Lazarussuchus | |
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Reconstruction of Lazarussuchus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Order: | † Choristodera |
Genus: | † Lazarussuchus Hecht, 1992 |
Type species | |
†Lazarussuchus inexpectatus Hecht, 1992 | |
Other species | |
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Lazarussuchus (meaning "Lazarus's crocodile") is an extinct genus of amphibious reptile, known from the Cenozoic of Europe. It is the youngest known member of Choristodera, an extinct order of aquatic reptiles that first appeared in the Middle Jurassic. Fossils have been found in Late Paleocene, Late Oligocene, Early Miocene and possibly Late Miocene deposits (~56-20 or possibly 11.6 million years ago) in France, Germany, and the Czech Republic. Two species have been named: the type species L. inexpectatus ("unexpected") (Hecht, 1992) from the late Oligocene of France. [1] and L. dvoraki from the early Miocene of the Czech Republic. [2] It was not a large animal; with the total preserved body and tail length of L. inexpectatus being just over 30 centimetres. [1] A complete specimen of Lazarussuchus with preserved soft tissue was found from the Late Paleocene of France, but has not been assigned to a species. [3]
The first remains of Lazarussuchus, belonging to the type species L. inexpectatus were described in 1992 from a mostly complete articulated skeleton (Claude Bernard University no Re 437, coll. Gennevaux 92813) found in the Upper Oligocene aged sediments of the Armissan limestone quarry near Narbonne in Aude, France. The genus was named after the "Lazarus Effect", as the remains were the youngest known choristodere fossils, and resembled the most primitive choristodere known, the Middle-Late Jurassic Cteniogenys . [1] In 2005 another species, L. dvoraki, was described from isolated skull bones and vertebrae from the Early-Middle Miocene sediments of the Merkur-North locality in the north-west Czech Republic. The species was named after Zdeněk Dvořák who had collected the specimens. [2] In 2008, remains of Lazarussuchus were reported from the Upper Oligocene sediments of Oberleichtersbach in northern Bavaria, Germany. Remains included 25 bones, and was suggested to probably represent a new species. [4] In 2013, a specimen of Lazurussuchus was described from the late Paleocene aged Menat Formation near Menat, Puy-de-Dôme in France. The specimen, which is an almost complete articulated skeleton (BDL 1819) is largely preserved as an impression, with remnants of disintegrating bone and some preserved soft tissue. The remains were not assigned to a species however, because it could not be robustly diagnosed separately from the two named species. [3] In 2019, in the supplementary information for the paper describing the remains of the extinct ape Danuvius, indeterminate remains of Lazarussuchus were reported from the Hammerschmiede clay pit near Pforzen, Bavaria, Germany. The Hammerschmiede locality has been dated magnetostratigraphically to the base of the Tortonian stage of the Late Miocene, approximately 11.62 million years ago. [5]
Lazarussuchus was small and superficially lizard like in appearance, with the total preserved body and tail length of L. inexpectatus being just over 30 centimetres (0.98 ft), and a skull length of around 4.5 centimetres (1.8 in) [1] According to Matsumoto and colleagues (2013) Lazarussuchus is distinguished from all other known choristoderes by the presence of paired nares high on the rostrum, elongated premaxillary bones which replace the maxillae in the anterior part of the rostrum, the slender paired nasal bones are overlapped anteriorly by the premaxillae and wedged between the anterior tips of the prefrontal bones, the nasal bones do not touch the maxillae, the bones on the hind portion of the skull are ornamented with a coarse tuberculous texture, the postorbital and postfrontal bones are fused into a combined postorbitofrontal, the lower temporal fenestrae are closed by the joining of the squamosal, quadratojugal, jugal and postorbitofrontal bones, the cervical ribs are broad and flattened, the trunk ribs are broad and sickle shaped, and are large relative to the size of the vertebrae, the ectepicondylar groove of the humerus is bridged to form a foramen, and the blade of the scapula is slender and parallel sided. [3]
L. inexpectatus is distinguished by having the postparietal processes of the parietal bone equal or nearly equal in length to the parietal plate, directed posteriorly with less than 30 degrees of angulation and only weakly concave, resulting in the upper temporal fenestra being elongated and almost rectangular in shape, with an anterior-posterior long axis, nine cervical vertebrae are present, the posterior portion of the trunk vertebrae have spinous processes below the postzygapophyses that act as additional articular surfaces, four functional sacral vertebrae are present, of which the first is a sacrodorsal, and a slender T-shaped interclavicle. L. dvoraki is distinguished by having postparietal processes only 1/3 of the length of the parietal plate, which are angulated greater than 45 degrees laterally and a concave lateral margin, resulting in the upper temporal fenestrae being smaller and more ovoid in comparison to L. inexpectatus, with a antero-medial to posterolateral long axis, and the trunk vertebrae lack articular spinal processes. [3]
