Kosmodraco Temporal range: | |
---|---|
Holotype skull of K. magnicornis | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Order: | † Choristodera |
Suborder: | † Neochoristodera |
Genus: | † Kosmodraco Brownstein, 2022 |
Type species | |
Kosmodraco dakotensis (Erickson, 1987) [1] | |
Other species | |
| |
Synonyms | |
|
Kosmodraco is a genus of large bodied choristodere from the Paleocene of North America. Originally described as species of the closely related 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.
The first specimen now known to belong to Kosmodraco was discovered in 1964 and 1968 in the Polecat Bench Formation (Wyoming). These two skulls, alongside others from Montana's Bear Creek, were reported on briefly by Sigogneau-Russell and Donald in 1978, who regarded them as evidence for the presence of the Eurasian genus Simoedosaurus in North America. [2] However, at the time, these four specimens, although thought to be diagnostic at a genus level, were still unprepared and not assigned to a species. They were subsequently stored in the collection of the Princeton University. It wasn't until a fifth specimen from North Dakota was prepared that the North American remains were named, with Bruce R. Erickson creating the new species Simoedosaurus dakotensis in 1987 and assigning the Princeton material and two additional specimens to it. Erickson, however, did realize there were differences in the specimens, noting that the North Dakota specimen was not just older (mid Tiffanian) than the Princeton fossils (late Tiffanian to Clarkforkian) but also larger, which he initially attributed to differences between growth stages. Later, in 2022, Chase Doran Brownstein reexamined the American Simoedosaurus material, finding that it differed enough to be described as its own genus. Brownstein further considered that the two best preserved Princeton specimens, YPM VPPU 19168 and YPM VPPU 18724, weren't juveniles but should instead be considered a second species. Subsequently, the genus Kosmodraco was erected with K. dakotensis as the type species and a second species in the form of K. magnicornis. [3]
The name Kosmodraco translates to "ornamented dragon" from the Greek κοσμος and Latin draco. The type species was named after the state of North Dakota, while K. magnicornis was named for its large squamosal spikes, derived from the Latin magnum (large) and cornum (horn). [3]
Unlike the well known Champsosaurus, Kosmodraco had an exceptionally brevirostrine skull with the blunt muzzle only making up approximately one third of the total skull length. The posterior region, mostly made up of the squamosal bone, is exaggeratedly broad, giving the skull an overall triangular form. Additionally, all of the skulls show that the cranium of Kosmodraco was exceptionally flat and covered by ridges and furrows along the dorsal and lateral surface. To add to this, four transverse bumps are present on the prefrontals before and between the eyes. The external nares make up around half the width of the bulbous premaxilla and are partly divided by the fused nasal bone. The nasal further extends into the nares through an internasal bar. The premaxilla is only slightly longer than wide and followed by a marked constriction where they meet the maxillae. Four premaxillary teeth of somewhat similar size are followed by forty-five maxillary teeth. In the holotype specimen, the anterior areas of the maxilla show an alternating patter between erupted teeth and empty alveoli, with the second maxilla showing the opposite distribution. Following the 16th tooth, this alternating pattern stops in favor of more continuous tooth rows. Tooth rows are additionally present on the vomers and palatines with small and blunt dentition. [3] The paired prefrontals are anteriorly bifurcated by the nasal and reach to almost the exact halfway point between the orbits where they meet the rectangular frontal bones. The orbits are subangular and directed dorsolaterally, in spite of the flattening of the skull. The shape of the orbits is owed to the fact that the skull roof overhangs large portions of them. Almost the entire medial border of the supratemporal fenestrae consists of the parietal bones, which display a long suture along the center of the skull. The central portion where the two parietals meet is located in a shallow through while the posterior elements of the bones are raised as a crest before meeting the squamosals. The squamosal bones themselves display prominent nodes towards the rear of the skull. The infratemporal and supratemporal fenestrae are similar in size and the maximum distance between the squamosals is only slightly larger than the width of the skull table. [1]
Kosmodraco magnicornis differs from the type species in several key traits. The margin of the rostrum is smooth and confluent unlike the constricted rostrum seen in K. dakotensis. The first three alveoli of the premaxilla are twice as large as any following (except for two where the maxilla is relatively inflated), which differs from the more consistent decrease in size observed in the teeth of the type species. At around the end of the anterior third of the maxilla, the bone is ventrally inflated by two enlarged maxillary alveoli in a condition comparable to the area right behind the premaxillary-maxillary suture in the type species, however greatly exaggerated in a manner more akin to modern crocodylians. This species has fewer teeth than K. dakotensis, with only thirty-one maxillary teeth opposed to forty-five. The orbits are more subcircular and possibly possessed higher raised orbital margins, however due to the crushing the holotype of K. dakotensis this cannot be said for certain. Much like the holotype, Kosmodraco magnicornis bears eight nodules along the posterior edge of the squamosal. However, unlike in the holotype, these nodules are much more developed, exceeding any other late Mesozoic or Cenozoic choristodere and much more resembles those of Coeruleodraco and Monjurosuchus . These ornaments are what give the species its name. [3]
