosaurier) mit Quetschzähnen aus der Grenzbitumenzone (Mitteltrias) des Monte San Giorgio (Schweiz, Kanton Tessin)" 1672:

are wide, with broadly expanded upper portions. Their front lower edges are short and mildly bowed inwards. Another pair of shoulder bones, the

4673: 7931: 5086: 3718: 2007:

was found to be the earliest diverging member, while within the clade it was either in a similar position or in a smaller nested group with

5477:"Microfacies, geochemistry and palaeoecology of the Middle Triassic Grenzbitumenzone from Monte San Giorgio (Canton Ticino, Switzerland)" 4970: 1575:

also present in the lower jaw. How the teeth are implanted varies through the tooth row; throughout most of the upper jaw, the teeth are

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was the result of the femora being incomplete. They also noted that the elements preserved in PMU 24584 showed differences from those of

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with a study of the postcranial anatomy published in 2024. Another study published earlier that year, led by Christian Klug, considered

1981:

In 2011, Philippa Thorne and colleagues ran an analysis based on an updated version of Motani's 1999 data matrix. Their analysis found

1479:, as at the time it was only known in the holotype skull of that taxon; however, Bindellini and colleagues identified this on multiple 1119:

should be reinstated as the name for the material due to its historical significance, in which case it could be the senior synonym of

1000:

Historically, a fairly rare species of large ichthyosaur was reported from the units informally termed the "Upper Saurian Niveau" of

7946: 6947: 4152: 3429:

are known from the Besano Formation; predominantly those that could have lived as plankton or on algae. Cephalopods present include

1761:, and phalanges of the hindflippers are less robust. Furthermore, while round, the phalanges of the pes are oblong and constricted. 1016:. The first remains of this taxon were a total of 11 vertebrae with some associated rib fragments found during the 1860s, and named 3250:, noting that a placement so far forwards in the body was not unheard of among ichthyosaurs. They also considered it doubtful that 3194:

would have been a predator of large animals, citing its long, slender snout, arguing that a diet of small animals was more likely.

925:

were distinct and valid in 2010, stating that there were was no morphological or phylogenetic support for their synonymy, and kept

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come from the Besano Formation (alternatively called the Grenzbitumenzone), a unit composed of oil shale, laminated dolomite, and

5969: 3670: 1545: 764: 414: 3368:

In the Triassic, when the Besano Formation was being deposited, the region where Monte San Giorgio is would have been a marine

120: 5373: 7684: 1468:

are large and quadrangular, as is typical for ichthyosaurs with longer cheek regions, and seen in many other shastasaurids.

1832:

was placed in an intermediate position between these two subfamilies by the analysis. While Dal Sasso and Pinna noted that

3076:. A 2020 survey of ichthyosaurs from Monte San Giorgio by Judith Pardo-Pérez and colleagues found that shastasaurids like 968:

as well, and were not able to find any distinguishing characteristics between the two genera. Therefore, they synonymized

3628:. Besides ichthyosaurs and sauropterygians, marine reptiles in the Besano Formation are represented by the thalattosaurs 3035:

were very well-adapted to their marine existence, spending their entire lives in the water and showing reduced levels of

818:

or include it in their comparisons, likely due to them not being aware of its relatively recent publication at the time.

3376:. This lagoon is estimated to be 30–130 metres (98–427 ft) deep. The upper waters of the lagoon contained abundant 1915:

Motani's concept of Shastasauridae to be a grade leading up to the post-Triassic ichthyosaurs and their relatives, with

550: 355:. Additional specimens from Monte San Giorgio that have previously been considered separate genera, including a partial 1740:, are rather wide, with their upper ends enlarged and directed inwards and their lower ends more heavily expanded. The 1127:, Motani argued that Wiman was probably right that only one large ichthyosaur was present in the Upper Saurian Niveau. 781:

In 1997, Michael Maisch and Andreas Matzke described an ichthyosaur skull housed at the Palaeontological Collection of

142: 6958: 5962: 976:, and referred the other four specimens to the latter species as well, since they also showed identifying features of 5844:

from the Alpine Triassic (Middle Triassic) and a Thalattosauroidea indet. from the Carnian of Oregon (Late Triassic)"

3039:

of their bones. Nevertheless, ichthyosaurs still would have breathed air. Ichthyosaurs were active animals with high

1379:. While large relative to the orbits, the rings are smaller relative to the animal's overall length than typical for 448:

indicate that it would have possessed strong jaw muscles, but its delicate snout suggests it would have fed on small

5265:"Heads or tails first? Evolution of fetal orientation in ichthyosaurs, with a scrutiny of the prevailing hypothesis" 4582: 4436:

Klug, C.; Sivgin, T.; Miedema, F.; Scheffold, B.; Reisdorf, A.G.; Stössel, I.; Maxwell, E.E.; Scheyer, T.M. (2024).

7721: 4557:"Memorandum on some fossil vertebrate remains collected by the Swedish expeditions to Spitzbergen in 1864 and 1868" 4273:

n. gen., n. sp., a new ichthyosaur from the Grenzbitumenzone (Anisian-Ladinian) of Monte San Giorgio (Switzerland)"

2022:

The following cladograms depict the three hypotheses recovered by the analysis of Bindellini and colleagues, 2021.

802:(Greek for "head"), referring to the way the skull was preserved, the authors likening its appearance to a game of 6964: 5901:

Spiekman, S. N. F.; Neenan, J. M.; Fraser, N. C.; Fernandez, V.; Rieppel, O.; Nosotti, S.; Scheyer, T. M. (2020).

5139:

Gutarra, S.; Moon, B. C.; Rahman, I. A.; Palmer, C.; Lautenschlager, S.; Brimacombe, A. J.; Benton, M. J. (2019).

3276:, the researchers found the orbit size and mandible length to increase at the same rate, and the tooth anatomy of 2019:. Due to this instability, Bindellini and colleagues considered shastasaurid relationships to still be ambiguous. 7936: 2719: 2433: 2099: 2015: 1189:

or a similar ichthyosaur, but agreed that it could not be distinguished from other representatives of the group.

5736:

Scheyer, T.M.; Neenan, J.M.; Renesto, S.; Saller, F.; Hagdorn, H.; Furrer, H.; Rieppel, O.; Tintori, A. (2012).

4510:"Triassic ichthyopterygian assemblages of the Svalbard archipelago: A reassessment of taxonomy and distribution" 3264:

and Matzke in 2000, arguing the specimens were too similar in size. Thus, they used this feature to distinguish

3080:

were more likely to show skeletal injuries than the smaller mixosaurids, but showed less frequent injuries than

1923:

was also coded into their analysis, and found to be more closely related to the post-Triassic ichthyosaurs than

1185:

since the late 20th century, Maisch and Matzke noted in 2000 the possibility that the fossil material came from

7716: 7672: 3171:-like hooklet preserved as stomach contents in the holotype. However, following the discovery of a specimen of 3157:, based on the shape and small size of its teeth. Bindellini and colleagues elaborated on the tooth anatomy of 3069: 767:. However, Cook's study of PIMUZ T 4376 did not lead to a paper, and the specimen remained incompletely known. 4674:"Evolution of fish-shaped reptiles (Reptilia: Ichthyopterygia) in their physical environments and constraints" 4196:, sp. nov., from the Middle Triassic of Monte San Giorgio (Switzerland), with a survey of the genus in Europe" 3469:

are also known in the formation, though the latter may have been washed in from elsewhere, as with many other

1203: 944:

was published. The authors studied six shastasaurid specimens from Monte San Giorgio, namely the holotypes of

5263:

Miedema, F.; Klein, N.; Blackburn, D.G.; Sander, P.M.; Maxwell, E.E.; Griebeler, E.M.; Scheyer, T.M. (2023).

5141:"Effects of body plan evolution on the hydrodynamic drag and energy requirements of swimming in ichthyosaurs" 3400:

is minimal and random, which also indicates undisturbed waters. The presence of terrestrial fossils, such as

1210:, a group of crushing-toothed marine reptiles. This resulted in temporary disagreement over whether the name 1384: 1260:

was a valid genus. Maxwell and Kear in 2013 doubted the assignment of some of the material considered to be

4171:

Cook, D.H.N. (1994). "A new ichthyosaur genus from the Middle Triassic of Monte San Giorgio, Switzerland".

4142: 7742: 6437: 6431: 3602: 3238: 782: 557:

group. In the spring of 1993, while extracting a block of dolomite containing the skeletons of a pair of

7903: 7823: 7711: 5191:

Jiang, D.Y.; Motani, R.; Tintori, A.; Rieppel, O.; Ji, C.; Zhou, M.; Wang, X.; Lu, H.; Li, Z.G. (2020).

4949: 4640:

n. g. (Ichthyosauria, Lower Triassic) and their bearing on the systematic position of the Omphalosauria"

4336: 3608: 3222:

may have consumed fish as well as coleoids. Bindellini and colleagues also noted that the large size of

3120: 1933: 598: 3797:

n. gen. n. sp., a new shastasaurid ichthyosaur from the Middle Triassic of Besano (Lombardy, N. Italy)"

3505: 3019: 1785: 1956: 1713:

being the larger of the two pairs. The roughly quadrangular radii have constricted middles, while the

1347: 6785: 6707: 6655: 5855: 5838:

Klein, N.; Sander, P.M.; Liu, J.; Druckenmiller, P.; Metz, E.T.; Kelley, N.P.; Scheyer, T.M. (2023).

5792: 5781:"A new pachypleurosaur from the Early Ladinian Prosanto Formation in the Eastern Alps of Switzerland" 5631: 5572: 5513: 5382: 5204: 5031: 4985: 4926: 4859: 4739: 4688: 4521: 4449: 4387: 4284: 4207: 4042: 3978: 3665: 3203: 2743: 2409: 2075: 2009: 1178: 749: 712: 278: 234: 6443: 4704: 4094: 1927:. They had previously found a similar result in a smaller study they conducted in 1997, which found 1230:

is indeed the correct name for the ichthyosaurian material. In 2003, McGowan and Motani argued that

913:

ichthyosaurs, would need to be described in greater detail before they could assess the validity of

409:

is long and thin, and contains numerous small pointed teeth. In the upper jaw, the teeth are mostly

363:

and a well-preserved, largely complete skeleton, have been reinterpreted as additional specimens of

7071: 6914: 6061: 4609: 4556: 1576: 1426: 1256:, and recovered the two of them as not being particularly close relatives. He also maintained that 1194: 1154: 889: 602: 410: 344: 230: 6928: 1070: 7184: 6603: 6488: 6238: 6123: 6110: 6073: 5985: 5761: 5718: 5647: 5563: 5539: 5408: 5344: 4930: 4875: 4537: 4300: 4058: 3994: 3495: 3338: 1591: 594: 352: 263: 137: 7908: 7828: 3708: 1383:

ichthyosaurs. Further back, the top of the skull is perforated by another pair of openings, the

770: 601:. While the preparation was not yet complete, a preliminary report on this large ichthyosaur by 6953: 3361:

too was a wide-ranging genus, despite all definite specimens being known from this unit in the

933:

separate as well. Otherwise, however, little further research was published on the taxonomy of

7890: 7882: 7810: 7802: 7502: 7377: 7342: 7298: 6823: 6203: 5940: 5883: 5820: 5600: 5559:"Early Mesozoic burst of morphological disparity in the slow-evolving coelacanth fish lineage" 5531: 5434: 5400: 5296: 5230: 5170: 5082: 5059: 5001: 4824:"First record of ichthyosaurs in Sicily (Upper Triassic of Monte Scalpello, Catania Province)" 4765: 4477: 4415: 4148: 3934: 3714: 3190:

would have been able to fit inside its mouth. However, Bindellini and colleagues doubted that

1966: 1704:, a midline bone located between them, does not appear to have been present. The forelimbs of 1656:

Front portion of holotype, showing skull as well as vertebral, shoulder, and forelimb elements

1637: 1490: 1307: 1174: 530: 394: 322: 7895: 7815: 3481:

being quite diverse, including abundant small species as well as larger representatives like

1268:

and considered it to be potentially valid. Bindellini and colleagues in 2024 also considered

1150:, indicating that it pertained to a different ichthyosaur, although similar or even related. 7622: 7571: 7539: 7422: 7227: 7083: 7028: 6582: 6519: 5930: 5920: 5873: 5863: 5810: 5800: 5753: 5708: 5639: 5590: 5580: 5521: 5390: 5334: 5286: 5276: 5220: 5212: 5160: 5152: 5119: 5049: 5039: 4993: 4922: 4914: 4867: 4804: 4755: 4747: 4696: 4651: 4529: 4467: 4457: 4405: 4395: 4292: 4215: 4050: 3986: 3924: 3914: 3073: 1944: 1825: 1774: 1652: 1604: 1448: 1443: 1139: 546: 472: 347:

and Giovanni Pinna in 1996, based on the nearly complete flattened skeleton BES SC 999, the

4700: 1611:

on the sides centra do not extend all the way forwards to their front edges. The frontmost

1367:, openings which housed the nostrils in life. Behind the external nares are the elliptical 871: 744:

was not the first shastasaur known from Monte San Giorgio. Two shastasaurid specimens from

726: 7465: 7457: 7449: 7407: 7311: 7259: 7160: 6921: 6900: 6893: 6771: 6750: 6610: 6320: 6281: 6247: 6219: 6168: 5779:

Klein, N.; Furrer, H.; Ehrbar, I.; Torres Ladeira, M.; Richter, H.; Scheyer, T.M. (2022).