The Menat specimen is distinguished by having 40 maxillary teeth and a total of 52 teeth in the upper jaw as opposed to 11 premaxillary and 24 maxillary tooth positions in L. inexpectatus, however, it cannot be ruled out that these are due to allometry. The Menat specimen is interpreted to have 10 cervical vertebrae, as opposed to the nine reported for L. inexpectatus, however the 10th cervical vertebra of the Menat specimen is similar to the 9th of L. inexpectatus, and there is a gap between the 8th and 9th vertebrae of the L. inexpectatus specimen suggesting a vertebra could be missing or obscured. The ribs tuberculum and capitulum are joined by a crest in L. inexpectatus, but the crest on the ribs of the 9th vertebra is limited. In the Menat specimen, the crest is present on all cervical ribs, including a particularly strong crest on the rib of the ninth vertebra. However, this contrast may be due to missalignment of vertebral counts as mentioned previously. The interclavicle of L. inexpectatus was described as T-shaped in its initial description, however the main part of the bone is not visible, and only a strong lateral process is visible. In contrast, the Menat specimen interclavicle is rhomboid in shape, but the lateral processes are incomplete, making comparison difficult. The type specimen of L. inexpectatus has plantar tubercles present on the proximal portion of the fifth metatarsal, while the Menat specimen does not. However, this difference can only be properly evaluated in 3 dimensional specimens. Matsumoto and colleagues (2013) declined to create a new species for the Menat specimen despite it being "most certainly specifically distinct" because "species diagnoses must be based on clear morphological differences and this is problematic for the Menat specimen" going on to state that "Size and preservation could .... explain many or all of the observed differences between the [L. inexpectatus type and Menat] specimens". [3]
The premaxilla of the Oberleichtersbach Lazarussuchus is shorter than that of L. inexpectatus, alongside other unspecified cranial bone differences. [4]
The Menat specimen of Lazarussuchus preserves some remnants of soft tissue, but no scales, which shows that the hindfoot (pes) was not webbed, and a dark stained region with a crenellated edge is present above the caudal vertebrae of the tail, suggestive of a crest similar to those found in some living reptiles, like the tuatara, lizards and crocodiles. [3]
Lazarussuchus is the youngest member of the group Choristodera and the only choristodere to have lived after the Eocene. The crocodile-like neochoristoderes went extinct at the start of the Eocene, possibly due to climatic changes happening at this time, known as the Paleocene-Eocene Thermal Maximum. The first discovered fossils of Lazarussuchus came from the Late Oligocene, meaning that it extended the fossil range of Choristodera by several million years. At the time of its discovery Lazarussuchus was considered an example of the Lazarus effect because its "unexpected" presence followed a long gap in the choristodere fossil record. [1]
Lazarussuchus was initially interpreted as the basalmost member of Choristodera, meaning that its lineage must have been the earliest to branch off from the group. [6] Since the first definitive choristoderes appear in the Middle Jurassic, the lineage represented by Lazarussuchus implied a very long ghost lineage of choristoderes spanning at least 100 million years. [2] However in the 2005 description of L. dvoraki, it was placed as the sister group of neochoristoderes. [2] Matsumoto and colleagues (2013) placed Cteniogenys, the oldest known choristodere, as the most basal member of the group and has Lazarussuchus in a more derived position within a clade of small-bodied choristoderes known from the Early Cretaceous of Asia, united by the shared condition of closed lower temporal fenestrae, which has been recovered by most subsequent analyses. The clade was informally named the "Allochoristodera" by Dong and colleagues (2020). [7] [8] This position significantly shortens the ghost lineage of Lazarussuchus, although there is still a gap between the disappearance of small-bodied, lizard-like choristoderes at the end of the Early Cretaceous and their reappearance in the form of Lazarussuchus in the Paleocene. Possible lizard-like choristoderes have been found in Late Cretaceous North American deposits, although their remains are very fragmentary. [3]
Phylogeny from the analysis of Dong and colleagues (2020): [8]
Choristodera |
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Although no gut contents have been found, Lazarussuchus is assumed to have fed on invertebrates. [6] At Oberleichtersbach and Merkur-North, Lazaurussuchus was found in continental lake deposits, with a subtropical climate. Occurring alongside fish, frogs, turtles, squamates and alligatoroid crocodillians belonging to the genus Diplocynodon. [9]
Archosauromorpha is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs rather than lepidosaurs. Archosauromorphs first appeared during the late Middle Permian or Late Permian, though they became much more common and diverse during the Triassic period.
