Lotosaurus Temporal range: Middle Triassic (Ladinian), | |
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A mounted skeleton of Lotosaurus at the Beijing Museum of Natural History. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauria |
Clade: | Pseudosuchia |
Clade: | Paracrocodylomorpha |
Superfamily: | † Poposauroidea |
Family: | † Lotosauridae Zhang, 1975 |
Genus: | † Lotosaurus Zhang, 1975 |
Type species | |
†Lotosaurus adentus Zhang, 1975 |
Lotosaurus is an extinct genus of sail-backed poposauroid known from Hunan Province of central China. [1]
Lotosaurus is known from the holotype IVPP V 4881 (or possibly V 4880), an articulated and well-preserved skeleton. Other referred specimens include IVPP V 48013 (a skull) as well as many articulated and disarticulated skeletal remains concentrated in a bonebed which is almost completely composed of Lotosaurus bones. All known specimens of this genus were collected from this bonebed, known as the Lotosaurus site, which belongs to the Batung Formation (or alternatively Xinlingzhen Formation of the Badong Group [2] ).
Further excavations in 2018 revealed many more specimens, as well as geological and environmental details of the Lotosaurus site. At least 38 individuals of various ages died within this one location. [3]
Lotosaurus was first named by Fa-kui Zhang in 1975 and the type species is Lotosaurus adentus. Lotosaurus was originally placed in its own family, Lotosauridae, which named by Zhang in 1975. The specific name is derived from the Greek a and denta, meaning "toothless", in reference to its toothless beak. [4]
Lotosaurus was 1.5 to 2.5 m (4.9 to 8.2 ft) long and a heavily built quadruped. It was a herbivore, shearing off leaves with its toothless, beaked jaws. Lotosaurus, like some other members of the Poposauroidea, had a sail on its back, granting it an appearance superficially similar to that of Permian pelycosaurs like Dimetrodon and Edaphosaurus , although not as high. [5]
Lotosaurus was originally thought to be a thecodont, probably related to Ctenosauriscus and other archosaur taxa with elongated neural spines (=Ctenosauriscidae). [4] Although modern cladistic analyses have abandoned the order Thecodontia, they have also supported the theory that Lotosaurus was a distant relative of ctenosauriscids. Nesbitt (2007) was the first to suggest that Lotosaurus is more closely related to Shuvosaurus (a shuvosaurid) than to Arizonasaurus (a ctenosauriscid). [6] In his massive revision of archosaurs which included a large cladistic analysis, Sterling J. Nesbitt (2011) found Lotosaurus to be a member of the group Poposauroidea. Poposauroids were part of Pseudosuchia, the lineage of reptiles which also includes aetosaurs, raisuchids, and modern crocodilians. [1]
Within Poposauroidea, Lotosaurus was placed as a relative of the Shuvosauridae, and therefore it is not included in the Ctenosauriscidae. Although shuvosaurids walked on two legs and lacked sails, their toothless beaks and presumed herbivorous habits resembled those of Lotosaurus. Ctenosauriscids were carnivores with many sharp teeth and high, theropod-like skulls, in contrast to Lotosaurus and shuvosaurids. Nevertheless, Lotosaurus is still considered a distant relative of ctenosauriscids, as that family is also placed within Poposauroidea. [1] Further studies confirmed these results. [2]
The Lotosaurus site was once believed to have been as old as the Anisian stage of the early Middle Triassic, about 245-237 million years ago. [1] However, a 2018 analysis suggested a younger date based on Uranium-Lead dating from 189 sampled Zircon grains. The youngest grain sampled gave a date of 225.6 mya, in the early Norian stage of the Late Triassic. However, this grain is a lone outlier in the sample, indicating that it may have leaked lead. Using the next youngest grains, which were clustered in a set and probably more reliable, the weighted mean age of the bonebed was found to be either 238.9 to 229.3 mya (considering the lower intercept of the U-Pb graph), or 244.3 to 228.3 mya (considering the mean age of the youngest cluster of grains). Of these two options, the former has less deviation between ages and may be more accurate. Regardless, the age of the bonebed has only a small chance of belonging to the Anisian stage, and the Lotosaurus specimens which died within the bonebed were more likely to have been alive during the Ladinian or Carnian stages of the Late Middle Triassic or early Late Triassic. [3] A subsequent 2019 study weighted mean age of 238.0 ± 1.4 Ma (Ladinian) as the maximum depositional age for the sandstones preserving fossils of Lotosaurus. [7]
The Lotosaurus site was once near the northern shore of the South China Craton, a subcontinent which drifted through Tethys ocean during the Middle Triassic. The South China Craton collided with the North China Craton and the rest of Laurasia at approximately the same time as some of the estimated ages of the site. As a result, it cannot be determined if Lotosaurus was endemic to the SCC while the subcontinent remained an island, or immigrated to the SCC from the NCC after the two merged. Considering that close relatives of Lotosaurus (the shuvosaurids) lived in other regions of Pangaea around that time, the latter hypothesis is currently believed to be more likely. The mineral and microfossil composition of the site suggests that it was a subtropical environment, with pronounced wet and dry seasons. The bonebed in particular would have been part of a floodplain or stream according to ripple marks within the rocks.
