Liliensternus

Liliensternus is an extinct genus of basal Neotheropod dinosaur that lived approximately 210 million years ago during the latter part of the Triassic Period in what is now Germany. Liliensternus was a moderate-sized, bipedal, ground-dwelling carnivore, that could grow up to 5.15 m (16.9 ft) long. It is the best represented Triassic theropod from Europe and one of the largest known.[1]

Liliensternus
Temporal range: Late Triassic, 228–201 Ma
Liliensternus liliensterni
Skeleton restoration
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Clade: Neotheropoda
Genus: Liliensternus
Welles, 1984
Type species
Halticosaurus liliensterni
Huene, 1934
Species

Liliensternus liliensterni
(Huene, 1934)

Synonyms

Halticosaurus liliensterni Huene, 1934

Description

Liliensternus NT
Liliensternus restored with a speculative crest.

Liliensternus was approximately 5.15 metres (16.9 feet) long, and may have weighed about 127 kilograms (280 pounds).[2] Other estimates suggest that Liliensternus was at best 5.2 m (17.1 ft) long and weighed 200 kg (441 lb) at most.[3] The remains of two specimens of Liliensternus together form a syntype series with inventory number MB.R.2175, and consist of the partial and fragmentary skeletons of at least two individuals, containing elements of the skull, the lower jaws, the vertebrae and the appendicular skeleton. The tibia (409 mm) is shorter than the femur (440 mm) in both Dilophosaurus and Liliensternus, unlike those of coelophysid taxa, such as Coelophysis. Paul (1988) noted that based on its appearance, Liliensternus could be considered to be an intermediate between Coelophysis and Dilophosaurus. Although the skull is not well known, many reconstructions have Liliensternus with a crest similar to that observed in Dilophosaurus. Its ilium (hip bone) is unusually short, as is the case with Dilophosaurus.[2] Liliensternus has five fingers, much like its contemporaries, but its fourth and fifth digits are smaller than the rest, a possible transitional stage between the five fingered theropods of the Triassic and the three fingered theropods of the Jurassic.[4]

Rauhut et al. (1998) noted that the remains may represent a juvenile or subadult individual based on the presence of only two fused sacrals and the fact that the neurocentral sutures are still visible in the vertebrae.[5]

A diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism or group. According to Rauhut (2000), Liliensternus can be distinguished based on the following features: the cervical vertebrae feature a broad rounded ridge that extends from the posterior end of the diapophyses to the posteroventral end of the vertebral centrum, one pair of pleurocoels in the cervical vertebrae, a less developed infradiapophyseal fossa, the absence of a horizontal ridge at the basis of the cervical neural spines, absence of a lateral bulge on the ilium.[6]

Discovery

Wappen Bedheim
Coat of Arms of Bedheim, depicting L. liliensterni[7]
Liliensternus
Liliensternus restored as a Coelophysoid

The specimens of Liliensternus, designated as the syntype series HMN BM.R.2175, were recovered near Großen Gleichberg in the Trossingen Formation of the Middle Keuper Group in Thuringia, Germany, together with remains of Ruehleia. The syntypes were discovered by Count Hugo Rühle von Lilienstern in the winter of 1932/1933[8] in marlstone (lime-rich mudstone) deposited in the Norian stage of the Late Triassic period, approximately 228 to 208 million years ago.[9] A left metatarsal later assigned to this genus, also deposited in the Norian, was collected in 1834 in the sandstone of the Trossingen Formation in Bayern, Germany. This metatarsal was originally described as a manual or pedal element by Meyer (1855) and a pubic fragment by Huene (1908) of Plateosaurus, and the material was re-identified as a proximal metatarsal belonging to Liliensternus by Moser in 2003.[5][10] Sander (1992) referred additional material to Liliensternus, which was thought to have been collected in 1961 grey/green marlstone from the Löwenstein Formation in Aargau, Switzerland, which is considered to also be from the Norian stage of the Late Triassic period.[11] A tooth referred to this Liliensternus, deposited in the Norian, and collected in 1913 in dark red mudstone from the Löwenstein Formation in Baden-Württemberg, Germany. The only material assigned to this genus from later strata was discovered in 1913 in blue claystone from the Rhaetian stage of the Late Triassic, in the Trossingen Formation from Sachsen-Anhalt, Germany, approximately 208 to 201 million years ago. The Liliensternus specimens remained in Hugo Rühle von Lilienstern's castle until 1969 when they were transferred to the collection of the Natural History Museum of Berlin, their present location.

