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''Liliensternus'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of basal
neotheropod Neotheropoda (meaning "new theropods") is a clade that includes coelophysoids and more advanced theropod dinosaurs, and is the only group of theropods that survived the Triassic–Jurassic extinction event. All neotheropods became extinct by the e ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
that lived approximately 210
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
during the latter part of the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
Period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
in what is now Germany. ''Liliensternus'' was a moderate-sized,
bipedal Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
, ground-dwelling
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
, that could grow up to long. It is the best represented Triassic theropod from Europe and one of the largest known.Rauhut, O.M.W. & A. Hungerbuhler, 1998, "A review of European Triassic theropods". ''Gaia'' 15. 75-88.


Description

''Liliensternus'' was approximately long, and may have weighed about . Other estimates suggest that ''Liliensternus'' was at best long and weighed at most. The remains of two specimens of ''Liliensternus'' together form a
syntype In biological nomenclature, a syntype is any one of two or more biological types that is listed in a description of a taxon where no holotype was designated. Precise definitions of this and related terms for types have been established as part of ...
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 ''Dilophosaurus'' ( ) is a genus of theropod dinosaurs that lived in what is now North America during the Early Jurassic, about 193 million years ago. Three skeletons were discovered in northern Arizona in 1940, and the two best preserved ...
'' and ''Liliensternus'', unlike those of coelophysid taxa, such as ''
Coelophysis ''Coelophysis'' ( traditionally; or , as heard more commonly in recent decades) is an extinct genus of coelophysid theropod dinosaur that lived approximately 228 to 201.3 million years ago during the latter part of the Triassic Period from t ...
''. 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''. ''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. 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
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e. 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 Skeletal pneumaticity is the presence of air spaces within bones. It is generally produced during development by excavation of bone by pneumatic diverticula (air sacs) from an air-filled space, such as the lungs or nasal cavity. Pneumatization is h ...
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.


Discovery

The specimens of ''Liliensternus'', designated as the
syntype In biological nomenclature, a syntype is any one of two or more biological types that is listed in a description of a taxon where no holotype was designated. Precise definitions of this and related terms for types have been established as part of ...
series HMN BM.R.2175, were recovered near
Großer Gleichberg At , the Großer Gleichberg is the higher of the two adjacent mountaintops known as the Gleichberge east of Römhild in the landscape of Grabfeld and the county of Hildburghausen in the German state of Thuringia. It was formed in the Tertiary vo ...
in the
Trossingen Formation The Trossingen Formation, formerly the Knollenmergel, is a geological formation in Germany and Switzerland. It dates back to the late Norian- Rhaetian.Weishampel et al., 2004, pp.521–525 Vertebrate paleofauna See also * List of dinosa ...
of the Middle Keuper Group in Thuringia,
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
, together with remains of ''
Ruehleia ''Ruehleia'' is a genus of sauropodomorph dinosaur from the Late Triassic period of Germany. The type species is ''R. bedheimensis'', described by Galton in 2001,Galton, P.M. (2001). "Prosauropod dinosaurs from the Upper Triassic of Germany", In ...
''. The syntypes were discovered by Count Hugo Rühle von Lilienstern in the winter of 1932/1933 in marlstone (lime-rich mudstone) deposited in the
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
stage of the Late
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
period, approximately 228 to 208 million years ago.F. v. Huene, 1934, "Ein neuer Coelurosaurier in der thüringischen Trias", ''Paläontologische Zeitschrift'' 16(3/4): 145-170 A left metatarsal later assigned to this genus, also deposited in the
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
, was collected in 1834 in the sandstone of the
Trossingen Formation The Trossingen Formation, formerly the Knollenmergel, is a geological formation in Germany and Switzerland. It dates back to the late Norian- Rhaetian.Weishampel et al., 2004, pp.521–525 Vertebrate paleofauna See also * List of dinosa ...
in
Bayern Bavaria ( ; ), officially the Free State of Bavaria (german: Freistaat Bayern, link=no ), is a state in the south-east of Germany. With an area of , Bavaria is the largest German state by land area, comprising roughly a fifth of the total lan ...
, Germany. This metatarsal was originally described as a manual or pedal element by Meyer (1855) and a pubic fragment by Huene (1908) of ''
Plateosaurus ''Plateosaurus'' (probably meaning "broad lizard", often mistranslated as "flat lizard") is a genus of plateosaurid dinosaur that lived during the Late Triassic period, around 214 to 204 million years ago, in what is now Central and Northern Eur ...
'', and the material was re-identified as a proximal metatarsal belonging to ''Liliensternus'' by Moser in 2003. 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 ). Swiss law does not designate a ''capital'' as such, but the federal parliament and government are installed in Bern, while other federal institutions, such as the federal courts, are in other cities (Bellinzona, Lausanne, Luzern, Neuchâtel ...
, which is considered to also be from the
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
stage of the Late Triassic period. A tooth referred to this ''Liliensternus'', deposited in the
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
, 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 The Rhaetian is the latest age of the Triassic Period (in geochronology) or the uppermost stage of the Triassic System (in chronostratigraphy). It was preceded by the Norian and succeeded by the Hettangian (the lowermost stage or earliest age ...
stage of the Late Triassic, in the
Trossingen Formation The Trossingen Formation, formerly the Knollenmergel, is a geological formation in Germany and Switzerland. It dates back to the late Norian- Rhaetian.Weishampel et al., 2004, pp.521–525 Vertebrate paleofauna See also * List of dinosa ...
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 Events January–February * January 1 – The International Telecommunication Union, a specialist agency of the League of Nations, is established. * January 15 – The 8.0 1934 Nepal–Bihar earthquake, Nepal–Bihar earthquake strik ...
. ''Liliensternus'' was described by
Friedrich von Huene Friedrich von Huene, born Friedrich Richard von Hoinigen, (March 22, 1875 – April 4, 1969) was a German paleontologist who renamed more dinosaurs in the early 20th century than anyone else in Europe. He also made key contributions about v ...
in
1934 Events January–February * January 1 – The International Telecommunication Union, a specialist agency of the League of Nations, is established. * January 15 – The 8.0 1934 Nepal–Bihar earthquake, Nepal–Bihar earthquake strik ...
.F. v. Huene. 1934. Ein neuer Coelurosaurier in der thüringischen Trias new coelurosaur in the Thuringian Trias ''Paläontologische Zeitschrift'' 16(3/4):145-170 Because it was originally named by von Huene as a member of the genus ''
Halticosaurus ''Halticosaurus'' (pron.:"HAL-tick-oh-SORE-us") is a dubious genus of theropod dinosaur from the late Triassic period (middle Norian stage, around 215.6–208 million years ago). It is known from a single fragmentary fossil specimen of the ...
'', the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of the genus ''Liliensternus'' is ''Halticosaurus liliensterni''; the ''
combinatio nova ''Combinatio nova'', abbreviated ''comb. nov.'' (sometimes ''n. comb.''), is Latin for "new combination". It is used in taxonomic biology literature when a new name is introduced based on a pre-existing name. The term should not to be confused wi ...
'' is ''Liliensternus liliensterni''. In 1993 Gilles Cuny and
Peter Galton Peter Malcolm Galton (born 14 March 1942 in London) is a British vertebrate paleontologist who has to date written or co-written about 190 papers in scientific journals or chapters in paleontology textbooks, especially on ornithischian and prosaur ...
described a new
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
that they assigned to this genus, ''Liliensternus airelensis.'' Other researchers began to notice differences between ''L. airelensis'' and the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
, ''L. liliensterni'', and in 2007, Martin Ezcurra and Cuny assigned the material to its own genus, '' Lophostropheus''.