5504: 4636:"Observations on Triassic ichthyosaurs. Part IX. The first associated skeletal remains of 3581: 3478: 2918: 2591: 2219: 1439: 1372: 1368: 1303: 956:. It was recovered from stratum 70 at Sasso Caldo, the same site at which the holotype of 755: 643: 542: 534: 512: 315: 41: 17: 3388:

organisms. However, water circulation within the lagoon was poor, resulting in typically

3287:

and the probable fetus present in the holotype's chest are subjects for future research.

1998:

was a shastasaur. In their 2021 study, Bindellini and colleagues updated the codings for

1729:(finger bones) are also rounded, the latter two types being having the shape of circles. 444:

and other members of this group are related to each other is unclear. The skull bones of

5878: 5859: 5839: 5796: 5635: 5595: 5576: 5558: 5517: 5475:

Röhl, H.J.; Schmid-Röhl, A.; Furrer, H.; Frimmel, A.; Oschmann, W.; Schwark, L. (2001).

5386: 5369:"Middle Triassic gastropods from the Besano Formation of Monte San Giorgio, Switzerland" 5291: 5264: 5208: 5035: 4989: 4863: 4743: 4692: 4525: 4472: 4453: 4437: 4410: 4391: 4371: 4288: 4211: 4046: 3982: 833:

in 2000, noting that they were anatomical similar. Additionally, he found the fact that

496: 7615: 7608: 7580: 7349: 7287: 7280: 7243: 7236: 7215: 7110: 7056: 6813: 6793: 6763: 6715: 6699: 6682: 6635: 6543: 6535: 6528: 6294: 5935: 5902: 5815: 5780: 5697:

from the latest Anisian (Middle Triassic) Besano formation (Grenzbitumenzone) of Italy"

5225: 5192: 5165: 5140: 5054: 5019: 4760: 4727: 3929: 3898: 3596: 3590: 3562: 3488: 3446: 3167: 3093: 3040: 2941: 2802: 2691: 2614: 2457: 2364: 2242: 2123: 2037: 1886: 1796: 1558: 1549: 1518: 1498: 1485: 1461: 1419: 1407: 1364: 1244: 1199: 1033: 826: 803: 651: 210: 5738:"Revised paleoecology of placodonts – with a comment on 'The shallow marine placodont 4808: 4370:

Bindellini, G.; Wolniewicz, A. S.; Miedema, F.; Dal Sasso, C.; Scheyer, T. M. (2024).

1351:

Skull reconstruction in oblique (A), rear (B), top (C), side (D), and bottom (E) views

940:

In 2021, a paper by Gabriele Bindellini and colleagues detailing the skull anatomy of

7925: 7601: 7479: 7386: 7365: 7169: 7102: 7092: 7049: 7009: 6551: 6503: 6474: 6461: 6391: 6263: 6256: 6179: 6161: 5737: 5666: 5651: 5643: 5543: 5476: 5412: 5348: 4784: 4635: 4541: 4509: 4304: 4268: 4234: 4191: 4030: 3998: 3962: 3792: 3652: 3630: 3533: 3138: 3056: 1919:

as the first of these taxa to branch off. As it was considered separate at the time,

1710: 1701: 1680:

heads, with their upper margins heavily expanded and their lower ends fan-shaped. In

1612: 1599: 1501:, which Bindellini and colleagues reconstructed as rounded and roughly pentagonal in 1472: 1452: 1435: 1415: 1226:

was considered to be the omphalosaurid. However, consensus subsequently emerged that

1207: 1018: 1013: 847:, another fragmentary ichthyosaur named in 1998 by Maisch and Matzke, was a juvenile 795: 597:

had been performed by three four preparators on the specimen, exposing the skull and

197: 78: 5765: 5722: 4934: 4879: 4655: 4612:[Contributions to the knowledge of ichthyosaurs in the German Muschelkalk]. 4062: 3897:

Bindellini, G.; Wolniewicz, A.S.; Miedema, F.; Scheyer, T.M.; Dal Sasso, C. (2021).

1845:, as their matrix did not include any non-shastasaurid ichthyopterygians other than 7560: 7553: 7532: 7517: 7472: 7442: 7327: 7273: 7146: 6883: 6831: 6731: 6723: 6594: 6568: 6349: 6342: 6302: 6270: 6212: 4997: 4219: 3570: 3413: 3393: 3389: 3373: 3176: 3048: 3036: 2826: 2481: 2147: 1895: 1852: 1806: 1737: 1595: 1567: 1540: 1423:

sagittal crest and their contributions to the rims of the supratemporal fenestrae.

1411: 1170: 905: 786: 703: 570: 437: 242: 4918: 4871: 1544:, Bindellini and colleagues interpretted the taller of the two projections as the 697:. While P. Martin Sander and Christiane Faber in 1998 considered it possible that 5757: 4533: 1911:(convergent evolution of features by unrelated groups) implied by his cladogram. 1434:

The front portions of the orbits are formed by a pair of crescentic bones called

964:, finding many of the features initially used to distinguish it to be present in 875:

Referred specimen PIMUZ T 4847 on display at the Paleontological Museum of Zurich

7435: 7400: 7319: 7125: 7118: 6995: 6982: 6907: 6842: 6801: 6738: 6662: 6192: 6148: 6137: 6030: 5840:"Comparative bone histology of two thalattosaurians (Diapsida: Thalattosauria): 4822:

Dal Sasso, C.; Insacco, G.; Chiarenza, A.A.; Di Franco, D.; Reitano, A. (2014).

4337:"Phylogeny, systematics, and origin of the Ichthyosauria – the state of the art" 3647: 3642: 3585: 3483: 3462: 3404:

and land-dwelling reptiles indicates that the region would have been near land.

3215: 3109: 2956: 2843: 2629: 2498: 2257: 2171: 1881: 1758: 1741: 1722: 1494: 1380: 1166: 1089:" specimen consisting of associated shoulder and forelimb material, including a 895: 745: 586: 561: 375: 330: 318: 53: 7855: 5907:(Tanystropheidae, Archosauromorpha) as revealed by synchrotron microtomography" 5868: 5805: 5585: 5395: 5368: 5281: 5216: 4751: 4462: 4400: 3300: 1288: 748:

were deposited in the collections of Paläontologisches Institut und Museum der

413:

but the rearmost teeth are implanted in a groove. The lower jaw bears enlarged

7629: 7587: 7494: 7487: 7393: 7334: 7201: 7042: 7016: 6778: 6624: 6617: 6369: 6360: 6309: 6046: 6015: 5713: 5692: 5526: 5499: 5193:"Evidence supporting predation of 4-m marine reptile by Triassic megapredator" 5020:"Resetting the evolution of marine reptiles at the Triassic-Jurassic boundary" 4235:"The paleobiogeography of Middle Triassic ichthyosaurs: The five major faunas" 3546: 3491: 3466: 3454: 3316: 3186: 3154: 3044: 2894: 2567: 2195: 1571: 1534: 1456: 1403: 1395: 1311: 1038: 1024: 732: 607: 574: 558: 460: 456: 398: 98: 63: 7765: 5535: 5404: 1566:

are all similar in shape, being small, cone-shaped, and acutely pointed. The

1394:

In the upper jaw, the snout is formed by the elongation of the tooth-bearing

899:, though remained tentative as the material was not studied firsthand. While 7524: 7509: 7357: 7176: 6329: 5044: 3566: 3474: 3458: 3438: 3430: 3426: 3089: 1951:

in 2010, though he considered it to be within Merriamosauria. He classified

1941: 1908: 1726: 1616: 1590:

has 60 vertebrae located in front of its hips (presacrals); 11 of these are

1530: 1526: 1514: 1214:

should be used for the ichthyosaur or omphalosaurid material, and the names

1049:, comprising all large ichthyosaur specimens from the Upper Saurian Niveau. 554: 538: 430: 418: 154: 103: 5944: 5887: 5824: 5604: 5339: 5318: 5300: 5234: 5174: 5156: 5077:

Naish, D. (2023). "Shark-shaped reptiles: The ichthyosaurs and their kin".

5063: 4769: 4481: 4419: 3938: 3594:

is known from the middle Besano Formation, while the particularly abundant

3210:

with similar feeding habits. As modern long-snouted predators such as some

2003:

former arrangement found with marginally more frequency. Within the grade,

1123:. Since PIMUZ T 4376 had a femur similar to those historically assigned to 7776: 5124: 5103: 3550:

are also known from Monte San Giorgio; both possess broad crushing teeth.

3341:

at Monte San Giorgio, and measures 5–16 metres (16–52 ft) thick. The

553:, with the excavation performed by the Gruppo Paleontologico di Besano, a 7849: 7759: 7546: 6670: 6495: 6009: 5500:"A Hiatus Obscures the Early Evolution of Modern Lineages of Bony Fishes" 3636: 3576: 3499: 3442: 3434: 3421: 3381: 3175:, a supposed cephalopod predator, with a considerable portion of a large 1863:

representing earlier branches of ichthyosaur evolution while others like

1693: 1673: 1399: 1292: 1005: 1001: 843: 647: 611: 501: 379: 348: 174: 129: 93: 88: 73: 68: 58: 5925: 3919: 1855:

group and the inclusion of highly fragmentary taxa was also criticized.

425:

are sickle-shaped, and the forelimbs are longer than the hindlimbs. The

7869: 7789: 7594: 6376: 6021: 5250:

European Association of Vertebrate Palaeontologists 18th Annual Meeting

4296: 4054: 3990: 3470: 3417: 3401: 3350: 3247: 3227: 3211: 3207: 3151: 3134: 3065: 1847: 1718: 1685: 1665: 1090: 1009: 953: 631: 619: 476: 453: 422: 383: 308: 184: 108: 83: 47: 1045:, to contain the species. Wiman assigned many additional specimens to 459:, which it could have caught with rapid, snapping bites. Ichthyosaurs 421:

in front of the hips, two in the hip region, and 139 in the tail. The

6003: 5954: 3450: 3385: 3377: 3369: 1753: 1661: 1570:

are lined with vertical ridges, though lack cutting edges, while the

1506: 635: 480: 464: 426: 372: 300: 164: 7736: 5079:

Ancient Sea Reptiles: Plesiosaurs, Ichthyosaurs, Mosasaurs, and More

4823: 4591:

Bulletin of the Geological Institutions of the University of Uppsala

3453:. Other, rarer invertebrate groups known from the formation include 3337:. This formation is one of a series of Middle Triassic units atop a 763:

in their description of the latter genus, following discussion with

4610:"Beiträge zur Kenntnis der Ichthyosaurier im deutschen Muschelkalk" 3554:

is understood to have consumed coleoids, while the larger teeth of

3498:

of the Besano Formation are uncommon as well and mainly consist of

5911: 3565:

are known from the Besano Formation, including the shell-crushing

3504: 3237: 3018: 1986: 1965: 1947:, Besanosauridae, to contain it. Maisch also used this family for 1842: 1749: 1745: 1651: 1425: 1346: 1287: 1069: 992:, though noted that more research would be needed to confim this. 910: 870: 769: 725: 615: 582: 565: 526: 495: 356: 326: 311: 5618:

Maisey, J.G. (2011). "The braincase of the Middle Triassic shark

4728:"Skeletal pathologies track body plan evolution in ichthyosaurs" 3362: 3334: 3310:

specimens have been found (yellow diamonds) at Monte San Giorgio

3283:

In 2024, Bindellini and colleagues mention that the ontogeny of

1714: 1517:, are broad-ended but slender-shafted. The openings between the 449: 7740: 7700: 6980: 6459: 6108: 5996: 5958: 5104:"Osteohistology of the Early Triassic ichthyopterygian reptile 1181:, Germany. Although this taxon has been widely recognized as a 4785:"Observations on Triassic ichthyosaurs. Part V. The skulls of 3226:

could have helped with prey capture as well by providing more

3084:, thought to have preyed upon larger animals. The holotype of 3052: 1677: 1608: 1101: 5317:

Renesto, S.; Dal Sasso, C.; Fogliazza, F.; Ragni, C. (2020).

4031:"Ichthyosauria: their diversity, distribution, and phylogeny" 1314:. While later ichthyosaurs developed fish-shaped body plans, 1027:

in 1873. The species was subsequently suggested to belong to

386:, although their attribution to this genus remains disputed. 4644:

Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen

3242:

The small vertebrae preserved within the holotype's rib cage

2025: 1871:

among them. Based on the results of his analysis, he placed

4561:

Bihang till Kongl. Svenska Vetenskaps-akademiens Handlingar

1940:

to fall outside of Merriamosauria in 2003, and named a new

1264:

by Maisch and Matzke, but agreed that it was distinct from

879:

In a 2003 book, Christopher McGowan and Motani listed both

774:

Disarticulated skull GPIT 1793/1, the holotype specimen of

5319:"New findings reveal that the Middle Triassic ichthyosaur 4797:

Neues Jahrbuch für Geologie und Paläontologie, Monatshefte

1875:

in Shastasauridae, but outside Shastasaurinae, formed by

4585:[Ichthyosaurs from the Triassic of Spitsbergen] 903:

had previously been considered nondiagnostic and thus a

500:

Skull and front part of the skeleton of BES SC 999, the

3967:

and a review of Triassic ichthyosaur paleobiogeography"

1371:, somewhat longer than tall. Within the orbits are the 980:. The same team of authors continued their revision of 5081:. Washington, DC: Smithsonian Books. pp. 94–123. 4900:"A new phylogeny of ichthyosaurs (Reptilia: Diapsida)" 1598:(bony projections of the vertebrae) and large, robust 1552:

to the surangular. The coronoid process is massive in

1471:

The jaw joint was formed by the robust, kidney-shaped

1165:, on the basis of a single vertebra discovered in the 4726:

Pardo-Pérez, J.M.; Kear, B.P.; Maxwell, E.E. (2020).

