Procompsognathus

Procompsognathus /ˌproʊkɒmpˈsɒɡnəθəs/ is an extinct genus of coelophysid theropod dinosaur that lived approximately 210 million years ago during the later part of the Triassic Period, in what is now Germany. Procompsognathus was a small-sized, lightly built, ground-dwelling, bipedal carnivore, that could grow up to 1 m (3.3 ft) long.

Procompsognathus
Temporal range: Late Triassic, 210 Ma
Procompsognathus triassicus
Holotype specimen (SMNS 12591)
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Family: Coelophysidae
Nopcsa, 1923
Genus: Procompsognathus
Fraas, 1913
Species:
P. triassicus
Binomial name
Procompsognathus triassicus
Fraas, 1913

Discovery and naming

The fragmentary and poorly preserved skeleton of Procompsognathus was found in the Middle Stubensandstein member of the Löwenstein Formation at the Weiße Steinbruch, the quarry of Albert Burrer on the northern slopes of the Stromberg region near Pfaffenheim in Württemberg, Germany.[1] The discovery was made by Albert Burrer in the spring of 1909 in white sandstone and gray/blue marl sediments that were deposited during the Norian stage of the Triassic period, approximately 210 million years ago.

The holotype SMNS 12591, consisted of three blocks of sandstone: one showed a small, seven-centimetre-long, severely crushed skull with lower jaws. The second and third contained the partly articulated remains of a postcranial skeleton, including twenty-nine vertebrae of the neck, back and tail; ribs; elements of the shoulder girdle and a forelimb; an ilium; both pubis and the hindlimbs. It represents an adult individual.

Burrer sent the specimen to Professor Eberhard Fraas of the königliche Stuttgarter Naturalienkabinett. In a lecture on 9 October 1911, Fraas referred to it by the name "Hallopus celerrimus", considering it jumping form of dinosaur that was approximately 60 cm (2.0 ft) long, and was associated with the origin of birds.[2] Later, Fraas decided to use a different name in the official publication.[3] Fraas in 1913 named the genus Procompsognathus with as type species, Procompsognathus triassicus.[4] The type specimen is housed in the collection of the State Museum of Natural History in Stuttgart, Germany.

In 1921, Friedrich von Huene referred two more specimens, both also found in the Burrer quarry, in 1908, to Procompsognathus: SMNS 12352, a partial skull and lower jaws from a larger individual than the holotype, and SMNS 12352a, an isolated left hand.[5]

The genus name Procompsognathus, means "before elegant jaw", and is derived from the name of another dinosaur, Compsognathus. A later (Jurassic) small predatory dinosaur, Compsognathus takes its name from the Greek word kompsos (κομψός) which can be rendered as "elegant", "refined" or "dainty" and the Greek word gnathos (γνάθος) which means "jaw". The prefix Pro (προ) implies "before" or "ancestor of", although this direct lineage was not supported by subsequent research. The specific name triassicus refers to the geologic time period to which this dinosaur belongs, the Triassic.[6]

Description

Procompsognathus may have been about 1 metre (3.3 ft) long,[7] though Fraas in 1913 estimated a length of 75 cm (2.5 ft). In 2010 Gregory S. Paul gave an estimate of 1 kilogram (2.2 lb) for the weight with a length of 1.1 metres (3.6 ft).[8] A biped, Procompsognathus had long hind legs, short arms, large clawed hands, a long slender snout with many small teeth, and a stiff tail. The femur discovered for the type specimen of this genus measures 93mm and the tibia, 112mm. The tibia is approximately 20% longer than the femur in Procompsognathus, an adaptation which has been strongly correlated with the development of cursorial habits in dinosaurs, suggesting that they were good runners.[9]

Distinguishing anatomical features

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. In 2000, Rauhut noted that its scapula is more slender than that of Coelophysis bauri.