Tanystropheus is an extinct genus of archosauromorph reptile which lived during the Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than the torso and tail combined. The neck was composed of 13 vertebrae strengthened by extensive cervical ribs. Tanystropheus is one of the most well-described non-archosauriform archosauromorphs, known from numerous fossils, including nearly complete skeletons. Some species within the genus may have reached a total length of 6 meters (20 ft), making Tanystropheus the longest non-archosauriform archosauromorph as well. Tanystropheus is the namesake of the family Tanystropheidae, a clade collecting many long-necked Triassic archosauromorphs previously described as "protorosaurs" or "prolacertiforms".
Champsosaurus is an extinct genus of crocodile-like choristodere reptile, known from the Late Cretaceous and early Paleogene periods of North America and Europe (Campanian–Paleocene). The name Champsosaurus is thought to come from champsai, (χαμψαι) said in an Ancient Greek source to be an Egyptian word for "crocodiles", and sauros, (σαύρος) Greek for "lizard". The morphology of Champsosaurus resembles that of gharials, with a long, elongated snout. It was native to freshwater environments where it likely preyed on fish, similar to living gharials.
Choristodera is an extinct order of semiaquatic diapsid reptiles that ranged from the Middle Jurassic, or possibly Triassic, to the Miocene. Choristoderes are morphologically diverse, with the best known members being the crocodile-like neochoristoderes such as Champsosaurus. Other choristoderans had lizard-like or long necked morphologies. Choristoderes appear to have been confined to the Northern Hemisphere, having been found in North America, Asia, and Europe, and possibly also North Africa. Choristoderes are generally thought to be derived neodiapsids that are close relatives or members of Sauria.
Simoedosaurus is an extinct reptile known from the Paleocene of North America, Europe and western Asia, and a member of the Choristodera, a group of aquatic reptiles that lived in the Northern Hemisphere from the Jurassic to the early Cenozoic.
Hyphalosaurus is a genus of freshwater aquatic reptiles, belonging to the extinct order Choristodera. They lived during the early Cretaceous period, about 123-120 million years ago. The genus contains two species, H. lingyuanensis from the Yixian Formation and H. baitaigouensis from both the Yixian and Jiufotang Formation of Liaoning Province, China. They are among the best-known animals from the Jehol Biota, with thousands of fossil specimens representing all growth stages in scientific and private collections.
Askeptosaurus is an extinct genus of askeptosauroid, a marine reptile from the extinct order Thalattosauria. Askeptosaurus is known from several well-preserved fossils found in Middle Triassic marine strata in what is now Italy and Switzerland.
Shokawa is an extinct genus of choristoderan diapsid reptile, known from the Lower Cretaceous of Japan. It is only known from one species, Shokawa ikoi. The only known remains are a postcranial specimen lacking the skull, discovered at the KO2 locality in sediments belonging to the Okurodani Formation near the village of Shokawa in Gifu Prefecture. Shokawa possessed a long neck with at least 16 cervical vertebrae, and closely resembles and is closely related to the smaller choristoderan, Hyphalosaurus. The generic name refers to the village near where it was found, while the specific name honors the collector of the first specimen, one Mr. Ikoi Shibata.
Pachystropheus is a genus of prehistoric reptile, from the Rhaetian of southwestern England. It was named by Erika von Huene in 1935; Huene described Pachystropheus as a choristodere, but this was overlooked for decades until its redescription by Storrs and Gower in 1993. This reevaluation would extend the fossil record of choristoderes back 45 million years. However, other authors consider attribution of Pachystropheus to Choristodera problematic, stating that it depends on vertebral and girdle characters that are also found in the skeletons of aquatic reptiles other than choristoderes; most of the diagnostic features of choristoderes are skull features, but the presence of these cannot be confirmed in Pachystropheus, as there is no confirmed skull material for this taxon. Silvio Renesto (2005) found similarities in the postcranial skeleton of Pachystropheus and the thalattosaur genus Endennasaurus; according to Renesto, these similarities may indicate that Pachystropheus and Endennasaurus are close relatives, but they might as well simply be a case of a convergent evolution triggered by the aquatic lifestyle of both taxa. The placement as a thalattosaur has been supported by other researchers.