1339:

is the largest known ichthyosaur in its environment.

1097:, they tentatively assigned the Svalbard specimen to 960:

was discovered. The authors reassessed the status of

4144:

Handbook of Paleoherpetology Part 8: Ichthyopterygia

1688:, however, no such openings are present in those of 1619:(belly ribs) are united along the animal's midline. 806:, while the species name comes from the Latin words 7839: 7749: 7570: 7421: 7376: 7297: 7257: 7226: 7199: 7159: 7144: 7091: 7070: 7026: 6993: 6881: 6841: 6812: 6749: 6681: 6634: 6593: 6566: 6518: 6472: 6390: 6359: 6328: 6319: 6280: 6246: 6237: 6202: 6178: 6147: 6121: 6072: 6045: 3165:feeding on cephalopods is present in the form of a 1430:

Photograph and diagram of the skull of PIMUZ T 4376

515:work began at a site known as Sasso Caldo (meaning 4141:McGowan, C.; Motani, R. (2003). Sues, H.D. (ed.). 3713:. Bloomington, Indiana: Indiana University Press. 1717:behind them are rounded and somewhat smaller. The 463:, and the holotype may contain the remains of an 5701:Rivista Italiana di Paleontologia e Stratigrafia 4954:: CS1 maint: bot: original URL status unknown ( 4828:Rivista Italiana di Paleontologia e Stratigrafia 3513:(top) compared with that of the contemporaneous 3394:bottom-dwelling organisms modifying the sediment 3372:, located in a basin on the western side of the 1700:are thin and broadest at their middles, and the 730:Referred specimen PIMUZ T 4376, together with a 5024:Proceedings of the National Academy of Sciences 2684:Cladogram showing monophyletic Shastasauridae: 2357:Cladogram showing monophyletic Shastasauridae: 2030:Cladogram showing paraphyletic Shastasauridae: 545:pertaining to the geological unit known as the 467:in its chest cavity. All definite specimens of 417:for the anchorage of jaw muscles. There are 60 3396:. The disarticulation seen in the holotype of 3150:to likely have been specialized on feeding on 3047:. Intermediate-grade ichthyosaurs likely used 1276:, and concluded that there was no evidence of 860:. Additionally, they referred PIMUZ T 4376 to 856:, listing multiple differences between it and 814:(snout). However, the authors did not mention 5970: 5108:: Implications for early ichthyosaur biology" 5102:Nakajima, Y.; Houssaye, A.; Endo, H. (2014). 5018:Thorne, P.M.; Ruta, M.; Benton, M.J. (2011). 4681:Annual Review of Earth and Planetary Sciences 1556:, only rivaled in its development by that of 8: 5671:Schweizerische Paläontologische Abhandlungen 4099:Stuttgarter Beiträge zur Naturkunde, Serie B 1640:, Y-shaped bones which are not very long in 1483:specimens in 2021, as well as a specimen of 996:Putative specimens from Svalbard and Germany 701:represented another specimen of the related 4583:"Ichthyosaurier aus der Trias Spitzbergens" 3477:have been recorded in this formation, with 3380:, and were inhabited by a diverse array of 3218:, Bindellini and colleagues suggested that 1078:that was suggested to have affinities with 7737: 7697: 7427: 7303: 7265: 7254: 7207: 7156: 7152: 7077: 7067: 7034: 7001: 6990: 6977: 6878: 6755: 6640: 6574: 6515: 6480: 6469: 6456: 6334: 6325: 6286: 6243: 6184: 6153: 6129: 6118: 6105: 6051: 6042: 5993: 5977: 5963: 5955: 4937:. Archived from the original on 2020-03-18 1107:. While Motani had previously agreed with 132:(top) and interpretative drawing (bottom) 119: 31: 5934: 5924: 5877: 5867: 5814: 5804: 5712: 5594: 5584: 5525: 5394: 5338: 5290: 5280: 5224: 5164: 5123: 5053: 5043: 4927:1983/463e9f78-10b7-4262-9643-0454b4aa7763 4759: 4471: 4461: 4409: 4399: 3928: 3918: 1529:is formed by the long, thin toothbearing 1192:Another large ichthyosaur from Svalbard, 5470: 5468: 5466: 5464: 5462: 5460: 5458: 5456: 5454: 5452: 5323:is the oldest amniote with a dorsal fin" 5312: 5310: 5186: 5184: 4845: 4843: 4841: 4721: 4719: 4717: 4431: 4429: 4365: 4363: 4361: 4359: 4357: 3892: 3890: 3888: 3886: 3884: 3882: 3880: 3878: 3876: 3874: 3872: 3870: 3868: 3866: 3864: 3862: 3860: 3858: 3856: 3854: 3852: 3850: 3848: 3846: 3844: 3842: 3840: 3838: 3836: 3834: 5362: 5360: 5358: 4629: 4627: 4503: 4501: 4499: 4497: 4495: 4493: 4491: 4262: 4260: 4258: 4256: 4254: 4252: 4136: 4134: 4132: 3832: 3830: 3828: 3826: 3824: 3822: 3820: 3818: 3816: 3814: 3786: 3784: 3782: 3780: 3778: 3776: 3774: 3772: 3770: 3768: 3766: 3764: 3762: 3760: 3758: 3756: 3754: 3752: 3750: 3682: 5686: 5684: 5428: 5426: 5424: 5422: 4947: 4893: 4891: 4889: 4701:10.1146/annurev.earth.33.092203.122707 4667: 4665: 4576: 4574: 4330: 4328: 4326: 4324: 4322: 4320: 4318: 4316: 4314: 4147:. Munich: Verlag Dr. Friedrich Pfeil. 4130: 4128: 4126: 4124: 4122: 4120: 4118: 4116: 4114: 4112: 4088: 4086: 4084: 4082: 4080: 4078: 4076: 4074: 4072: 3956: 3954: 3952: 3950: 3948: 3748: 3746: 3744: 3742: 3740: 3738: 3736: 3734: 3732: 3730: 3433:, coleoids, and the especially common 1851:. The usage of Cymbospondylinae for a 1497:entered through an opening called the 794:. The genus name comes from the words 4166: 4164: 4093:Maisch, M. W.; Matzke, A. T. (2000). 4024: 4022: 4020: 4018: 4016: 4014: 4012: 4010: 4008: 3702: 3700: 3698: 3696: 3694: 3692: 3690: 3688: 3686: 988:to probably be a distinct genus from 7: 4508:Maxwell, E. E.; Kear, B. P. (2013). 3624:may have been an apex predator like 1074:Diagram of the specimen assigned to 1008:stage of the Middle Triassic to the 4907:Journal of Systematic Palaeontology 4783:Maisch, M.W.; Matzke, A.T. (1999). 4634:Maisch, M.W.; Matzke, A.T. (2002). 4267:Maisch, M.W.; Matzke, A.T. (1997). 4239:Paleontologia Lombarda, Nuova Serie 3064:is the earliest known long-snouted 2934: 2910: 2886: 2818: 2794: 2735: 2711: 2704: 2697: 2687: 2607: 2583: 2559: 2473: 2449: 2425: 2401: 2377: 2370: 2360: 2235: 2211: 2187: 2163: 2139: 2115: 2091: 2067: 2043: 2033: 1974:, which may be a close relative of 1548:, formed through the fusion of the 864:, and assigned another specimen to 7942:Middle Triassic reptiles of Europe 5145:Proceedings of the Royal Society B 4978:Journal of Vertebrate Paleontology 4971:"Phylogeny of the Ichthyopterygia" 4852:Journal of Vertebrate Paleontology 4233:Sander, P.M.; Mazin, J.M. (1993). 4200:Journal of Vertebrate Paleontology 4173:Journal of Vertebrate Paleontology 3072:, and resulting in more efficient 1533:. Behind the dentaries, the large 25: 3791:Dal Sasso, C.; Pinna, G. (1996). 3441:known from the formation include 3416:from the Besano Formation is the 1355:About two-thirds of the skull of 759:and proportionally distinct from 711:has otherwise been accepted as a 7683: 7677: 7671: 6963: 6957: 6952: 6946: 6442: 6436: 6430: 5644:10.1111/j.1475-4983.2011.01035.x 5557:Ferrante, C.; Cavin, L. (2023). 5367:Pieroni, V.; Furrer, H. (2020). 3961:Sander, P.M.; Faber, C. (1998). 3671:Timeline of ichthyosaur research 3329:All known definite specimens of 3315: 3299: 3119: 3108: 3045:maintain their body temperatures 1936:. McGowan and Motani considered 1784: 1773: 1684:, each coracoid is pierced by a 1410:, whose border is formed by the 1318:is more elongate, and resembles 1067:and indeterminate shastasaurid. 736:(top) preserved on the same slab 593:By February 1996, 2500 hours of 333:, containing the single species 141: 3612:, and an additional species of 3146:Dal Sasso and Pinna considered 1985:to be part of the shastasaurid 1794:While superficially resembling 1631:and suggested that the tail of 1363:skulls show drawn-out slotlike 1322:in overall shape. The skull of 440:-type ichthyosaur, how exactly 436:While it is understood to be a 5848:Swiss Journal of Palaeontology 5785:Swiss Journal of Palaeontology 5374:Swiss Journal of Palaeontology 4998:10.1080/02724634.1999.10011160 4442:Swiss Journal of Palaeontology 4438:"Swiss ichthyosaurs: a review" 4380:Swiss Journal of Palaeontology 4220:10.1080/02724634.1989.10011750 3349:Zone, and dates to the latest 1: 5327:Acta Palaeontologica Polonica 5112:Acta Palaeontologica Polonica 4919:10.1080/14772019.2017.1394922 4872:10.1080/02724634.2015.1025956 4271:Mikadocephalus gracilirostris 3461:, which lived on the seabed. 3412:Among the most common of the 3280:to not change with ontogeny. 970:Mikadocephalus gracilirostris 881:Mikadocephalus gracilirostris 862:Mikadocephalus gracilirostris 792:Mikadocephalus gracilirostris 776:Mikadocephalus gracilirostris 429:are round and squat, and the 274:Mikadocephalus gracilirostris 27:Genus of Triassic ichthyosaur 7932:Middle Triassic ichthyosaurs 5758:10.1080/08912963.2011.621083 5439:Eclogae Geologicae Helvetiae 4534:10.1080/11035897.2012.759145 4277:Paläontologische Zeitschrift 4035:Paläontologische Zeitschrift 3971:Paläontologische Zeitschrift 1387:, which are rather small in 1157:erected a second species of 551:Milan Natural History Museum 371:have been discovered in the 5903:"The cranial morphology of 4809:10.1127/njgpm/1999/1999/345 3322:A view of Monte San Giorgio 2720:"Callawayia" wolonggangense 2434:"Callawayia" wolonggangense 2100:"Callawayia" wolonggangense 2016:"Callawayia" wolonggangense 1804:is more closely related to 1222:were sometimes employed if 642:, referencing a village in 569:shale was exposed by using 537:and consisted of layers of 257:Dal Sasso & Pinna, 1996 7963: 5869:10.1186/s13358-023-00277-3 5806:10.1186/s13358-022-00254-2 5586:10.1038/s41598-023-37849-9 5505:Frontiers in Earth Science 5396:10.1186/s13358-019-00201-8 5282:10.1186/s12862-023-02110-4 5217:10.1016/j.isci.2020.101347 4752:10.1038/s41598-020-61070-7 4463:10.1186/s13358-024-00327-4 4401:10.1186/s13358-024-00330-9 4374:Besanosaurus leptorhynchus 3901:Besanosaurus leptorhynchus 3795:Besanosaurus leptorhynchus 3536:. The stomach contents of 3206:multiple times in aquatic 1828:leading up to the latter. 