In 1998 Chatterjee noted that the skull of Procompsognathus possesses a suite of theropod synapomorphies, which included:

  • the presence of an accessory maxillary fenestra
  • the vomers are fused rostrally and are extended considerably forward to the choana
  • the single quadrate head is received completely by squamosal without paroccipital contact
  • the lateroshenoid possesses a transverse postorbital process
  • the orbitosphenoids are fused

Classification

Type specimen SMNS 12591

While it is undoubtedly a small, bipedal carnivore, the extremely poor preservation of the only known Procompsognathus fossil makes its exact identity difficult to determine. Fraas originally assigned it to the Dinosauria. In 1923 Franz Nopcsa coined a Procompsognathinae, and in 1929 von Huene created a Procompsognathidae, though these concepts are today rarely used. In 1932 von Huene saw it as a member of the non-dinosaurian Pseudosuchia.[10] It has since then usually been considered a theropod dinosaur, with some exceptions. In 1992 Paul Sereno and Rupert Wild stated that the holotype specimen consisted of fossils from two separate animals: the postcranial skeleton would be a theropod, likely a coelophysoid related to Segisaurus, but the skull and the von Huene specimens they referred to the basal crocodylomorph Saltoposuchus connectens.[11] However, in 1993 Sankar Chatterjee after further preparation refuted their assessment and regarded the skull as that of a theropod similar to Syntarsus, and demonstrated that it could not have been a crocodylomorph, as it lacks the landmark features of this group.[12][13] Sereno (1997) and Ezcurra and Novas (2007) conducted phylogenetic analysis that supported the placement of Procompsognathus in the taxon Coelophysidae.[14][15] This genus may be most closely related to Segisaurus halli.[16]

Referred specimens SMNS 12352 and SMNS 12352a

Much controversy has arisen however, about the association with the later material referred by von Huene. In 1982 John Ostrom claimed that SMNS 12352 and SMNS 12352a originated from a taxon different from the holotype. In 2006 and 2008 Fabien Knoll concluded that SMNS 12352 represented a crocodylomorph and SMNS 12352a a crocodylomorph or some other basal archosaur. The postcranial skeleton, for which she reserved the inventory number SMNS 12591, was a coelophysoid; and the skull, now indicated with the number SMNS 12591a, a perhaps more derived theropod, possibly a basal member of the Tetanurae.[17][18] In 2012 Knoll after a CAT-scan reaffirmed that SMNS 12352 was a crocodylomorph, but established it was different from Saltoposuchus.[19]

Oliver Rauhut and Axel Hungerbuhler (2000) noted features of the vertebrae which suggest that Procompsognathus may be a coelophysid or ceratosaur,[20] and Carrano et al. (2005), in their re-study of the related genus Segisaurus, found both Segisaurus and Procompsognathus to belong to the Coelophysidae within Dinosauria.[21] In 2004 David Allen considered Procompsognathus to be a primitive, non-dinosaurian ornithodiran.[22]

Paleoecology

Procompsognathus lived in a relatively dry, inland environment and may have eaten insects, lizards and other small prey.[23] Contemporaries of Prompsognathus included the coelophysoids Halticosaurus and Dolichosuchus, as well as the Sauropodomorphs Plateosaurus gracilis and Efraasia minor. Weishampel, et al. (2004) noted that theropod tracks and fossils of an unnamed herrerasaur genus are known from the Lower Stubensandstein.[24]

In popular culture

Procompsognathus appears in the novel Jurassic Park and its sequel The Lost World by Michael Crichton. The species is sometimes referred to as "compys" by the characters. While the author invents a venomous bite for Procompsognathus with soporific effects, there is no evidence to support venom in Procompsognathus.[25]