The genus and specific names Liliensternus liliensterni are derived from the last name of count, amateur paleontologist, and medical doctor, Hugo Rühle von Lilienstern. This dinosaur was named in his honor for his furthering paleontology in Germany by founding a paleontological museum in his castle in Bedheim, on 1 July 1934. Liliensternus was described by Friedrich von Huene in 1934. Because it was originally named by von Huene as a member of the genus Halticosaurus, the type species of the genus Liliensternus is Halticosaurus liliensterni; the combinatio nova is Liliensternus liliensterni.

In 1993 Gilles Cuny and Peter Galton described a new species that they assigned to this genus, Liliensternus airelensis.[12] Other researchers began to notice differences between L. airelensis and the type species, L. liliensterni,[13] and in 2007, Martin Ezcurra and Cuny assigned the material to its own genus, Lophostropheus.[14]

Classification

Liliensternus sp
Teeth assigned to Liliensternus sp.

In 1934, Huene described two skeletons assigning them the name Halticosaurus liliensterni, but in 1984 Samuel Paul Welles concluded that the type species of Halticosaurus, H. longotarsus, was a nomen dubium. Most what had been written in the literature about Halticosaurus in fact regarded H. liliensterni. Welles therefore erected a new genus: Liliensternus, the name again honoring Rühle von Lilienstern.[15] The new species name became Liliensternus liliensterni. Rowe (1989) found that Liliensternus is more derived than Dilophosaurus. A second species named in 1993 by Cuny and Galton for fragmentary remains found in France, Liliensternus airelensis, which had an extra pair of cervical pleurocoels, was in 2007 reassigned to a separate genus, Lophostropheus.[16] Originally assigned to the Halticosauridae, Liliensternus is today considered a basal member of the Neotheropoda.

The following evolutionary tree illustrates a synthesis of the relationships of the early theropod groups compiled by Hendrickx et al. in 2015, including the position of Liliensternus in which all studies concur.[17]

Neotheropoda

Coelophysidae

Liliensternus

Zupaysaurus

Dilophosauridae

Dilophosaurus

Dracovenator

Averostra

Ceratosauria

Tetanurae

Cryolophosaurus

Sinosaurus

Monolophosaurus

Orionides

Paleoecology

Liliensternus abelov2014
Restoration of Liliensternus in its environment

Liliensternus was an active bipedal carnivore that could have preyed on the larger herbivores like Plateosaurus, which were present in its paleoenvironment. The material discovered in Switzerland, Sachsen-Anhalt, Germany and the tooth from Baden-Württemberg, Germany, suggest that Liliensternus inhabited ancient floodplains that were abundant with reptiles, therapsids, and Plateosaurus.[18] Paul (1988) noted that Liliensternus used its slashing tooth arrays to disable prosauropods and its speed to catch swift ornithischians.[2]

See also

References

  • F. v. Huene. 1934. Ein neuer Coelurosaurier in der thüringischen Trias [A new coelurosaur in the Thuringian Trias]. Paläontologische Zeitschrift 16(3/4):145-170
  1. ^ Rauhut, O.M.W. & A. Hungerbuhler, 1998, "A review of European Triassic theropods". Gaia 15. 75-88.
  2. ^ a b c Paul, Gregory S. (1988). Predatory Dinosaurs of the World. Simon & Schuster. p. 267. ISBN 0-671-61946-2.
  3. ^ "LILIENSTERNUS". Dinochecker.com. Retrieved 8 July 2013.
  4. ^ Dixon, Dougal (2015). The Complete Illustrated Encyclopedia of Dinosaurs. London: Hermes House.
  5. ^ a b Mortimer, Mickey (2012). "Coelophysoidea". Archived from the original on 2013-05-04.
  6. ^ Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.]. 440 pp.
  7. ^ http://www.ngw.nl/int/dld/b/bedheim.htm
  8. ^ Barbara A. R. Mohr, Evelyn Kustatscher, Cornelia Hiller and Gottfried Böhme, 2008, "Hugo Rühle v. Lilienstern and his palaeobotanical collection – an East-West German story", Earth Sciences History, 27: 278-296
  9. ^ F. v. Huene, 1934, "Ein neuer Coelurosaurier in der thüringischen Trias", Paläontologische Zeitschrift 16(3/4): 145-170
  10. ^ Moser, 2003. Plateosaurus engelhardti Meyer, 1837 (Dinosauria: Sauropodomorpha) aus dem Feuerletten (Mittelkeuper; Obertrias) von Bayern. Zitteliana B 24, 3-186.
  11. ^ Sander, 1992. The Norian Plateosaurus bonebeds of central Europe and their taphonomy. Palaeogeography, Palaeoclimatology, Palaeoecology. 93:255-296.
  12. ^ Cuny, Gilles; Galton, Peter M. (1993). "Revision of the Airel theropod dinosaur from the Triassic-Jurassic boundary (Normandy, France)". Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 187 (3): 261–288.
  13. ^ Rauhut, Oliver W.M.; Hungerbühler, A. (2002). "A review of European Triassic theropods". Gaia. 15: 75–88.
  14. ^ Ezcurra, Martin D.; Cuny, Gilles (2007). "The coelophysoid Lophostropheus airelensis, gen. nov.: a review of the systematics of "Liliensternus" airelensis from the Triassic-Jurassic boundary outcrops of Normandy (France)". Journal of Vertebrate Paleontology. 27 (1): 73–86. doi:10.1671/0272-4634(2007)27[73:TCLAGN]2.0.CO;2.
  15. ^ S.P. Welles, 1984, "Dilophosaurus wetherilli (Dinosauria, Theropoda): osteology and comparisons", Palaeontographica Abteilung A 185: 85-180
  16. ^ Ezcurra, M.D, and Cuny, G. (2007). "The coelophysoid Lophostropheus airelensis, gen. nov.: a review of the systematics of "Liliensternus" airelensis from the Triassic-Jurassic boundary outcrops of Normandy (France)." Journal of Vertebrate Paleontology, 27(1): 73-86.
  17. ^ Hendrickx, C., Hartman, S.A., & Mateus, O. (2015). An Overview of Non- Avian Theropod Discoveries and Classification. PalArch’s Journal of Vertebrate Palaeontology, 12(1): 1-73.
  18. ^ O. Jaekel. 1913. Über die Wirbeltierfunde in der oberen Trias von Halberstadt [On the vertebrate finds in the Upper Triassic of Halberstadt]. Paläontologische Zeitschrift 1:155-215