Postcranial material is known from both species. Kosmodraco dakotensis is associated with eight cervical vertebrae and at least sixteen additional centra thought to be presacrals. Three sacral vertebrae were described (however one of them was lost) and multiple caudals from various sections of the tail. Among the cervicals, the axis has a stout neural arch and large crest that projected over the centrum of the atlas, likely as an anchor for the musculature of the head. This differs significantly from the low and broad neural spine observed in Champsosaurus gigas. Thirty-four complete vertebrae are known from K. magnicornis and show very similar morphology to the type species. [3] The shoulder girdle is known from Kosmodraco magnicornis with a long blade of the scapula and a longer coracoid blade. Of the pubic bones, the ischium and pubis are only well enough preserved for a very general reconstruction. The ilium is known from more complete material. The acetabulum has a rounded rather than pointed bottom and rises almost vertically towards the iliac blade unlike in Simoedosaurus. Although several limb bones are known from fragments, only a single femur is complete. This femur is longer and more slender than fossils known from Europe and the condyles are relatively weak, with narrow articular surfaces. [1]
Kosmodraco was perhaps among the largest amphibious predatory animals of its time and range with the type species having an estimated body length of 3–4 m (9.8–13.1 ft), [1] perhaps even 5 m (16 ft), [4] rivaling even most crocodylomorphs of the Paleocene. K. magnicornis was on the smaller side of the spectrum with a skull length of 431 mm (17.0 in), which still leaves it as the fourth largest known choristodere behind K. dakotensis, Champsosaurus gigas and Simoedosaurus . [3]
Phylogenetic analysis consistently recovered Kosmodraco as clading together with the Old World choristodere Simoedosaurus and Tchoiria , forming the family Simoedosauridae within Neochoristodera. Within the phylogenetic analysis conducted by Brownstein, differences within Simoedosauridae are restricted to the relationships between the two basalmost taxa, both species of Tchoiria. However, their placement as basal simoedosaurids is consistent amongst all of them. Depicted below are the results of the Bayesian phylogenetic analysis. [3]
| |||||||||||||||||||||||||||||||||||||
Although niche partitioning is a common explanation for predator-rich ecosystems and would fit the distinctive skull shapes of both Kosmodraco and the contemporary Champsosaurus, Brownstein argues that in the case of Kosmodraco the vastly different skull shapes have an evolutionary rather than strictly ecological origin. The brevirostrine skull shape of the animal has repeatedly been compared to both alligatoroids and gars of the genus Atractosteus (including the alligator gar), with all of them having evolved relatively short but robust skulls. [1] Kosmodraco additionally shows that both the orbits and nares were raised, an adaption to a semi-aquatic lifestyle commonly seen in crocodilians and absent in gars. In the reverse case, gars and Kosmodraco share the presence of palatal teeth which are not found in alligators. The overall skull shape is also closer to that of gars with a more triangular shape, while the skulls of alligatoroids are typically broader and more subrectangular. Kosmodraco however differs from both with its extremely flattened skull and characteristically expanded postorbital region. As far as the postcrania are concerned, Kosmodraco possessed three sacral vertebrae bearing distinctive prominences for ligaments and muscles to attach to. These attachments for sacral and caudal soft tissue differ from what is observed in crocodilians today and make it subsequently difficult to infer the precise ecological niche these animals would have inhabited. [3]
Given the differing skull shapes, it is likely that the contemporary Champsosaurus and Kosmodraco, despite being similar in size, preyed on different animals; the former filling a niche possibly similar to extant gharials, while the latter, with its more robust and broader skull, had a generalized diet of fish and small vertebrates. More similar to Kosmodraco were the various brevirostrine crocodilians found in Paleocene North America. In the Fort Union Formation, for example, Champsosaurus gigas and a species of Kosmodraco coexisted with possibly as many as four different crocodilians. Most of these species, however, like Ceratosuchus and Allognathosuchus, were notably smaller than either choristodere. The exception is Borealosuchus , which with a length of up to 4 meters rivaled Kosmodraco in size. [4] Given Kosmodraco's size and robust morphology, Erickson considers it possible that it was capable of competing with crocodilians. This could explain injuries found on the bones of Kosmodraco as described by Erickson, although they might have been caused by intraspecific combat. [1]
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.