1202:Vendomdalen Member of the 974:Besanosaurus leptorhynchus 852:defeneded the validity of 624:Besanosaurus leptorhynchus 252:Besanosaurus leptorhynchus 18:Besanosaurus leptorhynchus 7707: 7696: 7669: 7430: 7306: 7268: 7253: 7210: 7155: 7080: 7066: 7037: 7004: 6989: 6976: 6944: 6877: 6758: 6643: 6577: 6514: 6483: 6468: 6455: 6428: 6337: 6289: 6187: 6156: 6132: 6117: 6104: 6054: 6041: 5992: 5527:10.3389/feart.2020.618853 5269:BMC Ecology and Evolution 4656:10.1127/njgpa/226/2002/59 3471:bottom-dwelling organisms 2954: 2939: 2932: 2915: 2908: 2891: 2884: 2840: 2823: 2816: 2799: 2792: 2757: 2740: 2733: 2716: 2709: 2702: 2695: 2627: 2612: 2605: 2588: 2581: 2564: 2557: 2495: 2478: 2471: 2454: 2447: 2430: 2423: 2406: 2399: 2382: 2375: 2368: 2255: 2240: 2233: 2216: 2209: 2192: 2185: 2168: 2161: 2144: 2137: 2120: 2113: 2096: 2089: 2072: 2065: 2048: 2041: 1513:, a pair of bones in the 1418:on the sides and back. A 269: 262: 248: 241: 138:Scientific classification 136: 127: 118: 34: 7947:Ichthyosauromorph genera 4372:"Postcranial anatomy of 3542:Mixosaurus kuhnschnyderi 3070:anti-predator adaptation 1836:superficially resembled 461:gave birth to live young 367:. Putative specimens of 128:Cast of BES SC 999, the 5905:Tanystropheus hydroides 5714:10.13130/2039-4942/5946 5045:10.1073/pnas.1018959108 4194:Cymbospondylus buchseri 4192:"The large ichthyosaur 3530:Cymbospondylus buchseri 3494:were also present. The 3234:Reproduction and growth 1732:The upper hip bones of 1385:supratemporal fenestrae 626:. The genus name means 5842:Askeptosaurus italicus 5665:Brinkmann, W. (2004). 5620:Acronemus tuberculatus 5340:10.4202/app.00731.2020 5321:Mixosaurus cornalianus 5157:10.1098/rspb.2018.2786 4608:von Huene, F. (1916). 4029:Sander, P. M. (2000). 3801:Paleontologia Lombarda 3707:Dal Sasso, C. (2004). 3603:Silvestrosaurus buzzii 3588:. The pachypleurosaur 3538:Mixosaurus cornalianus 3522: 3243: 3028: 1978: 1955:in a separate family, 1657: 1431: 1352: 1295: 1082: 885:Wimanius odontopalatus 876: 778: 737: 719:Further specimens and 508: 7904:Paleobiology Database 7824:Paleobiology Database 5125:10.4202/app.2012.0045 4335:Maisch, M.W. (2010). 4190:Sander, P.M. (1989). 3609:Nothosaurus giganteus 3508: 3408:Contemporaneous biota 3241: 3022: 1969: 1655: 1648:Appendicular skeleton 1429: 1350: 1291: 1204:Vikinghøgda Formation 1073: 917:. Maisch argued that 874: 773: 729: 599:appendicular skeleton 499: 277:Maisch & Matzke, 6786:Guizhouichthyosaurus 6708:Guizhouichthyosaurus 6656:Guizhouichthyosaurus 5691:Renesto, S. (2010). 4555:Hulke, J.W. (1873). 3899:"Cranial anatomy of 3666:List of ichthyosaurs 3204:convergently evolved 3173:Guizhouichthyosaurus 3096:fused) metatarsals. 2744:Guizhouichthyosaurus 2410:Guizhouichthyosaurus 2076:Guizhouichthyosaurus 2010:Guizhouichthyosaurus 1972:Guizhouichthyosaurus 1272:to be distinct from 1004:, spanning from the 479:, this region was a 7072:Temnodontosauroidea 5926:10.7717/peerj.10299 5860:2023SwJP..142...15K 5797:2022SwJP..141...12K 5693:"A new specimen of 5636:2011Palgy..54..417M 5577:2023NatSR..1311356F 5518:2021FrEaS...8.8853R 5498:Romano, C. (2021). 5433:Furrer, H. (1995). 5387:2020SwJP..139....2P 5209:2020iSci...23j1347J 5106:Utatsusaurus hataii 5036:2011PNAS..108.8339T 4990:1999JVPal..19..473M 4969:Motani, R. (1999). 4898:Moon, B.C. (2017). 4864:2016JVPal..36E5956J 4744:2020NatSR..10.4206P 4693:2005AREPS..33..395M 4672:Motani, R. (2005). 4526:2013GFF...135...85M 4454:2024SwJP..143...31K 4392:2024SwJP..143...32B 4289:1997PalZ...71..267M 4212:1989JVPal...9..163S 4095:"The Ichthyosauria" 4047:2000PalZ...74....1S 3983:1998PalZ...72..149S 3920:10.7717/peerj.11179 3646:in addition to the 3133:resembles those of 1963:in the same paper. 1232:Pessopteryx nisseri 1195:Pessopteryx nisseri 1155:Friedrich von Huene 1041:named a new genus, 901:Pessosaurus polaris 890:species inquirendae 821:Sander argued that 783:Tübingen University 603:Cristiano Dal Sasso 492:Holotype and naming 389:As an ichthyosaur, 345:Cristiano Dal Sasso 7702:Related categories 7185:Protoichthyosaurus 6604:Barracudasauroides 6489:Quasianosteosaurus 6239:Ichthyosauriformes 6124:Ichthyosauromorpha 6111:Ichthyosauromorpha 6074:Ichthyosauromorpha 5986:Ichthyosauromorpha 5746:Historical Biology 5622:(Bassani, 1886)". 5564:Scientific Reports 5481:Geologia Insubrica 5151:(1898): 20182786. 4732:Scientific Reports 4581:Wiman, C. (1910). 4297:10.1007/BF02988496 4055:10.1007/BF02987949 3991:10.1007/BF02987823 3710:Dinosaurs of Italy 3523: 3509:The head shape of 3496:cartilaginous fish 3339:carbonate platform 3244: 3129:The long snout of 3031:Ichthyosaurs like 3029: 1979: 1658: 1583:Vertebrae and ribs 1493:. Above this, the 1455:of the genus. The 1432: 1353: 1335:, indicating that 1298:Ichthyosaurs like 1296: 1138:, since the eight 1083: 877: 779: 750:Universität Zürich 738: 610:, and made it the 509: 415:coronoid processes 7919: 7918: 7891:Open Tree of Life 7811:Open Tree of Life 7743:Taxon identifiers 7734: 7733: 7730: 7729: 7722:Triassic reptiles 7692: 7691: 7667: 7666: 7663: 7662: 7659: 7658: 7655: 7654: 7651: 7650: 7647: 7646: 7643: 7642: 7639: 7638: 7503:Kazakhstanosaurus 7417: 7416: 7378:Ophthalmosaurinae 7343:Myobradypterygius 7299:Ophthalmosauridae 7195: 7194: 7140: 7139: 7136: 7135: 6972: 6971: 6942: 6941: 6938: 6937: 6873: 6872: 6869: 6868: 6865: 6864: 6861: 6860: 6824:Qianichthyosaurus 6562: 6561: 6451: 6450: 6426: 6425: 6422: 6421: 6418: 6417: 6414: 6413: 6410: 6409: 6386: 6385: 6233: 6232: 6229: 6228: 6204:Parahupehsuchinae 6100: 6099: 6096: 6095: 6092: 6091: 5088:978-1-58834-727-5 5030:(20): 8339–8344. 4614:Palaeontographica 3720:978-0-253-34514-1 3291:Palaeoenvironment 3011: 3010: 3003: 3002: 2994: 2993: 2985: 2984: 2976: 2975: 2967: 2966: 2873: 2872: 2864: 2863: 2855: 2854: 2781: 2780: 2772: 2771: 2676: 2675: 2667: 2666: 2658: 2657: 2649: 2648: 2640: 2639: 2546: 2545: 2537: 2536: 2528: 2527: 2519: 2518: 2510: 2509: 2349: 2348: 2340: 2339: 2331: 2330: 2322: 2321: 2313: 2312: 2304: 2303: 2295: 2294: 2286: 2285: 2277: 2276: 2268: 2267: 1696:(collarbones) of 1491:occipital condyle 1198:, comes from the 1179:Baden-Württemberg 531:Monte San Giorgio 323:Monte San Giorgio 287: 286: 281: 237: 16:(Redirected from 7954: 7937:Fossils of Italy 7912: 7911: 7899: 7898: 7886: 7885: 7873: 7872: 7860: 7859: 7858: 7832: 7831: 7819: 7818: 7806: 7805: 7793: 7792: 7780: 7779: 7770: 7769: 7768: 7738: 7698: 7687: 7681: 7675: 7623:Simbirskiasaurus 7572:Platypterygiinae 7540:Pervushovisaurus 7428: 7423:Platypterygiidae 7304: 7266: 7255: 7228:Stenopterygiidae 7208: 7157: 7153: 7084:Temnodontosaurus 7078: 7068: 7035: 7029:Neoichthyosauria 7002: 6991: 6978: 6967: 6961: 6956: 6950: 6879: 6756: 6641: 6583:Cymbospondylidae 6575: 6520:Cymbospondylidae 6516: 6481: 6470: 6457: 6446: 6440: 6434: 6335: 6326: 6287: 6244: 6185: 6154: 6130: 6119: 6106: 6052: 6043: 6036: 6035: 5994: 5979: 5972: 5965: 5956: 5949: 5948: 5938: 5928: 5898: 5892: 5891: 5881: 5871: 5835: 5829: 5828: 5818: 5808: 5776: 5770: 5769: 5733: 5727: 5726: 5716: 5688: 5679: 5678: 5662: 5656: 5655: 5615: 5609: 5608: 5598: 5588: 5554: 5548: 5547: 5529: 5495: 5489: 5488: 5472: 5447: 5446: 5430: 5417: 5416: 5398: 5364: 5353: 5352: 5342: 5314: 5305: 5304: 5294: 5284: 5260: 5254: 5253: 5245: 5239: 5238: 5228: 5188: 5179: 5178: 5168: 5136: 5130: 5129: 5127: 5099: 5093: 5092: 5074: 5068: 5067: 5057: 5047: 5015: 5009: 5008: 5006: 5000:. Archived from 4975: 4966: 4960: 4959: 4953: 4945: 4943: 4942: 4904: 4895: 4884: 4883: 4847: 4836: 4835: 4819: 4813: 4812: 4780: 4774: 4773: 4763: 4723: 4712: 4711: 4709: 4703:. Archived from 4678: 4669: 4660: 4659: 4631: 4622: 4621: 4605: 4599: 4598: 4588: 4578: 4569: 4568: 4552: 4546: 4545: 4505: 4486: 4485: 4475: 4465: 4433: 4424: 4423: 4413: 4403: 4367: 4352: 4351: 4341: 4332: 4309: 4308: 4264: 4247: 4246: 4230: 4224: 4223: 4187: 4181: 4180: 4168: 4159: 4158: 4138: 4107: 4106: 4090: 4067: 4066: 4026: 4003: 4002: 3958: 3943: 3942: 3932: 3922: 3894: 3809: 3808: 3788: 3725: 3724: 3704: 3582:pachypleurosaurs 3552:M. kuhnschnyderi 3479:actinopterygians 3447:thylacocephalans 3319: 3303: 3123: 3112: 3100:Diet and feeding 3074:thermoregulation 2935: 2911: 2887: 2819: 2795: 2736: 2712: 2705: 2698: 2688: 2608: 2584: 2560: 2474: 2450: 2426: 2402: 2378: 2371: 2361: 2236: 2212: 2188: 2164: 2140: 2116: 2092: 2068: 2044: 2034: 2026: 1957:Guanlingsauridae 1788: 1777: 1676:are shaped like 1577:set into sockets 1546:coronoid process 1310:for limbs and a 1293:Life restoration 1059:they considered 1037:instead, before 740:The holotype of 691: 688: 685: 677: 674: 671: 667: 664: 661: 641: 638: 629: 573:, wedges, and a 547:Besano Formation 524: 521: 518: 487:History of study 473:Besano Formation 433:are elliptical. 