Classification

In 1934, Huene described two skeletons assigning them the name ''Halticosaurus liliensterni'', but in 1984
Samuel Paul Welles Samuel Paul Welles (November 9, 1907 – August 6, 1997) was an American palaeontologist. Welles was a research associate at the Museum of Palaeontology, University of California, Berkeley. He took part in excavations at the Placerias Quarry in ...
concluded that the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of ''Halticosaurus'', ''H. longotarsus'', was a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium'' it may be impossible to determine whether a s ...
''. Most of 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. The new species name became ''Liliensternus liliensterni''. Rowe (1989) found that ''Liliensternus'' is more derived than ''
Dilophosaurus ''Dilophosaurus'' ( ) is a genus of theropod dinosaurs that lived in what is now North America during the Early Jurassic, about 193 million years ago. Three skeletons were discovered in northern Arizona in 1940, and the two best preserved ...
''. A second species named in 1993 by Cuny and Galton for fragmentary remains found in
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
, ''Liliensternus airelensis'', which had an extra pair of cervical pleurocoels, was in 2007 reassigned to a separate genus, '' Lophostropheus''. Originally assigned to the Halticosauridae, ''Liliensternus'' is today considered a basal member of the
Neotheropoda Neotheropoda (meaning "new theropods") is a clade that includes coelophysoids and more advanced theropod dinosaurs, and is the only group of theropods that survived the Triassic–Jurassic extinction event. All neotheropods became extinct by the ...
. 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.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.


Paleoecology

''Liliensternus'' was an active bipedal carnivore that could have preyed on the larger
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s like ''
Plateosaurus ''Plateosaurus'' (probably meaning "broad lizard", often mistranslated as "flat lizard") is a genus of plateosaurid dinosaur that lived during the Late Triassic period, around 214 to 204 million years ago, in what is now Central and Northern Eur ...
'', 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,
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
s, and ''
Plateosaurus ''Plateosaurus'' (probably meaning "broad lizard", often mistranslated as "flat lizard") is a genus of plateosaurid dinosaur that lived during the Late Triassic period, around 214 to 204 million years ago, in what is now Central and Northern Eur ...
''.O. Jaekel. 1913. Über die Wirbeltierfunde in der oberen Trias von Halberstadt n the vertebrate finds in the Upper Triassic of Halberstadt Paläontologische Zeitschrift 1:155-215 Paul (1988) noted that ''Liliensternus'' used its slashing tooth arrays to disable prosauropods and its speed to catch swift ornithischians.


See also

* Timeline of coelophysoid research


References


External links


''Liliensternus'' on the Dinosaur Mailing List
{{Taxonbar, from=Q132620 Coelophysoids Late Triassic dinosaurs of Europe Triassic Germany Fossils of Germany Fossil taxa described in 1984 Taxa named by Samuel Paul Welles