Mekosuchus is a genus of extinct Australasian mekosuchine crocodilian. Species of Mekosuchus were generally small-sized, terrestrial animals with short, blunt-snouted heads and strong limbs. Four species are currently recognized, M. inexpectatus, M. whitehunterensis, M. sanderi and M. kalpokasi, all known primarily from fragmentary remains.
Mahajangasuchus is an extinct genus of crocodyliform which had blunt, conical teeth. The type species, M. insignis, lived during the Late Cretaceous; its fossils have been found in the Maevarano Formation in northern Madagascar. It was a fairly large predator, measuring up to 4 metres (13 ft) long.
Arambourgisuchus is an extinct genus of dyrosaurid crocodylomorph from the late Palaeocene of Morocco, found in the region of Sidi Chenane in 2000, following collaboration by French and Moroccan institutions, and described in 2005 by a team led by palaeontologist Stéphane Jouve. Arambourgisuchus was a large animal with an elongated skull 1 meter in length.
Lazarussuchus 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 in France, Germany, and the Czech Republic. Two species have been named: the type species L. inexpectatus ("unexpected") from the late Oligocene of France. and L. dvoraki from the early Miocene of the Czech Republic. It was not a large animal; with the total preserved body and tail length of L. inexpectatus being just over 30 centimetres. 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.
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.
Anthracosuchus is an extinct genus of dyrosaurid crocodyliform from the Paleocene of Colombia. Remains of Anthracosuchus balrogus, the only known species, come from the Cerrejón Formation in the Cerrejón mine, and include four fossil specimens with partial skulls. Anthracosuchus differs from other dyrosaurids in having an extremely short (brevirostrine) snout, widely spaced eye sockets with bony protuberances around them, and osteoderms that are smooth and thick. It is one of the most basal dyrosaurids along with Chenanisuchus and Cerrejonisuchus.
Knoetschkesuchus is a genus of small atoposaurid crocodylomorph from the Late Jurassic of Germany and Portugal. Two species are known: the German species K. langenbergensis, described by Schwarz and colleagues in 2017 based on two partial skeletons and various isolated bones; and the Portuguese species K. guimarotae, named from over 400 specimens including several partial skeletons. Knoetschkesuchus was a small and short-snouted crocodilian, measuring about 55 centimetres (22 in) in length, that primarily fed on small prey, including invertebrates, amphibians, and mammals. This specialization towards small prey ecologically separated Knoetschkesuchus from most of the other diverse crocodilians that it lived with in the island ecosystem of Jurassic Europe.
Kadimakara is an extinct genus of early archosauromorph reptile from the Arcadia Formation of Queensland, Australia. It was seemingly a very close relative of Prolacerta, a carnivorous reptile which possessed a moderately long neck. The generic name Kadimakara references prehistoric creatures from Aboriginal myths which may have been inspired by ice-age megafauna. The specific name K. australiensis relates to the fact that it was found in Australia. Prolacerta and Kadimakara were closely related to the Archosauriformes, a successful group which includes archosaurs such as crocodilians, pterosaurs, and dinosaurs.