411:set into sockets 397:for limbs and a 336:B. leptorhynchus 306: 303: 297: 276: 229: 222: 209: 196: 146: 145: 123: 113: 50: 40:Temporal range: 32: 21: 7962: 7961: 7957: 7956: 7955: 7953: 7952: 7951: 7922: 7921: 7920: 7915: 7907: 7902: 7894: 7889: 7881: 7876: 7868: 7863: 7854: 7853: 7848: 7835: 7827: 7822: 7814: 7809: 7801: 7796: 7788: 7783: 7775: 7773: 7764: 7763: 7758: 7745: 7735: 7726: 7717:Marine reptiles 7703: 7688: 7635: 7566: 7466:Brachypterygius 7458:Athabascasaurus 7450:Arthropterygius 7413: 7408:Ophthalmosaurus 7372: 7312:Arthropterygius 7293: 7262: 7260:Ophthalmosauria 7249: 7222: 7204: 7191: 7161:Ichthyosauridae 7149: 7132: 7087: 7062: 7031: 7022: 6998: 6985: 6968: 6934: 6922:Svalbardosaurus 6901:Pachygonosaurus 6894:Barracudasaurus 6857: 6837: 6808: 6772:Californosaurus 6751:Euichthyosauria 6745: 6677: 6630: 6611:Contectopalatus 6589: 6571: 6558: 6510: 6477: 6464: 6447: 6406: 6382: 6355: 6321:Eoichthyosauria 6315: 6282:Ichthyopterygia 6276: 6248:Omphalosauridae 6225: 6220:Parahupehsuchus 6198: 6174: 6169:Nanchangosaurus 6143: 6126: 6113: 6088: 6068: 6037: 5999: 5998: 5988: 5983: 5953: 5952: 5900: 5899: 5895: 5837: 5836: 5832: 5778: 5777: 5773: 5735: 5734: 5730: 5690: 5689: 5682: 5664: 5663: 5659: 5617: 5616: 5612: 5556: 5555: 5551: 5497: 5496: 5492: 5474: 5473: 5450: 5432: 5431: 5420: 5366: 5365: 5356: 5316: 5315: 5308: 5262: 5261: 5257: 5247: 5246: 5242: 5190: 5189: 5182: 5138: 5137: 5133: 5101: 5100: 5096: 5089: 5076: 5075: 5071: 5017: 5016: 5012: 5004: 4973: 4968: 4967: 4963: 4946: 4940: 4938: 4902: 4897: 4896: 4887: 4858:(1): e1025956. 4849: 4848: 4839: 4821: 4820: 4816: 4782: 4781: 4777: 4725: 4724: 4715: 4707: 4676: 4671: 4670: 4663: 4633: 4632: 4625: 4607: 4606: 4602: 4586: 4580: 4579: 4572: 4554: 4553: 4549: 4507: 4506: 4489: 4435: 4434: 4427: 4369: 4368: 4355: 4344:Palaeodiversity 4339: 4334: 4333: 4312: 4266: 4265: 4250: 4232: 4231: 4227: 4189: 4188: 4184: 4170: 4169: 4162: 4155: 4140: 4139: 4110: 4092: 4091: 4070: 4028: 4027: 4006: 3960: 3959: 3946: 3896: 3895: 3812: 3790: 3789: 3728: 3721: 3706: 3705: 3684: 3679: 3662: 3563:sauropterygians 3487:, though rarer 3410: 3353:, meaning that 3327: 3326: 3325: 3324: 3323: 3320: 3312: 3311: 3304: 3293: 3236: 3144: 3143: 3142: 3141: 3126: 3125: 3124: 3115: 3114: 3113: 3102: 3041:metabolic rates 3017: 3012: 3004: 2995: 2986: 2977: 2968: 2919:Californosaurus 2874: 2865: 2856: 2782: 2773: 2679: 2677: 2668: 2659: 2650: 2641: 2592:Californosaurus 2547: 2538: 2529: 2520: 2511: 2352: 2350: 2341: 2332: 2323: 2314: 2305: 2296: 2287: 2278: 2269: 2220:Californosaurus 1865:Californosaurus 1814: 1813: 1812: 1811: 1791: 1790: 1789: 1780: 1779: 1778: 1767: 1650: 1585: 1562:. The teeth of 1373:sclerotic rings 1345: 1312:fin on the tail 1304:marine reptiles 1286: 1280:from Svalbard. 998: 825:was probably a 756:Californosaurus 724: 689: 686: 683: 675: 672: 669: 665: 662: 659: 644:Varese Province 639: 630: 627: 585:analysis. Each 522: 519: 516: 513:paleontological 494: 489: 399:fin on the tail 316:Middle Triassic 304: 298: 295: 258: 255: 228: 220: 207: 194: 140: 114: 112: 111: 106: 101: 96: 91: 86: 81: 76: 71: 66: 61: 56: 46: 45: 42:Middle Triassic 38: 28: 23: 22: 15: 12: 11: 5: 7960: 7958: 7950: 7949: 7944: 7939: 7934: 7924: 7923: 7917: 7916: 7914: 7913: 7900: 7887: 7874: 7861: 7845: 7843: 7841:Mikadocephalus 7837: 7836: 7834: 7833: 7820: 7807: 7794: 7781: 7771: 7755: 7753: 7747: 7746: 7741: 7732: 7731: 7728: 7727: 7725: 7724: 7719: 7714: 7708: 7705: 7704: 7701: 7694: 7693: 7690: 7689: 7670: 7668: 7665: 7664: 7661: 7660: 7657: 7656: 7653: 7652: 7649: 7648: 7645: 7644: 7641: 7640: 7637: 7636: 7634: 7633: 7626: 7619: 7616:Plutoniosaurus 7612: 7609:Platypterygius 7605: 7598: 7591: 7584: 7581:Caypullisaurus 7576: 7574: 7568: 7567: 7565: 7564: 7557: 7550: 7543: 7536: 7529: 7521: 7514: 7506: 7499: 7491: 7484: 7476: 7469: 7462: 7454: 7446: 7439: 7431: 7425: 7419: 7418: 7415: 7414: 7412: 7411: 7404: 7397: 7390: 7382: 7380: 7374: 7373: 7371: 7370: 7362: 7354: 7350:Nannopterygius 7346: 7339: 7331: 7324: 7316: 7307: 7301: 7295: 7294: 7292: 7291: 7288:Macropterygius 7284: 7281:Delphinosaurus 7277: 7269: 7263: 7258: 7251: 7250: 7248: 7247: 7244:Stenopterygius 7240: 7237:Magnipterygius 7232: 7230: 7224: 7223: 7221: 7220: 7216:Chacaicosaurus 7211: 7205: 7200: 7197: 7196: 7193: 7192: 7190: 7189: 7181: 7173: 7165: 7163: 7150: 7145: 7142: 7141: 7138: 7137: 7134: 7133: 7131: 7130: 7122: 7115: 7111:Excalibosaurus 7107: 7098: 7096: 7089: 7088: 7081: 7075: 7064: 7063: 7061: 7060: 7057:Suevoleviathan 7053: 7046: 7038: 7032: 7027: 7024: 7023: 7021: 7020: 7013: 7005: 6999: 6994: 6987: 6986: 6981: 6974: 6973: 6970: 6969: 6945: 6943: 6940: 6939: 6936: 6935: 6933: 6932: 6925: 6918: 6911: 6904: 6897: 6889: 6887: 6875: 6874: 6871: 6870: 6867: 6866: 6863: 6862: 6859: 6858: 6856: 6855: 6854: 6853: 6847: 6845: 6839: 6838: 6836: 6835: 6828: 6819: 6817: 6814:Toretocnemidae 6810: 6809: 6807: 6806: 6798: 6794:Mikadocephalus 6790: 6782: 6775: 6768: 6764:Auroroborealia 6759: 6753: 6747: 6746: 6744: 6743: 6735: 6728: 6720: 6716:Himalayasaurus 6712: 6704: 6700:Guanlingsaurus 6696: 6687: 6685: 6683:Shastasauridae 6679: 6678: 6676: 6675: 6667: 6659: 6652: 6644: 6638: 6636:Merriamosauria 6632: 6631: 6629: 6628: 6621: 6614: 6607: 6599: 6597: 6591: 6590: 6588: 6587: 6578: 6572: 6567: 6564: 6563: 6560: 6559: 6557: 6556: 6548: 6544:Thalattoarchon 6540: 6536:Phantomosaurus 6532: 6529:Cymbospondylus 6524: 6522: 6512: 6511: 6509: 6508: 6500: 6492: 6484: 6478: 6473: 6466: 6465: 6460: 6453: 6452: 6449: 6448: 6429: 6427: 6424: 6423: 6420: 6419: 6416: 6415: 6412: 6411: 6408: 6407: 6405: 6404: 6403: 6402: 6396: 6394: 6388: 6387: 6384: 6383: 6381: 6380: 6373: 6365: 6363: 6357: 6356: 6354: 6353: 6346: 6338: 6332: 6323: 6317: 6316: 6314: 6313: 6306: 6299: 6295:Isfjordosaurus 6290: 6284: 6278: 6277: 6275: 6274: 6267: 6260: 6252: 6250: 6241: 6235: 6234: 6231: 6230: 6227: 6226: 6224: 6223: 6216: 6208: 6206: 6200: 6199: 6197: 6196: 6188: 6182: 6176: 6175: 6173: 6172: 6165: 6157: 6151: 6145: 6144: 6142: 6141: 6133: 6127: 6122: 6115: 6114: 6109: 6102: 6101: 6098: 6097: 6094: 6093: 6090: 6089: 6087: 6086: 6085: 6084: 6078: 6076: 6070: 6069: 6067: 6066: 6065: 6064: 6055: 6049: 6039: 6038: 6034: 6033: 6024: 6018: 6012: 6006: 5997: 5990: 5989: 5984: 5982: 5981: 5974: 5967: 5959: 5951: 5950: 5893: 5830: 5771: 5752:(3): 257–267. 5728: 5707:(2): 145–160. 5680: 5657: 5630:(2): 417–428. 5610: 5549: 5490: 5448: 5418: 5354: 5333:(3): 511–522. 5306: 5255: 5240: 5180: 5131: 5118:(2): 343–352. 5094: 5087: 5069: 5010: 5007:on 2012-04-15. 4984:(3): 473–496. 4961: 4885: 4837: 4814: 4803:(6): 345–356. 4793:reconstructed" 4787:Mikadocephalus 4775: 4713: 4710:on 2018-12-23. 4661: 4638:Merriamosaurus 4623: 4600: 4570: 4547: 4487: 4425: 4353: 4310: 4283:(3): 267–289. 4248: 4225: 4206:(2): 163–173. 4182: 4160: 4153: 4108: 4068: 4004: 3977:(1): 149–162. 3963:"New finds of 3944: 3810: 3726: 3719: 3681: 3680: 3678: 3675: 3674: 3673: 3668: 3661: 3658: 3650:, long-necked 3626:Cymbospondylus 3620:. While rare, 3616:, potentially 3597:Serpianosaurus 3591:Odoiporosaurus 3519:Cymbospondylus 3489:sarcopterygian 3409: 3406: 3321: 3314: 3313: 3305: 3298: 3297: 3296: 3295: 3294: 3292: 3289: 3266:Mikadocephalus 3235: 3232: 3168:Phragmoteuthis 3139:river dolphins 3128: 3127: 3118: 3117: 3116: 3107: 3106: 3105: 3104: 3103: 3101: 3098: 3094:pathologically 3082:Cymbospondylus 3057:spindle-shaped 3016: 3013: 3009: 3008: 3001: 3000: 2997: 2996: 2992: 2991: 2988: 2987: 2983: 2982: 2979: 2978: 2974: 2973: 2970: 2969: 2965: 2964: 2961: 2960: 2953: 2950: 2949: 2946: 2945: 2942:Toretocnemidae 2938: 2933: 2931: 2928: 2927: 2924: 2923: 2914: 2909: 2907: 2904: 2903: 2900: 2899: 2890: 2885: 2883: 2880: 2879: 2876: 2875: 2871: 2870: 2867: 2866: 2862: 2861: 2858: 2857: 2853: 2852: 2849: 2848: 2839: 2836: 2835: 2832: 2831: 2822: 2817: 2815: 2812: 2811: 2808: 2807: 2803:Guanlingsaurus 2798: 2793: 2791: 2788: 2787: 2784: 2783: 2779: 2778: 2775: 2774: 2770: 2769: 2766: 2765: 2756: 2753: 2752: 2749: 2748: 2739: 2734: 2732: 2729: 2728: 2725: 2724: 2715: 2710: 2708: 2703: 2701: 2696: 2694: 2692:Merriamosauria 2686: 2681: 2674: 2673: 2670: 2669: 2665: 2664: 2661: 2660: 2656: 2655: 2652: 2651: 2647: 2646: 2643: 2642: 2638: 2637: 2634: 2633: 2626: 2623: 2622: 2619: 2618: 2615:Toretocnemidae 2611: 2606: 2604: 2601: 2600: 2597: 2596: 2587: 2582: 2580: 2577: 2576: 2573: 2572: 2563: 2558: 2556: 2553: 2552: 2549: 2548: 2544: 2543: 2540: 2539: 2535: 2534: 2531: 2530: 2526: 2525: 2522: 2521: 2517: 2516: 2513: 2512: 2508: 2507: 2504: 2503: 2494: 2491: 2490: 2487: 2486: 2477: 2472: 2470: 2467: 2466: 2463: 2462: 2458:Guanlingsaurus 2453: 2448: 2446: 2443: 2442: 2439: 2438: 2429: 2424: 2422: 2419: 2418: 2415: 2414: 2405: 2400: 2398: 2395: 2394: 2391: 2390: 2381: 2376: 2374: 2369: 2367: 2365:Merriamosauria 2359: 2354: 2347: 2346: 2343: 2342: 2338: 2337: 2334: 2333: 2329: 2328: 2325: 2324: 2320: 2319: 2316: 2315: 2311: 2310: 2307: 2306: 2302: 2301: 2298: 2297: 2293: 2292: 2289: 2288: 2284: 2283: 2280: 2279: 2275: 2274: 2271: 2270: 2266: 2265: 2262: 2261: 2254: 2251: 2250: 2247: 2246: 2243:Toretocnemidae 2239: 2234: 2232: 2229: 2228: 2225: 2224: 2215: 2210: 2208: 2205: 2204: 2201: 2200: 2191: 2186: 2184: 2181: 2180: 2177: 2176: 2167: 2162: 2160: 2157: 2156: 2153: 2152: 2143: 2138: 2136: 2133: 2132: 2129: 2128: 2124:Guanlingsaurus 2119: 2114: 2112: 2109: 2108: 2105: 2104: 2095: 2090: 2088: 2085: 2084: 2081: 2080: 2071: 