The large number of individuals within a small area indicate that all of the animals died at around the same time. However, their bones are well preserved and minimally scattered, indicating that the cause of death was not a catastrophic event such as a flood or mudslide. It has been suggested that the group perished due to thirst or sickness during the dry season, perhaps while gathered around an evaporating or contaminated watering hole. Their rotting carcasses would have soon afterwards been covered in water and sediment upon the arrival of the wet season, which would explain the apparent but slight scattering of bones. [3]
"Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilians than to birds and other dinosaurs. First named in the 1940s, Rauisuchia was a name exclusive to Triassic archosaurs which were generally large, carnivorous, and quadrupedal with a pillar-erect hip posture, though exceptions exist for all of these traits. Rauisuchians, as a traditional taxonomic group, were considered distinct from other Triassic archosaur groups such as early dinosaurs, phytosaurs, aetosaurs, and crocodylomorphs.
Effigia was an extinct genus of shuvosaurid known from the Late Triassic of New Mexico, south-western USA. With a bipedal stance, long neck, and a toothless beaked skull, Effigia and other shuvosaurids bore a resemblance to the ornithomimid dinosaurs of the Cretaceous Period. However, shuvosaurids were not dinosaurs, but were instead a specialized family of poposauroid pseudosuchians, meaning that their closest living relatives are crocodilians.
Shuvosaurus is a genus of beaked reptile from the Late Triassic of western Texas. Despite looking superficially similar to a theropod dinosaur, it is actually more closely related to crocodilians.The vertebrae of Gojirasaurus now are now known to belong to Shuvosaurus.
Arizonasaurus was a ctenosauriscid archosaur from the Middle Triassic. Arizonasaurus is found in the Middle Triassic Moenkopi Formation of northern Arizona. A fairly complete skeleton was found in 2002 by Sterling Nesbitt. The taxon has a large sailback formed by elongated neural spines of the vertebrae. The type species, Arizonasaurus babbitti, was named by Samuel Paul Welles in 1947.
Hypselorhachis is a genus of extinct reptile, possibly a ctenosauriscid archosaur related to Ctenosauriscus. It lived during the Triassic Period. It is currently known only from a single vertebra found from the Middle Triassic Manda Beds in Tanzania. The vertebra is preserved in reasonably good condition, as although the tall neural spine is chipped in several places it is not broken despite being quite slender, only around 20 mm thick transversely.
Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Despite Pseudosuchia meaning "false crocodiles", the name is a misnomer as true crocodilians are now defined as a subset of the group.
Rauisuchidae is a group of large predatory Triassic archosaurs. Some disagreement exists over which genera should be included in the Rauisuchidae and which should be in the related Prestosuchidae and Poposauridae, and indeed whether these should even be thought of as separate valid families. Rauisuchids occurred throughout much of the Triassic, and may have first occurred in the Early Triassic if some archosaurian taxa such as Scythosuchus and Tsylmosuchus are considered to be within the family.
Poposaurus is an extinct genus of pseudosuchian archosaur from the Late Triassic of the southwestern United States. It belongs to the clade Poposauroidea, an unusual group of Triassic pseudosuchians that includes sail-backed, beaked, and aquatic forms. Fossils have been found in Wyoming, Utah, Arizona, and Texas. Except for the skull, most parts of the skeleton are known. The type species, P. gracilis, was described and named by Maurice Goldsmith Mehl in 1915. A second species, P. langstoni, was originally the type species of the genus Lythrosuchus. Since it was first described, Poposaurus has been variously classified as a dinosaur, a phytosaur, and a "rauisuchian".
Bromsgroveia is an extinct genus of predatory ctenosauriscid archosaur from the Middle Triassic Bromsgrove Sandstone of England. Ctenosauriscids were a group of rauisuchians that was related to the ancestors of modern crocodiles and alligators.
Ctenosauriscus is an extinct genus of sail-backed poposauroid archosaur from Early Triassic deposits of Lower Saxony in northern Germany. It gives its name to the family Ctenosauriscidae, which includes other sail-backed poposauroids such as Arizonasaurus. Fossils have been found in latest Olenekian deposits around 247.5-247.2 million years old, making it one of the first known archosaurs.
Ctenosauriscidae is an extinct family of pseudosuchian archosaurs within the clade Poposauroidea. Ctenosauriscids existed in Africa, Asia, Europe and North America during the Early Triassic to the Middle Triassic period. All species had large "sails" on their backs. Ctenosauriscids are among some of the earliest archosaurs and represent the first global radiation of the group.