External links

Averostra

Averostra, or "bird snouts", is a clade that includes most theropod dinosaurs that have a promaxillary fenestra (fenestra promaxillaris), an extra opening in the front outer side of the maxilla, the bone that makes up the upper jaw. Two groups of averostrans, the Ceratosauria and the Orionides, survived into the Cretaceous period. When the Cretaceous–Paleogene extinction event occurred, ceratosaurians and two groups of orionideans within the clade Coelurosauria, the Tyrannosauroidea and Maniraptoriformes, were still extant. Only one subgroup of maniraptoriformes, Aves, survived the extinction event and persisted to the present day.

Avetheropoda

Avetheropoda, or "bird theropods", is a clade that includes carnosaurians and coelurosaurs to the exclusion of other dinosaurs.

Cerapoda

Cerapoda ("ceratopsians and ornithopods") is a clade of the dinosaur order Ornithischia.

Coelophysoidea

Coelophysoidea were common dinosaurs of the Late Triassic and Early Jurassic periods. They were widespread geographically, probably living on all continents. Coelophysoids were all slender, carnivorous forms with a superficial similarity to the coelurosaurs, with which they were formerly classified, and some species had delicate cranial crests. Sizes range from about 1 to 6 m in length. It is unknown what kind of external covering coelophysoids had, and various artists have portrayed them as either scaly or feathered. Some species may have lived in packs, as inferred from sites where numerous individuals have been found together.

Examples of coelophysoids include Coelophysis, Procompsognathus and Liliensternus. Most dinosaurs formerly referred to as being in the dubious taxon "Podokesauridae" are now classified as coelophysoids.

Daemonosaurus

Daemonosaurus (pron.:"DAY-mow-no-SORE-us") is an extinct genus of theropod dinosaur from the Late Triassic of New Mexico. Fossils have been found from deposits in the Chinle Formation, which is latest Triassic in age. While theropods had diversified into several specialized groups by this time, Daemonosaurus is a basal theropod that lies outside the clade Neotheropoda. Daemonosaurus is unusual among early theropods in that it had a short skull and long protruding teeth.

Dinosauriformes

Dinosauriformes is a clade of archosaurian reptiles that include the dinosaurs and their most immediate relatives. All dinosauriformes are distinguished by several features, such as shortened forelimbs and a partially to fully perforated acetabulum, the hole in the hip socket traditionally used to define dinosaurs. The oldest known member is Asilisaurus, dating to about 245 million years ago in the Anisian age of the middle Triassic period.

Dolichosuchus

Dolichosuchus (meaning "long crocodile") is the name given to a genus of dinosaur from the Triassic. It was originally classified in the disused family Hallopodidae, but has since been reclassified as a coelophysoid. A single fossil (consisting of a single lower leg bone, or tibia) was found in Germany. Since only one bone was discovered, the genus is considered a nomen dubium. Some scientists have noted that the tibia closely resembles those of Liliensternus and Dilophosaurus.The type species is D. cristatus, described by Huene in 1932. The bone was recovered from the Lower or Middle Stubensandstein formation.

Gojirasaurus

Gojirasaurus (meaning "Godzilla lizard") is a dubious genus of coelophysoid theropod dinosaur named after the giant monster movie character Gojira (the Japanese name for the monster Godzilla).