2066: 2064: 2061: 2060: 2057: 2056: 2047: 2042: 2040: 2038:Merriamosauria 2032: 2024: 1961:Guanlingsaurus 1953:Mikadocephalus 1929:Mikadocephalus 1921:Mikadocephalus 1900:Cymbospondylus 1887:Merriamosauria 1861:Cymbospondylus 1838:Cymbospondylus 1797:Cymbospondylus 1793: 1792: 1783: 1782: 1781: 1772: 1771: 1770: 1769: 1768: 1766: 1765:Classification 1763: 1682:Cymbospondylus 1649: 1646: 1629:Cymbospondylus 1592:neck vertebrae 1584: 1581: 1559:Phantomosaurus 1499:foramen magnum 1486:Guanlingsaurus 1477:Mikadocephalus 1420:sagittal crest 1408:pineal foramen 1365:external nares 1344: 1341: 1320:Cymbospondylus 1285: 1282: 1245:Isfjordosaurus 1220:Merriamosaurus 1208:omphalosaurids 1200:Early Triassic 1121:Mikadocephalus 1105:gracilirostris 1099:Mikadocephalus 1095:Mikadocephalus 1080:Mikadocephalus 1034:Cymbospondylus 997: 994: 962:Mikadocephalus 950:Mikadocephalus 931:Mikadocephalus 923:Mikadocephalus 854:Mikadocephalus 839:Mikadocephalus 827:junior synonym 823:Mikadocephalus 810:(slender) and 804:pick-up sticks 765:Robert Appleby 723: 721:Mikadocephalus 717: 493: 490: 488: 485: 471:come from the 361:Mikadocephalus 285: 284: 283: 282: 267: 266: 260: 259: 256: 246: 245: 239: 238: 218: 214: 213: 211:Shastasauridae 205: 201: 200: 192: 188: 187: 182: 178: 177: 172: 168: 167: 162: 158: 157: 152: 148: 147: 134: 133: 125: 124: 116: 115: 107: 102: 97: 92: 87: 82: 77: 72: 67: 62: 57: 52: 51: 39: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 7959: 7948: 7945: 7943: 7940: 7938: 7935: 7933: 7930: 7929: 7927: 7910: 7905: 7901: 7897: 7892: 7888: 7884: 7879: 7875: 7871: 7866: 7862: 7857: 7851: 7847: 7846: 7844: 7842: 7838: 7830: 7825: 7821: 7817: 7812: 7808: 7804: 7799: 7795: 7791: 7786: 7782: 7778: 7772: 7767: 7761: 7757: 7756: 7754: 7752: 7748: 7744: 7739: 7723: 7720: 7718: 7715: 7713: 7710: 7709: 7706: 7699: 7695: 7686: 7682: 7680: 7674: 7632: 7631: 7627: 7625: 7624: 7620: 7618: 7617: 7613: 7611: 7610: 7606: 7604: 7603: 7602:Maiaspondylus 7599: 7597: 7596: 7592: 7590: 7589: 7585: 7583: 7582: 7578: 7577: 7575: 7573: 7569: 7563: 7562: 7558: 7556: 7555: 7551: 7549: 7548: 7544: 7542: 7541: 7537: 7535: 7534: 7530: 7527: 7526: 7522: 7520: 7519: 7515: 7512: 7511: 7507: 7505: 7504: 7500: 7497: 7496: 7492: 7490: 7489: 7485: 7482: 7481: 7480:Cryopterygius 7477: 7475: 7474: 7470: 7468: 7467: 7463: 7460: 7459: 7455: 7452: 7451: 7447: 7445: 7444: 7440: 7438: 7437: 7433: 7432: 7429: 7426: 7424: 7420: 7410: 7409: 7405: 7403: 7402: 7398: 7396: 7395: 7391: 7389: 7388: 7387:Acamptonectes 7384: 7383: 7381: 7379: 7375: 7368: 7367: 7366:Thalassodraco 7363: 7360: 7359: 7355: 7352: 7351: 7347: 7345: 7344: 7340: 7337: 7336: 7332: 7330: 7329: 7325: 7322: 7321: 7317: 7314: 7313: 7309: 7308: 7305: 7302: 7300: 7296: 7290: 7289: 7285: 7283: 7282: 7278: 7276: 7275: 7271: 7270: 7267: 7264: 7261: 7256: 7252: 7246: 7245: 7241: 7239: 7238: 7234: 7233: 7231: 7229: 7225: 7218: 7217: 7213: 7212: 7209: 7206: 7203: 7198: 7187: 7186: 7182: 7179: 7178: 7174: 7172: 7171: 7170:Ichthyosaurus 7167: 7166: 7164: 7162: 7158: 7154: 7151: 7148: 7143: 7128: 7127: 7123: 7121: 7120: 7116: 7113: 7112: 7108: 7105: 7104: 7103:Eurhinosaurus 7100: 7099: 7097: 7094: 7093:Leptonectidae 7090: 7086: 7085: 7079: 7076: 7073: 7069: 7065: 7059: 7058: 7054: 7052: 7051: 7050:Hauffiopteryx 7047: 7045: 7044: 7040: 7039: 7036: 7033: 7030: 7025: 7019: 7018: 7014: 7012: 7011: 7010:Hudsonelpidia 7007: 7006: 7003: 7000: 6997: 6992: 6988: 6984: 6979: 6975: 6966: 6962: 6960: 6955: 6949: 6931: 6930: 6926: 6924: 6923: 6919: 6917: 6916: 6915:Sphaerodontes 6912: 6910: 6909: 6905: 6903: 6902: 6898: 6896: 6895: 6891: 6890: 6888: 6886: 6885: 6880: 6876: 6851: 6850: 6849: 6848: 6846: 6844: 6840: 6834: 6833: 6829: 6826: 6825: 6821: 6820: 6818: 6815: 6811: 6804: 6803: 6799: 6796: 6795: 6791: 6788: 6787: 6783: 6781: 6780: 6776: 6774: 6773: 6769: 6766: 6765: 6761: 6760: 6757: 6754: 6752: 6748: 6741: 6740: 6736: 6734: 6733: 6729: 6726: 6725: 6721: 6718: 6717: 6713: 6710: 6709: 6705: 6702: 6701: 6697: 6694: 6693: 6689: 6688: 6686: 6684: 6680: 6673: 6672: 6668: 6665: 6664: 6660: 6658: 6657: 6653: 6651: 6650: 6646: 6645: 6642: 6639: 6637: 6633: 6627: 6626: 6622: 6620: 6619: 6615: 6613: 6612: 6608: 6606: 6605: 6601: 6600: 6598: 6596: 6592: 6585: 6584: 6580: 6579: 6576: 6573: 6570: 6565: 6554: 6553: 6552:Xinminosaurus 6549: 6546: 6545: 6541: 6538: 6537: 6533: 6531: 6530: 6526: 6525: 6523: 6521: 6517: 6513: 6506: 6505: 6504:Xinminosaurus 6501: 6498: 6497: 6493: 6491: 6490: 6486: 6485: 6482: 6479: 6476: 6475:Ichthyosauria 6471: 6467: 6463: 6462:Ichthyosauria 6458: 6454: 6445: 6441: 6439: 6433: 6400: 6399: 6398: 6397: 6395: 6393: 6392:Ichthyosauria 6389: 6379: 6378: 6374: 6372: 6371: 6367: 6366: 6364: 6362: 6358: 6352: 6351: 6347: 6345: 6344: 6340: 6339: 6336: 6333: 6331: 6327: 6324: 6322: 6318: 6312: 6311: 6307: 6305: 6304: 6300: 6297: 6296: 6292: 6291: 6288: 6285: 6283: 6279: 6273: 6272: 6268: 6266: 6265: 6264:Omphalosaurus 6261: 6259: 6258: 6257:Cartorhynchus 6254: 6253: 6251: 6249: 6245: 6242: 6240: 6236: 6222: 6221: 6217: 6215: 6214: 6210: 6209: 6207: 6205: 6201: 6195: 6194: 6190: 6189: 6186: 6183: 6181: 6180:Hupehsuchidae 6177: 6171: 6170: 6166: 6164: 6163: 6162:Eohupehsuchus 6159: 6158: 6155: 6152: 6150: 6146: 6140: 6139: 6135: 6134: 6131: 6128: 6125: 6120: 6116: 6112: 6107: 6103: 6082: 6081: 6080: 6079: 6077: 6075: 6071: 6063: 6059: 6058: 6057: 6056: 6053: 6050: 6048: 6044: 6040: 6032: 6028: 6025: 6023: 6019: 6017: 6013: 6011: 6007: 6005: 6001: 6000: 5995: 5991: 5987: 5980: 5975: 5973: 5968: 5966: 5961: 5960: 5957: 5946: 5942: 5937: 5932: 5927: 5922: 5918: 5914: 5913: 5908: 5906: 5897: 5894: 5889: 5885: 5880: 5875: 5870: 5865: 5861: 5857: 5853: 5849: 5845: 5843: 5834: 5831: 5826: 5822: 5817: 5812: 5807: 5802: 5798: 5794: 5790: 5786: 5782: 5775: 5772: 5767: 5763: 5759: 5755: 5751: 5747: 5743: 5741: 5732: 5729: 5724: 5720: 5715: 5710: 5706: 5702: 5698: 5696: 5687: 5685: 5681: 5676: 5672: 5668: 5661: 5658: 5653: 5649: 5645: 5641: 5637: 5633: 5629: 5625: 5624:Palaeontology 5621: 5614: 5611: 5606: 5602: 5597: 5592: 5587: 5582: 5578: 5574: 5570: 5566: 5565: 5560: 5553: 5550: 5545: 5541: 5537: 5533: 5528: 5523: 5519: 5515: 5511: 5507: 5506: 5501: 5494: 5491: 5486: 5482: 5478: 5471: 5469: 5467: 5465: 5463: 5461: 5459: 5457: 5455: 5453: 5449: 5445:(3): 827–852. 5444: 5440: 5436: 5429: 5427: 5425: 5423: 5419: 5414: 5410: 5406: 5402: 5397: 5392: 5388: 5384: 5380: 5376: 5375: 5370: 5363: 5361: 5359: 5355: 5350: 5346: 5341: 5336: 5332: 5328: 5324: 5322: 5313: 5311: 5307: 5302: 5298: 5293: 5288: 5283: 5278: 5274: 5270: 5266: 5259: 5256: 5251: 5244: 5241: 5236: 5232: 5227: 5222: 5218: 5214: 5210: 5206: 5203:(9): 101347. 5202: 5198: 5194: 5187: 5185: 5181: 5176: 5172: 5167: 5162: 5158: 5154: 5150: 5146: 5142: 5135: 5132: 5126: 5121: 5117: 5113: 5109: 5107: 5098: 5095: 5090: 5084: 5080: 5073: 5070: 5065: 5061: 5056: 5051: 5046: 5041: 5037: 5033: 5029: 5025: 5021: 5014: 5011: 5003: 4999: 4995: 4991: 4987: 4983: 4979: 4972: 4965: 4962: 4957: 4951: 4936: 4932: 4928: 4924: 4920: 4916: 4912: 4908: 4901: 4894: 4892: 4890: 4886: 4881: 4877: 4873: 4869: 4865: 4861: 4857: 4853: 4846: 4844: 4842: 4838: 4833: 4829: 4825: 4818: 4815: 4810: 4806: 4802: 4798: 4794: 4792: 4788: 4779: 4776: 4771: 4767: 4762: 4757: 4753: 4749: 4745: 4741: 4737: 4733: 4729: 4722: 4720: 4718: 4714: 4706: 4702: 4698: 4694: 4690: 4686: 4682: 4675: 4668: 4666: 4662: 4657: 4653: 4649: 4645: 4641: 4639: 4630: 4628: 4624: 4619: 4616:(in German). 4615: 4611: 4604: 4601: 4596: 4593:(in German). 4592: 4584: 4577: 4575: 4571: 4566: 4562: 4558: 4551: 4548: 4543: 4539: 4535: 4531: 4527: 4523: 4519: 4515: 4511: 4504: 4502: 4500: 4498: 4496: 4494: 4492: 4488: 4483: 4479: 4474: 4469: 4464: 4459: 4455: 4451: 4447: 4443: 4439: 4432: 4430: 4426: 4421: 4417: 4412: 4407: 4402: 4397: 4393: 4389: 4385: 4381: 4377: 4375: 4366: 4364: 4362: 4360: 4358: 4354: 4349: 4345: 4338: 4331: 4329: 4327: 4325: 4323: 4321: 4319: 4317: 4315: 4311: 4306: 4302: 4298: 4294: 4290: 4286: 4282: 4278: 4274: 4272: 4263: 4261: 4259: 4257: 4255: 4253: 4249: 4244: 4240: 4236: 4229: 4226: 4221: 4217: 4213: 4209: 4205: 4201: 4197: 4195: 4186: 4183: 4179:(3): 21A–22A. 4178: 4174: 4167: 4165: 4161: 4156: 4154:3-89937-007-4 4150: 4146: 4145: 4137: 4135: 4133: 4131: 4129: 4127: 4125: 4123: 4121: 4119: 4117: 4115: 4113: 4109: 4104: 4100: 4096: 4089: 4087: 4085: 4083: 4081: 4079: 4077: 4075: 4073: 4069: 4064: 4060: 4056: 4052: 4048: 4044: 4040: 4036: 4032: 4025: 4023: 4021: 4019: 4017: 4015: 4013: 4011: 4009: 4005: 4000: 3996: 3992: 3988: 3984: 3980: 3976: 3972: 3968: 3966: 3965:Omphalosaurus 3957: 3955: 3953: 3951: 3949: 3945: 3940: 3936: 3931: 3926: 3921: 3916: 3912: 3908: 3904: 3902: 3893: 3891: 3889: 3887: 3885: 3883: 3881: 3879: 3877: 3875: 3873: 3871: 3869: 3867: 3865: 3863: 3861: 3859: 3857: 3855: 3853: 3851: 3849: 3847: 3845: 3843: 3841: 3839: 3837: 3835: 3833: 3831: 3829: 3827: 3825: 3823: 3821: 3819: 3817: 3815: 3811: 3806: 3802: 3798: 3796: 3787: 3785: 3783: 3781: 3779: 3777: 3775: 3773: 3771: 3769: 3767: 3765: 3763: 3761: 3759: 3757: 3755: 3753: 3751: 3749: 3747: 3745: 3743: 3741: 3739: 3737: 3735: 3733: 3731: 3727: 3722: 3716: 3712: 3711: 3703: 3701: 3699: 3697: 3695: 3693: 3691: 3689: 3687: 3683: 3676: 3672: 3669: 3667: 3664: 3663: 3659: 3657: 3655: 