Cteniogenys is a genus of choristodere, a morphologically diverse group of aquatic reptiles. It is part of the monotypic family Cteniogenidae. The type and only named species, C. antiquus, was named in 1928 by Charles W. Gilmore. The holotype, VP.001088, was collected in the Morrison Formation, Wyoming in 1881 by William H. Reed. More specimens have been discovered since then, including specimens from the Late Jurassic of Portugal and Middle Jurassic of Britain, which have not been assigned to species.
Monjurosuchus is a genus of choristoderan reptile that lived in what is now China and Japan during the Early Cretaceous. It has large eyes, a rounded skull, robust legs with short claws, and a long, thin tail. Fossils have been found that preserve soft tissue, showing that it had soft skin and webbed feet.
Khurendukhosaurus is a genus of choristodere, a type of amphibious reptile. It is known from Lower Cretaceous rocks of Mongolia and Russia. Two species have been named. The type species, K. orlovi, was named in 1984 by Sigogneau–Russell and Efimov for the fragmentary postcranial skeleton PIN 3386/3. This specimen was discovered in the Albian-age Lower Cretaceous Khuren Dukh Formation Formation at Hüren Dukh, central Mongolia. The lake deposits at this site also contain fossils of the choristoderes Irenosaurus and Tchoiria. Other postcranial bones of K. orlovi have been found at this site as well.
Tchoiria () is a genus of neochoristoderan reptile from the Early Cretaceous of Mongolia. The name Tchoiria comes from the city of Choir which is nearby to where the holotype was found. Tchoiria is thought to have a similar diet to another neochoristoderan reptile, Champsosaurus, due to morphology of the skull. It would hunt in freshwater environments, like the living gharials, where it would prey on many different types of fish and turtles.
Neochoristodera is a lineage of specialised crocodile-like fully aquatic choristodere reptiles. Noted for their long jaws and large size, these animals were predominant across the Northern Hemisphere, occurring in freshwater and coastal environments across the Cretaceous and early Cenozoic.
Philydrosaurus is an extinct genus of choristoderan which existed in China during the Early Cretaceous. The type species P. proseilus was named in 2005. Philydrosaurus was found from the Jiufotang Formation and is slightly younger than Monjurosuchus, which was found from the Yixian Formation.
Jesairosaurus is an extinct genus of early archosauromorph reptile known from the Illizi Province of Algeria. It is known from a single species, Jesairosaurus lehmani. Although a potential relative of the long-necked tanystropheids, this lightly-built reptile could instead be characterized by its relatively short neck as well as various skull features.
The Menat Formation is a geologic formation in France. It preserves fossils dating back to the Paleocene. It is a maar deposit located on top of an ancient volcano, the extent is very localised with the outcropping area being around 600 by 1000 metres. Numerous species of fossil insects, plants and fish are known, as well as some isolated mammals, including the primate Plesiadapis insignis, the choristodere reptile Lazarussuchus, and birds, including members of Halcyornithidae and Messelasturidae and relatives of Songziidae.
Coeruleodraco is an extinct genus of choristoderan known from the Late Jurassic (Oxfordian) Tiaojishan Formation in China. Coeruleodraco is significant as the most complete Jurassic choristodere taxon, as the only other named Jurassic choristodere Cteniogenys is based on fragmentary remains. Although similar to Philydrosaurus in its proportions and postcranial characters, it is distinct in retaining several apparently plesiomorphic characters, including a short snout, paired external nares and an open lower temporal fenestra.
Heishanosaurus is an extinct genus of choristodere reptile from the Early Cretaceous of China. The type and currently only known species is Heishanosaurus pygmaeus. It is unusual as it is much more primitive than other known choristoderes from the Early Cretaceous of Asia, and retains many plesiomorphic characters.
Kosmodraco is a genus of large bodied choristodere from the Paleocene of North America. Originally described as a species of Simoedosaurus, it was found to represent a distinct genus in 2022. Multiple fossil skulls show a relatively short and robust snout and a skull that is considerably wider behind the eyes. Two species are currently recognized, K. dakotensis and K. magnicornis.