Halticosaurus

Halticosaurus (pron.:"HAL-tick-oh-SORE-us") is a dubious genus of theropod dinosaur from the late Triassic period (middle Norian stage, around 208 million years ago). It is known from a single fragmentary fossil specimen of the species H. longotarsus, found in the Middle Stubensandstein formation of what is present-day Germany. The only known specimen was poorly preserved and may have been put together from bones of unrelated animals. Further research would be required to determine which of the bones belonged together, and what kind of theropod Halticosaurus was. However, most of the bones have been lost. For these reasons, Halticosaurus is considered to be a nomen dubium.

Herrerasauridae

Herrerasauridae is a family of carnivorous basal saurischian dinosaurs. They are among the oldest known dinosaurs, first appearing in the fossil record around 233.23 million years ago (Late Triassic), before becoming extinct by the end of the Triassic period. Herrerasaurids were relatively small-sized dinosaurs, normally not more than 4 metres (13 ft) long. The best known representatives of this group are from South America (Brazil, Argentina), where they were first discovered in the 1960s. A nearly complete skeleton of Herrerasaurus ischigulastensis was discovered in the Ischigualasto Formation in San Juan, Argentina, in 1988. Less complete herrerasaurids have been found in North America, and they may have inhabited other continents as well.

Herrerasaurid anatomy is unusual and specialized, and they are not considered to be ancestral to any later dinosaur group. They only superficially resemble theropods and often present a mixture of very primitive and derived traits. The acetabulum is only partly open, and there are only two sacral vertebrae, the lowest number among dinosaurs. The pubic bone has a derived structure, being rotated somewhat posteriorly and folded to create a superficially tetanuran-like terminal expansion, especially prominent in H. ischigulastensis. The hand is primitive in having five metacarpals and the third finger longer than the second, but resembles those of theropods in having only three long fingers, with curved claws. Herrerasaurids also have a hinged mandible, which is also found in theropods.

Jingshanosaurus

Jingshanosaurus (meaning "Jingshan lizard") is a genus of sauropodomorph dinosaurs from the early Jurassic period.

Lepidus praecisio

Lepidus is a genus of extinct coelophysoidean theropod from the Upper Triassic of the United States. It lived in the Otis Chalk localities of the Dockum Group in Texas, around 223 million years ago.

Lophostropheus

Lophostropheus (pron.:" LOAF-oh-STRO-fee-us") is an extinct genus of coelophysoid theropod dinosaur that lived approximately 200 million years ago during the boundary between the Late Triassic Period and the Early Jurassic Period, in what is now Normandy, France. Lophostropheus is one of the few dinosaurs that may have survived the Triassic–Jurassic extinction event.

Lophostropheus was a small to medium-sized, moderately-built, ground-dwelling, bipedal carnivore, that could grow up to 3 m (9.8 ft) long. Over the years it had been incorrectly classified as Halticosaurus and Liliensternus, but was later recognized as a new genus and was reassigned to Lophostropheus in 2007.

Neotheropoda

Neotheropoda (meaning "new theropods") is a clade that includes coelophysoids and more advanced theropod dinosaurs, and the only group of theropods who survived the Triassic–Jurassic extinction event. Yet all of the neotheropods became extinct during the early Jurassic period except for Averostra.

Norian

The Norian is a division of the Triassic geological period. It has the rank of an age (geochronology) or stage (chronostratigraphy). The Norian lasted from ~227 to 208.5 million years ago. It was preceded by the Carnian and succeeded by the Rhaetian.

Orionides

Orionides is a clade of tetanuran theropod dinosaurs from the Middle Jurassic to the Present. The clade includes most theropod dinosaurs, including birds.

Rhaetian

See Raetians for the Alpine people of antiquity. See Raetian language for their language.The Rhaetian is, in geochronology, the latest age of the Triassic period or in chronostratigraphy the uppermost stage of the Triassic system. It lasted from 208.5 to 201.3 million years ago. It was preceded by the Norian and succeeded by the Hettangian (the lowermost stage or earliest age of the Jurassic).In this age, Pangaea began to break up, though the Atlantic Ocean was not yet formed.

Trossingen Formation

The Trossingen Formation, formerly the Knollenmergel, is a geological formation in Germany. It dates back to the late Norian.

Zupaysaurus

Zupaysaurus (; "ZOO-pay-SAWR-us") is a genus of early theropod dinosaur living during the Norian stage of the Late Triassic in what is now Argentina. Fossils of the dinosaur were found in the Los Colorados Formation of the Ischigualasto-Villa Unión Basin in northwestern Argentina. Although a full skeleton has not yet been discovered, Zupaysaurus can be considered a bipedal predator, up to 4 metres (13 ft) long. It may have had two parallel crests running the length of its snout.

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