3654: 3653:Tanystropheus 3649: 3645: 3644: 3639: 3638: 3633: 3632: 3631:Askeptosaurus 3627: 3623: 3619: 3615: 3611: 3610: 3605: 3604: 3599: 3598: 3593: 3592: 3587: 3583: 3579: 3578: 3573: 3572: 3568: 3564: 3559: 3557: 3553: 3549: 3548: 3543: 3539: 3535: 3534:apex predator 3531: 3527: 3520: 3517:(middle) and 3516: 3512: 3507: 3503: 3501: 3497: 3493: 3490: 3486: 3485: 3480: 3476: 3472: 3468: 3464: 3460: 3456: 3452: 3448: 3444: 3440: 3436: 3432: 3428: 3424: 3423: 3419: 3415: 3414:invertebrates 3407: 3405: 3403: 3399: 3395: 3391: 3387: 3386:free swimming 3383: 3379: 3375: 3371: 3366: 3364: 3360: 3356: 3352: 3348: 3347:N. secedensis 3344: 3340: 3336: 3332: 3318: 3309: 3306:Map of where 3302: 3290: 3288: 3286: 3281: 3279: 3275: 3271: 3267: 3261: 3258: 3253: 3249: 3240: 3233: 3231: 3229: 3225: 3221: 3217: 3213: 3209: 3205: 3200: 3195: 3193: 3189: 3188: 3183: 3178: 3174: 3170: 3169: 3164: 3160: 3156: 3153: 3149: 3140: 3136: 3132: 3122: 3111: 3099: 3097: 3095: 3091: 3087: 3083: 3079: 3075: 3071: 3067: 3063: 3058: 3054: 3050: 3046: 3042: 3038: 3034: 3026: 3023:Skull of the 3021: 3014: 3007: 2999: 2998: 2990: 2989: 2981: 2980: 2972: 2971: 2963: 2962: 2959: 2958: 2952: 2951: 2948: 2947: 2944: 2943: 2937: 2936: 2930: 2929: 2926: 2925: 2922: 2921: 2920: 2913: 2912: 2906: 2905: 2902: 2901: 2898: 2897: 2896: 2889: 2888: 2882: 2881: 2878: 2877: 2869: 2868: 2860: 2859: 2851: 2850: 2847: 2846: 2845: 2838: 2837: 2834: 2833: 2830: 2829: 2828: 2821: 2820: 2814: 2813: 2810: 2809: 2806: 2805: 2804: 2797: 2796: 2790: 2789: 2786: 2785: 2777: 2776: 2768: 2767: 2764: 2763: 2762: 2755: 2754: 2751: 2750: 2747: 2746: 2745: 2738: 2737: 2731: 2730: 2727: 2726: 2723: 2722: 2721: 2714: 2713: 2707: 2706: 2700: 2699: 2693: 2690: 2689: 2685: 2682: 2680: 2672: 2671: 2663: 2662: 2654: 2653: 2645: 2644: 2636: 2635: 2632: 2631: 2625: 2624: 2621: 2620: 2617: 2616: 2610: 2609: 2603: 2602: 2599: 2598: 2595: 2594: 2593: 2586: 2585: 2579: 2578: 2575: 2574: 2571: 2570: 2569: 2562: 2561: 2555: 2554: 2551: 2550: 2542: 2541: 2533: 2532: 2524: 2523: 2515: 2514: 2506: 2505: 2502: 2501: 2500: 2493: 2492: 2489: 2488: 2485: 2484: 2483: 2476: 2475: 2469: 2468: 2465: 2464: 2461: 2460: 2459: 2452: 2451: 2445: 2444: 2441: 2440: 2437: 2436: 2435: 2428: 2427: 2421: 2420: 2417: 2416: 2413: 2412: 2411: 2404: 2403: 2397: 2396: 2393: 2392: 2389: 2388: 2387: 2380: 2379: 2373: 2372: 2366: 2363: 2362: 2358: 2355: 2353: 2345: 2344: 2336: 2335: 2327: 2326: 2318: 2317: 2309: 2308: 2300: 2299: 2291: 2290: 2282: 2281: 2273: 2272: 2264: 2263: 2260: 2259: 2253: 2252: 2249: 2248: 2245: 2244: 2238: 2237: 2231: 2230: 2227: 2226: 2223: 2222: 2221: 2214: 2213: 2207: 2206: 2203: 2202: 2199: 2198: 2197: 2190: 2189: 2183: 2182: 2179: 2178: 2175: 2174: 2173: 2166: 2165: 2159: 2158: 2155: 2154: 2151: 2150: 2149: 2142: 2141: 2135: 2134: 2131: 2130: 2127: 2126: 2125: 2118: 2117: 2111: 2110: 2107: 2106: 2103: 2102: 2101: 2094: 2093: 2087: 2086: 2083: 2082: 2079: 2078: 2077: 2070: 2069: 2063: 2062: 2059: 2058: 2055: 2054: 2053: 2046: 2045: 2039: 2036: 2035: 2031: 2028: 2027: 2023: 2020: 2018: 2017: 2012: 2011: 2006: 2001: 1997: 1992: 1988: 1984: 1977: 1973: 1968: 1964: 1962: 1958: 1954: 1950: 1946: 1943: 1939: 1935: 1930: 1926: 1922: 1918: 1912: 1910: 1905: 1901: 1897: 1893: 1888: 1884: 1883: 1878: 1874: 1870: 1866: 1862: 1856: 1854: 1850: 1849: 1844: 1843:natural group 1839: 1835: 1831: 1827: 1823: 1819: 1809: 1808: 1803: 1799: 1798: 1787: 1776: 1764: 1762: 1760: 1755: 1751: 1747: 1743: 1739: 1735: 1730: 1728: 1724: 1720: 1716: 1712: 1707: 1703: 1702:interclavicle 1699: 1695: 1691: 1687: 1683: 1679: 1675: 1671: 1667: 1663: 1654: 1647: 1645: 1643: 1639: 1634: 1630: 1625: 1620: 1618: 1614: 1613:cervical ribs 1610: 1606: 1601: 1600:neural spines 1597: 1593: 1589: 1582: 1580: 1578: 1573: 1569: 1565: 1561: 1560: 1555: 1551: 1547: 1543: 1542: 1536: 1532: 1528: 1523: 1520: 1516: 1512: 1508: 1504: 1500: 1496: 1492: 1488: 1487: 1482: 1478: 1474: 1469: 1467: 1463: 1458: 1454: 1453:autapomorphic 1450: 1445: 1441: 1437: 1428: 1424: 1421: 1417: 1414:in front and 1413: 1409: 1405: 1401: 1397: 1392: 1390: 1386: 1382: 1378: 1374: 1370: 1366: 1362: 1358: 1349: 1342: 1340: 1338: 1334: 1329: 1325: 1321: 1317: 1313: 1309: 1305: 1301: 1294: 1290: 1283: 1281: 1279: 1275: 1271: 1267: 1263: 1259: 1255: 1251: 1247: 1246: 1241: 1237: 1233: 1229: 1225: 1221: 1217: 1216:Rotundopteryx 1213: 1209: 1205: 1201: 1197: 1196: 1190: 1188: 1184: 1180: 1176: 1172: 1168: 1164: 1160: 1156: 1151: 1149: 1145: 1141: 1137: 1133: 1128: 1126: 1122: 1118: 1114: 1110: 1106: 1103: 1100: 1096: 1092: 1088: 1081: 1077: 1072: 1068: 1066: 1062: 1057: 1052: 1048: 1044: 1040: 1036: 1035: 1030: 1026: 1022: 1020: 1019:Ichthyosaurus 1015: 1014:Late Triassic 1012:stage of the 1011: 1007: 1003: 995: 993: 991: 987: 983: 979: 975: 971: 967: 963: 959: 955: 951: 947: 943: 938: 936: 932: 928: 924: 920: 916: 912: 908: 907: 902: 898: 897: 892: 891: 886: 882: 873: 869: 867: 863: 859: 855: 850: 846: 845: 840: 836: 832: 828: 824: 819: 817: 813: 809: 805: 801: 797: 793: 788: 784: 777: 772: 768: 766: 762: 758: 757: 751: 747: 743: 735: 734: 728: 722: 718: 716: 714: 710: 706: 705: 700: 696: 681: 657: 653: 649: 645: 637: 633: 625: 621: 617: 613: 609: 604: 600: 596: 591: 588: 584: 578: 576: 572: 571:sledgehammers 567: 564:, a piece of 563: 560: 556: 552: 548: 544: 540: 536: 532: 528: 514: 507: 503: 498: 491: 486: 484: 482: 478: 475:. During the 474: 470: 466: 462: 458: 455: 451: 447: 443: 439: 434: 432: 428: 424: 420: 416: 412: 408: 404: 400: 396: 392: 387: 385: 381: 377: 374: 370: 366: 362: 358: 354: 350: 346: 343:was named by 342: 338: 337: 332: 328: 324: 320: 317: 313: 310: 302: 299:reptile from 293: 292: 280: 275: 271: 270: 268: 265: 261: 254: 253: 247: 244: 240: 236: 233:& Pinna, 232: 227: 226: 219: 216: 215: 212: 206: 203: 202: 199: 198:Ichthyosauria 193: 190: 189: 186: 183: 180: 179: 176: 173: 170: 169: 166: 163: 160: 159: 156: 153: 150: 149: 144: 139: 135: 131: 126: 122: 117: 110: 105: 100: 95: 90: 85: 80: 75: 70: 65: 60: 55: 49: 43: 37: 33: 30: 19: 7840: 7751:Besanosaurus 7750: 7712:Ichthyosaurs 7676: 7628: 7621: 7614: 7607: 7600: 7593: 7586: 7579: 7561:Undorosaurus 7559: 7554:Sveltonectes 7552: 7545: 7538: 7533:Parrassaurus 7531: 7523: 7518:Muiscasaurus 7516: 7508: 7501: 7493: 7486: 7478: 7473:Catutosaurus 7471: 7464: 7456: 7448: 7443:Aegirosaurus 7441: 7434: 7406: 7399: 7392: 7385: 7364: 7356: 7348: 7341: 7333: 7328:Jabalisaurus 7326: 7318: 7310: 7286: 7279: 7274:Argovisaurus 7272: 7242: 7235: 7214: 7183: 7175: 7168: 7147:Thunnosauria 7124: 7117: 7109: 7101: 7082: 7055: 7048: 7041: 7015: 7008: 6951: 6929:Tibetosaurus 6927: 6920: 6913: 6906: 6899: 6892: 6884:Nomina dubia 6882: 6832:Toretocnemus 6830: 6822: 6800: 6792: 6784: 6777: 6770: 6762: 6737: 6732:Shastasaurus 6730: 6724:Ichthyotitan 6722: 6714: 6706: 6698: 6692:Besanosaurus 6691: 6690: 6669: 6661: 6654: 6649:Besanosaurus 6648: 6647: 6623: 6616: 6609: 6602: 6595:Mixosauridae 6581: 6569:Hueneosauria 6550: 6542: 6534: 6527: 6502: 6494: 6487: 6435: 6375: 6368: 6350:Utatsusaurus 6348: 6343:Parvinatator 6341: 6308: 6303:Chaohusaurus 6301: 6293: 6271:Sclerocormus 6269: 6262: 6255: 6218: 6213:Eretmorhipis 6211: 6191: 6167: 6160: 6136: 6026: 5916: 5910: 5904: 5896: 5851: 5847: 5841: 5833: 5788: 5784: 5774: 5749: 5745: 5739: 5731: 5704: 5700: 5694: 5674: 5670: 5660: 5627: 5623: 5619: 5613: 5571:(1): 11356. 5568: 5562: 5552: 5509: 5503: 5493: 5484: 5480: 5442: 5438: 5378: 5372: 5330: 5326: 5320: 5272: 5268: 5258: 5249: 5243: 5200: 5196: 5148: 5144: 5134: 5115: 5111: 5105: 5097: 5078: 5072: 5027: 5023: 5013: 5002:the original 4981: 4977: 4964: 4950:cite journal 4939:. Retrieved 4910: 4906: 4855: 4851: 4831: 4827: 4817: 4800: 4796: 4790: 4786: 4778: 4735: 4731: 4705:the original 4684: 4680: 4650:(1): 59–94. 4647: 4643: 4637: 4617: 4613: 4603: 4594: 4590: 4564: 4560: 4550: 4520:(1): 85–94. 4517: 4513: 4445: 4441: 4383: 4379: 4373: 4347: 4343: 4280: 4276: 4270: 4242: 4238: 4228: 4203: 4199: 4193: 4185: 4176: 4172: 4143: 4102: 4098: 4038: 4034: 3974: 3970: 3964: 3910: 3906: 3900: 3804: 3800: 3794: 3709: 3651: 3641: 3635: 3629: 3625: 3622:N. giganteus 3621: 3618:N. juvenilis 3617: 3613: 3607: 3601: 3595: 3589: 3586:nothosaurids 3575: 3571:Paraplacodus 3569: 3560: 3555: 3551: 3545: 3541: 3537: 3529: 3526:Besanosaurus 3525: 3524: 3518: 3514: 3511:Besanosaurus 3510: 3482: 3463:Radiolarians 3420: 3411: 3398:Besanosaurus 3397: 3390:anoxic water 3374:Tethys Ocean 3367: 3359:Besanosaurus 3358: 3355:Besanosaurus 3354: 3346: 3343:Besanosaurus 3342: 3331:Besanosaurus 3330: 3328: 3308:Besanosaurus 3307: 3285:Besanosaurus 3284: 3282: 3278:Besanosaurus 3277: 3274:Besanosaurus 3273: 3270:Besanosaurus 3269: 3265: 3262: 3257:Besanosaurus 3256: 3252:Besanosaurus 3251: 3245: 3224:Besanosaurus 3223: 3220:Besanosaurus 3219: 3199:Besanosaurus 3198: 3196: 3192:Besanosaurus 3191: 3185: 3182:Besanosaurus 3181: 3177:thalattosaur 3172: 3166: 3163:Besanosaurus 3162: 3159:Besanosaurus 3158: 3148:Besanosaurus 3147: 3145: 3131:Besanosaurus 3130: 3086:Besanosaurus 3085: 3081: 3078:Besanosaurus 3077: 3062:Besanosaurus 3061: 3049:anguilliform 3037:ossification 3033:Besanosaurus 3032: 3030: 3025:Besanosaurus 3024: 3015:Paleobiology 3005: 2955: 2940: 2917: 2916: 2893: 2892: 2842: 2841: 2827:Shastasaurus 2825: 2824: 