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.
Rugarhynchos is an extinct genus of doswelliid archosauriform from the Late Triassic of New Mexico. The only known species is Rugarhynchos sixmilensis. It was originally described as a species of Doswellia in 2012, before receiving its own genus in 2020. Rugarhynchos was a close relative of Doswellia and shared several features with it, such as the absence of an infratemporal fenestra and heavily textured skull bones. However, it could also be distinguished by many unique characteristics, such as a thick diagonal ridge on the side of the snout, blunt spikes on its osteoderms, and a complex suture between the quadratojugal, squamosal, and jugal. Non-metric multidimensional scaling and tooth morphology suggest that Rugarhynchos had a general skull anatomy convergent with some crocodyliforms, spinosaurids, and phytosaurs. However, its snout was somewhat less elongated than those other reptiles.
Alligator hailensis, or Haile alligator, is a large, extinct species of Alligator known from the early Pleistocene of Florida. It is named after the town of Haile, Florida, where it was found. Its age and skeletal morphology is intermediate between the geologically older Alligator mefferdi and the modern American alligator, making it a transitional fossil.
Chinatichampsus is an extinct genus of crocodilian from the Devil's Graveyard Formation of Texas, specifically the Dalquest Desert Research Site. It is a monotypic genus, containing only the type species Chintanichampsus wilsonorum. A single specimen, TMM 45911–1, was first discovered in 2010. Chinatichampsus is the most basal Eocene caimanine, dating to between 42.8 and 41.5 million years ago, and is considered to be more basal than Protocaiman.
Ultrastenos is an extinct genus of Australian mekosuchine crocodilian that lived during the Late Oligocene in northwestern Queensland, Australia. Following its discovery, it was speculated that Ultrastenos was a slender-snouted animal similar to modern gharials or freshwater crocodiles due to the seemingly abruptly narrowing mandible. However, a later study found that this was a missinterpretation of the fossil specimen and that Ultrastenos instead had a more generalized lower jaw. The same publication also provided evidence that the fossils of Ultrastenos belonged to the same animal previously named "Baru" huberi, adding further evidence to the idea that the animal was short snouted, contrary to the initial hypothesis. Given that "Baru" huberi was named first, the type species of Ultrastenos changed from U. willisi to U. huberi in accordance with the rules of the ICZN. Ultrastenos was a small mekosuchine, measuring upwards of 1.5 m long.
Qianshanosuchus is a genus of basal crocodyloid from the Paleocene of the Qianshan Basin, China. The fossil material, which includes an incomplete skull and parts of the lower jaw, show various features usually associated with juvenile crocodiles alongside various unique traits that were used to erect a new genus. It is the first and only basal crocodyloid currently known from the Paleocene of China, which had previously only yielded alligatoroids and planocraniids. Its presence in this part of the world and its basal position to species of the genus Asiatosuchus supports the idea that crocodyloids dispersed from Asia into Europe. Qianshanosuchus only includes a single species, Qianshanosuchus youngi.
Eurycephalosuchus is an extinct genus of orientalosuchine alligatoroid from the Late Cretaceous Jiangxi Province of China. Known from a well preserved skull and mandible alongside various postcranial remains, Eurycephalosuchus possessed a short and broad skull with a very short skulltable. Eurycephalosuchus lived with at least one other crocodilian, an indetermined member of the clade Brevirostres. The genus is monotypic, containing only the species Eurycephalosuchus gannanensis.
Varanosuchus is an extinct genus of atoposaurid neosuchian from the Early Cretaceous Sao Khua Formation of Thailand. Varanosuchus is known from three individuals which preserve assorted postcranial material and a complete skull. The skull of Varanosuchus was altirostral, meaning it wasn't flattened like in modern crocodilians and instead much deeper, while the limbs were slender and straight, leading to it somewhat resembling a monitor lizard. Little is known about the ecology of atoposaurids, however, based on the slender, erect limbs, the altirostral skull and the well ornamented osteoderms it has been suggested that Varanosuchus was a terrestrial animal with some semi-aquatic affinities. Only a single species is recognized, Varanosuchus sakonnakhonensis.