Sillosuchus is a genus of shuvosaurid poposauroid archosaur that lived in South America during the Late Triassic period. Shuvosaurids were an unusual family of reptiles belonging to the group Poposauroidea; although their closest modern relatives are crocodilians, they were bipedal and lightly armored, with dinosaur-like hip and skull structures. Based on skull remains from members of the family such as Effigia, they were also toothless and likely beaked herbivores.
Dongusuchus is an extinct genus of archosaur. Fossils have been found from the Donguz Formation outcropping on the banks of the Donguz River in the Orenburg Oblast of Russia. They are associated with a fossil assemblage called the Eryosuchus Fauna, named after the capitosaurid Eryosuchus, the most common organism found from the assemblage. The locality dates back to the Anisian and early Ladinian stages of the Middle Triassic.
Vytshegdosuchus is an extinct genus of paracrocodylomorph archosaur known from the Early Triassic Yarenskian Gorizont of the Komi Republic of the European section of Russia. It contains a single species, Vytshegdosuchus zheshartensis. Vytshegdosuchus was named by Andreii Sennikov in 1988.
Poposauroidea is a clade of advanced pseudosuchians. It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as Qianosuchus and Lotosaurus. It excludes most large predatory quadrupedal "rauisuchians" such as rauisuchids and "prestosuchids". Those reptiles are now allied with crocodylomorphs in a clade known as Loricata, which is the sister taxon to the poposauroids in the clade Paracrocodylomorpha. Although it was first formally defined in 2007, the name "Poposauroidea" has been used for many years. The group has been referred to as Poposauridae by some authors, although this name is often used more narrowly to refer to the family that includes Poposaurus and its close relatives.
Paracrocodylomorpha is a clade of pseudosuchian archosaurs. The clade includes the diverse and unusual group Poposauroidea as well as the generally carnivorous and quadrupedal members of Loricata, including modern crocodylians. Paracrocodylomorpha was named by paleontologist J. Michael Parrish in 1993, although the group is now considered to encompass more reptiles than his original definition intended. The most recent definition of Paracrocodylomorpha, as defined by Sterling Nesbitt in 2011, is "the least inclusive clade containing Poposaurus and Crocodylus niloticus. Most groups of paracrocodylomorphs became extinct at the end of the Triassic period, with the exception of the crocodylomorphs, from which crocodylians such as crocodiles and alligators evolved in the latter part of the Mesozoic.
Xilousuchus is an extinct genus of poposauroid from lower Triassic deposits of Fugu County of northeastern Shanxi Province, China. It is known from the holotype, IVPP V 6026, a single well-preserved partial skeleton including the skull. It was found from the Heshanggou Formation of the Ordos Basin, Hazhen commune. It was first named by Xiao-Chun Wu in 1981 and the type species is Xilousuchus sapingensis. Wu (1981) referred Xilousuchus to the Proterosuchia. Gower and Sennikov (1996) found it to be an erythrosuchian based strictly on the braincase. A more detailed re-description of the genus was provided by Nesbitt et al. (2010) and found poposauroid affinities. In his massive revision of archosaurs which included a large cladistic analysis, Sterling J. Nesbitt (2011) found Xilousuchus to be a poposauroid which is most closely related to Arizonasaurus. Xilousuchus is the oldest archosaur to date, although Ctenosauriscus and Vytshegdosuchus might be even older by less than one million year. Since Xilousuchus is a suchian archosaur, its early age suggests that most of the major groups of archosaurs developed by the Early Triassic, soon after the appearance of the first archosaur.
Diandongosuchus is an extinct genus of archosauriform reptile, possibly a member of the Phytosauria, known from the Middle Triassic of China. The type species Diandongosuchus fuyuanensis was named in 2012 from the Zhuganpo Formation of Yunnan Province. It is a marine species that shows similarities with another Chinese Triassic species called Qianosuchus mixtus, although it has fewer adaptations toward marine life. It was originally classified as the basal-most member of the pseudosuchian clade Poposauroidea. However, a subsequent study conducted by Stocker et al. indicated it to be the basalmost known phytosaur instead.
Wangisuchus is an extinct genus of archosauriform reptile from the Middle Triassic of China that is known from fragmentary fossil jaw bones. These bones were found at the Hsishihwa locality in the upper Ermaying Formation, which dates to the late Anisian stage about 242 million years ago. Wangisuchus was named in 1964 by Chinese paleontologist Yang Zhongjian, who described a single species, Wangisuchus tzeyii, on the basis of these bones.
Mambawakale is a genus of large sized basal paracrocodylomorph, possibly a poposauroid, from the Manda Beds of Tanzania. It was informally named Pallisteria before being officially published under its current name almost 60 years after its discovery. It contains a single species, Mambawakale ruhuhu.