2801: 2800: 2761:Besanosaurus 2760: 2759: 2758: 2742: 2741: 2718: 2717: 2683: 2678: 2628: 2613: 2590: 2589: 2566: 2565: 2497: 2496: 2482:Shastasaurus 2480: 2479: 2456: 2455: 2432: 2431: 2408: 2407: 2386:Besanosaurus 2385: 2384: 2383: 2356: 2351: 2256: 2241: 2218: 2217: 2194: 2193: 2170: 2169: 2148:Shastasaurus 2146: 2145: 2122: 2121: 2098: 2097: 2074: 2073: 2052:Besanosaurus 2051: 2050: 2049: 2029: 2021: 2014: 2008: 2005:Besanosaurus 2004: 2000:Besanosaurus 1999: 1996:Besanosaurus 1995: 1991:Besanosaurus 1990: 1983:Besanosaurus 1982: 1980: 1976:Besanosaurus 1975: 1971: 1970:Skeleton of 1960: 1959:, alongside 1952: 1949:Besanosaurus 1948: 1938:Besanosaurus 1937: 1928: 1925:Besanosaurus 1924: 1920: 1917:Besanosaurus 1916: 1913: 1903: 1899: 1896:sister taxon 1892:Besanosaurus 1891: 1880: 1877:Shastasaurus 1876: 1873:Besanosaurus 1872: 1869:Besanosaurus 1868: 1864: 1860: 1857: 1853:paraphyletic 1846: 1837: 1834:Besanosaurus 1833: 1830:Besanosaurus 1829: 1822:Besanosaurus 1821: 1818:Besanosaurus 1817: 1815: 1807:Shastasaurus 1805: 1802:Besanosaurus 1801: 1795: 1734:Besanosaurus 1733: 1731: 1706:Besanosaurus 1705: 1698:Besanosaurus 1697: 1690:Besanosaurus 1689: 1681: 1670:Besanosaurus 1669: 1659: 1642:Besanosaurus 1641: 1633:Besanosaurus 1632: 1628: 1624:Besanosaurus 1623: 1621: 1596:zygapophyses 1588:Besanosaurus 1587: 1586: 1568:tooth crowns 1564:Besanosaurus 1563: 1557: 1554:Besanosaurus 1553: 1541:Chaohusaurus 1539: 1525:Most of the 1524: 1511:Besanosaurus 1510: 1503:Besanosaurus 1502: 1484: 1481:Besanosaurus 1480: 1476: 1470: 1466:Besanosaurus 1465: 1449:postorbitals 1444:postfrontals 1433: 1393: 1389:Besanosaurus 1388: 1381:parvipelvian 1377:Besanosaurus 1376: 1361:Besanosaurus 1360: 1357:Besanosaurus 1356: 1354: 1337:Besanosaurus 1336: 1333:Besanosaurus 1332: 1328:Besanosaurus 1327: 1324:Besanosaurus 1323: 1319: 1316:Besanosaurus 1315: 1300:Besanosaurus 1299: 1297: 1278:Besanosaurus 1277: 1274:Besanosaurus 1273: 1269: 1266:Besanosaurus 1265: 1261: 1257: 1254:Besanosaurus 1253: 1249: 1243: 1240:Besanosaurus 1239: 1236:nomen dubium 1235: 1231: 1227: 1223: 1219: 1215: 1211: 1193: 1191: 1187:Besanosaurus 1186: 1183:nomen dubium 1182: 1171:Black Forest 1162: 1158: 1152: 1148:Besanosaurus 1147: 1143: 1136:nomen dubium 1135: 1131: 1129: 1124: 1120: 1116: 1113:nomen dubium 1112: 1108: 1104: 1098: 1094: 1086: 1084: 1079: 1075: 1065:nomen dubium 1064: 1060: 1055: 1050: 1046: 1042: 1032: 1029:Shastasaurus 1028: 1017: 999: 990:Besanosaurus 989: 985: 982:Besanosaurus 981: 978:Besanosaurus 977: 973: 969: 966:Besanosaurus 965: 961: 958:Besanosaurus 957: 949: 946:Besanosaurus 945: 942:Besanosaurus 941: 939: 935:Besanosaurus 934: 930: 927:Besanosaurus 926: 922: 918: 914: 906:nomen dubium 904: 900: 894: 888: 884: 880: 878: 866:Besanosaurus 865: 861: 858:Besanosaurus 857: 853: 849:Besanosaurus 848: 842: 838: 835:Besanosaurus 834: 831:Besanosaurus 830: 822: 820: 816:Besanosaurus 815: 811: 807: 799: 791: 787:air abrasion 780: 775: 761:Besanosaurus 760: 754: 742:Besanosaurus 741: 739: 731: 720: 709:Besanosaurus 708: 704:Shastasaurus 702: 699:Besanosaurus 698: 695:Besanosaurus 694: 679: 655: 650:, while the 623: 592: 579: 562:ichthyosaurs 510: 506:Besanosaurus 505: 469:Besanosaurus 468: 446:Besanosaurus 445: 442:Besanosaurus 441: 438:shastasaurid 435: 431:finger bones 407:Besanosaurus 406: 403:Besanosaurus 402: 391:Besanosaurus 390: 388: 369:Besanosaurus 368: 365:Besanosaurus 364: 360: 341:Besanosaurus 340: 335: 334: 291:Besanosaurus 290: 289: 288: 273: 251: 250: 243:Type species 225:Besanosaurus 224: 223: 36:Besanosaurus 35: 29: 7436:Acuetzpalin 7401:Mollesaurus 7320:Gengasaurus 7126:Wahlisaurus 7119:Leptonectes 6996:Parvipelvia 6983:Parvipelvia 6908:Pessosaurus 6843:Parvipelvia 6802:Shonisaurus 6739:Shonisaurus 6663:Pessopteryx 6193:Hupehsuchus 6149:Hupehsuchia 6138:Baisesaurus 6031:Neodiapsida 5695:Nothosaurus 4913:(2): 1–27. 4834:(1): 71–82. 4738:(1): 4206. 4687:: 395–420. 4041:(1): 1–35. 3648:semiaquatic 3643:Hescheleria 3614:Nothosaurus 3580:as well as 3492:coelacanths 3484:Saurichthys 3455:brachiopods 3216:piscivorous 3155:cephalopods 2957:Parvipelvia 2844:Shonisaurus 2630:Parvipelvia 2499:Shonisaurus 2258:Parvipelvia 2172:Shonisaurus 1904:Shonisaurus 1882:Shonisaurus 1759:metatarsals 1742:pubic bones 1723:metacarpals 1572:tooth roots 1535:surangulars 1495:spinal cord 1440:prefrontals 1396:premaxillae 1284:Description 1270:Pessopteryx 1262:Pessopteryx 1258:Pessopteryx 1250:Pessopteryx 1228:Pessopteryx 1224:Pessopteryx 1212:Pessopteryx 1167:Muschelkalk 1163:P. suevicus 1159:Pessosaurus 1144:Pessosaurus 1132:Pessosaurus 1125:Pessosaurus 1117:Pessosaurus 1109:Pessosaurus 1087:Pessosaurus 1076:Pessosaurus 1061:Pessosaurus 1056:Pessosaurus 1051:Pessosaurus 1043:Pessosaurus 896:Pessosaurus 746:Switzerland 713:valid taxon 608:shastasaurs 595:preparation 457:cephalopods 376:archipelago 331:Switzerland 319:ichthyosaur 7926:Categories 7630:Sisteronia 7588:Kyhytysuka 7495:Janusaurus 7488:Grendelius 7394:Baptanodon 7335:Janusaurus 7202:Baracromia 7043:Dearcmhara 7017:Macgowania 6852:see below↓ 6779:Callawayia 6625:Phalarodon 6618:Mixosaurus 6401:see below↓ 6370:Gulosaurus 6361:Grippiidae 6310:Thaisaurus 6083:see below↓ 6062:Sauropsida 6047:Sauropsida 6020:Subclass: 6016:Sauropsida 5919:: e10299. 5512:: 618853. 4941:2023-09-30 4597:: 124–148. 4567:(9): 1–12. 4350:: 151–214. 4245:: 145–152. 3913:: e11179. 3677:References 3567:placodonts 3556:Phalarodon 3547:Phalarodon 3515:Mixosaurus 3467:macroalgae 3439:Arthropods 3431:nautiloids 3427:gastropods 3187:Mixosaurus 3043:, able to 2895:Callawayia 2568:Callawayia 2196:Callawayia 1894:being the 1519:pterygoids 1462:squamosals 1047:P. polaris 1039:Carl Wiman 1025:John Hulke 915:P. polaris 733:Mixosaurus 575:jackhammer 559:mixosaurid 520:warm stone 7525:Palvennia 7510:Keilhauia 7358:Palvennia 7177:Malawania 6330:Grippidia 6002:Kingdom: 5854:(1): 15. 5791:(1): 12. 5652:140697673 5544:231713997 5536:2296-6463 5413:211089125 5405:1664-2384 5349:222285117 4542:129092001 4448:(1). 31. 4386:(1). 32. 4305:129713553 3999:129184174 3500:hybodonts 3475:bony fish 3459:echinoids 3443:ostracods 3435:ammonoids 3090:ankylosed 3088:exhibits 1942:monotypic 1909:homoplasy 1727:phalanges 1694:clavicles 1674:coracoids 1617:gastralia 1531:dentaries 1527:lower jaw 1515:braincase 1473:quadrates 1436:lacrimals 1416:parietals 1153:In 1916, 614:of a new 587:radiogram 555:volunteer 539:oil shale 525:) on the 511:In 1985, 419:vertebrae 373:Norwegian 294:(meaning 231:Dal Sasso 161:Kingdom: 155:Eukaryota 44:, latest 7856:Q6845573 7850:Wikidata 7774:BioLib: 7760:Wikidata 7547:Sumpalla 6671:Wimanius 6496:Tholodus 6022:Diapsida 6010:Chordata 6008:Phylum: 6004:Animalia 5945:33240633 5888:37601161 5879:10432342 5825:35844249 5766:83707481 5740:Cyamodus 5723:86049393 5605:37443368 5596:10345187 5381:(1): 2. 5301:37072698 5292:10114408 5235:32822565 5197:iScience 5175:30836867 5064:21536898 4935:90912678 4880:85621052 4791:Wimanius 4770:32144303 4482:39229570 4473:11366730 4420:39263671 4411:11384637 4105:: 1–159. 4063:85352593 3939:33996277 3660:See also 3637:Clarazia 3577:Cyamodus 3561:Various 3521:(bottom) 3422:Daonella 3402:conifers 3382:plankton 3212:dolphins 3208:amniotes 3135:gharials 3027:holotype 1934:outgroup 1810:(bottom) 1666:scapulae 1664:-shaped 1638:chevrons 1550:coronoid 1507:stapedes 1412:frontals 1400:maxillae 1308:flippers 1111:being a 1063:to be a 1006:Ladinian 1002:Svalbard 986:Wimanius 954:CT scans 919:Wimanius 844:Wimanius 808:gracilis 800:kephalos 680:rhynchos 648:Lombardy 612:holotype 543:dolomite 535:faulting 529:side of 502:holotype 423:scapulae 395:flippers 380:Svalbard 349:holotype 264:Synonyms 204:Family: 185:Reptilia 175:Chordata 171:Phylum: 165:Animalia 151:Domain: 130:holotype 7896:4128739 7883:1318811 7870:4818015 7816:4130217 7803:1138622 7790:4818063 7766:Q829430 7595:Leninia 6377:Grippia 6014:Class: 5936:7682440 5856:Bibcode 5816:9276568 5793:Bibcode 5677:: 1–86. 5632:Bibcode 5573:Bibcode 5514:Bibcode 5487:: 1–13. 5383:Bibcode 5226:7520894 5205:Bibcode 5166:6458325 5055:3100925 5032:Bibcode 4986:Bibcode 4860:Bibcode 4761:7060314 4740:Bibcode 4689:Bibcode 4620:: 1–68. 4522:Bibcode 4450:Bibcode 4388:Bibcode 4285:Bibcode 4208:Bibcode 4043:Bibcode 3979:Bibcode 3930:8106916 3807:: 3–23. 3473:. Many 3425:. Many 3418:bivalve 3351:Anisian 3248:embryos 3228:inertia 3152:coleoid 3066:diapsid 1848:Grippia 1800:(top), 1754:fibulae 1719:carpals 1686:foramen 1306:, with 1173:in the 1169:of the 1091:humerus 1021:polaris 1010:Carnian 911:Chinese 812:rostrum 673:slender 632:reptile 620:species 527:Italian 477:Anisian 454:coleoid 384:Germany 353:prepare 309:extinct 307:) is a 217:Genus: 191:Order: 181:Class: 48:Anisian 7909:170846 7829:170856 7777:418411 5943: 5933: 5886: 5876: 5823: 5813: 5764: 5721: 5650: 5603: 5593: 5542: 5534: 5411: 5403: 5347: 5299: 5289: 5275:(12). 5252:: 116. 5233: 5223: 5173: 5163: 5085: 5062: 5052: 4933: 4878: 4768: 4758: 4540: 4480: 4470: 4418: 4408: 4303: 4151: 4061: 3997: 3937: 3927: 3717: 3640:, and 3451:shrimp 3449:, and 3378:oxygen 3370:lagoon 3006: 1945:family 1816:While 1750:tibiae 1746:femora 1736:, the 1725:, and 1692:. The 1662:sickle 1605:centra 1505:. The 1457:jugals 1438:. 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