References

  1. ^ E. Fraas. 1907. Aetosaurus crassicauda n. sp. nebst Beobachtungen über das Becken der Aetosaurier [Aetosaurus crassicauda n. sp. together with observations on the pelvis of aetosaurs]. Mitteilungen aus dem Königlichen Naturalien-Kabinett zu Stuttgart 42:101-109
  2. ^ Fraas, E., 1911, "Die schwäbische Dinosaurier", Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg 1912: LXVI-LXVII
  3. ^ Pompeckj, J.F., 1915, "Zur Erinnerung an Eberhard Fraas und an sein Werk", Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg 1924: XXXIII
  4. ^ Fraas, E., 1913, "Die neuesten Dinosaurierfunde in der schwäbischen Trias", Naturwissenschaften 1(45): 1097-1100
  5. ^ Von Huene, F., 1921, "Neue Pseudosuchier und Coelurosaurier aus dem württembergischen Keuper", Acta Zoologica 2: 329-403
  6. ^ Liddell, Henry George and Robert Scott (1980). A Greek-English Lexicon (Abridged Edition). United Kingdom: Oxford University Press. ISBN 0-19-910207-4.
  7. ^ Holtz, Thomas R. Jr. (2011) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages, Winter 2010 Appendix.
  8. ^ Paul, G.S., 2010, The Princeton Field Guide to Dinosaurs, Princeton University Press p. 72
  9. ^ Coombs, W. P., Jr. 1978. Theoretical aspects of cursorial adaptations in dinosaurs. The Quarterly Review of Biology 53:393–418.
  10. ^ von Huene, F., 1932, Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien für Geologie und Paläontologie, 4, pp. 361
  11. ^ Sereno, Paul C.; Wild, Rupert (December 15, 1992). "Procompsognathus: theropod, "thecodont" or both?". Journal of Vertebrate Paleontology. 12 (4): 435–458. doi:10.1080/02724634.1992.10011473.
  12. ^ Chatterjee, S., 1993, "Procompsognathus from the Triassic of Germany is not a crocodylomorph", Journal of Vertebrate Paleontology 13(3): 29A
  13. ^ Chatterjee, S., 1998, "Reassessment of the Procompsognathus skull", p. 6 in: Wolberg, D.L., Gittis, K., Miller, S., Carey, L., and A. Raynor (eds.), Dinofest International. The Academy of Natural Sciences, Philadelphia
  14. ^ P. C. Sereno. 1997. The origin and evolution of dinosaurs. Annual Review of Earth and Planetary Sciences 25:435-489
  15. ^ M. D. Ezcurra and F. E. Novas. 2007. Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina. Historical Biology 19(1):35-72
  16. ^ Carrano, M.T., Hutchinson, J.R., & Sampson, S.D. 2005. New information on Segisaurus halli, a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate Paleontology 25(4): 835-849.
  17. ^ Knoll, F. & Schoch, R., 2006, "Does Procompsognathus have a head? Systematics of an enigmatic Triassic taxon", Journal of Vertebrate Paleontology 26: 86A
  18. ^ Knoll, F., 2008, "On the Procompsognathus postcranium (Late Triassic, Germany)", Geobios 41: 779-786
  19. ^ Knoll, F. & Schoch, R., 2012, "CT scanning, rapid prototyping and re-examination of a partial skull of a basal crocodylomorph from the Late Triassic of Germany", Swiss Journal of Geosciences 105: 109-115
  20. ^ Rauhut, O. and Hungerbuhler, A. (2000). "A review of European Triassic theropods." Gaia 15, 75-88
  21. ^ Carrano, M.T, Hutchinson, J.R, Sampson, S.D. (2005). "New information on Segisaurus halli, a small theropod dinosaur from the Early Jurassic of Arizona." Journal of Vertebrate Paleontology. 25(4):835-849
  22. ^ Allen, D. (2004). "The phylogenetic status of Procompsognathus revisited." Journal of Vertebrate Paleontology, 24(3): 34A
  23. ^ Frances Freedman & Tony Gibbons, 1997, Looking at-- Procompsognathus: a dinosaur from the Triassic period, Gareth Stevens Pub., 1 January 1997, pp. 24
  24. ^ Baden-Wurrtemberg, Germany; 1. Lower Stubensandstein," in Weishampel, et al. (2004). Page 524.
  25. ^ Bennington, J Bret (1996). "Errors in the Movie "Jurassic Park"". American Paleontologist. 4 (2): 4–7.
1913 in paleontology

Paleontology or palaeontology is the study of prehistoric life forms on Earth through the examination of plant and animal fossils. This includes the study of body fossils, tracks (ichnites), burrows, cast-off parts, fossilised feces (coprolites), palynomorphs and chemical residues. Because humans have encountered fossils for millennia, paleontology has a long history both before and after becoming formalized as a science. This article records significant discoveries and events related to paleontology that occurred or were published in the year 1913.

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.

Coelophysidae

Coelophysidae is a family of primitive carnivorous theropod dinosaurs. Most species were relatively small in size. The family flourished in the Late Triassic and Early Jurassic periods, and has been found on numerous continents. Many members of Coelophysidae are characterized by long, slender skulls and light skeletons built for speed. One member, Coelophysis, displays the earliest known furcula in a dinosaur.Under cladistic analysis, Coelophysidae was first defined by Paul Sereno in 1998 as the most recent common ancestor of Coelophysis bauri and Procompsognathus triassicus, and all of that common ancestor's descendants. However, Tykoski (2005) has advocated for the definition to change to include the additional taxa of "Syntarsus" kayentakatae and Segisaurus halli. Coelophysidae is part of the superfamily Coelophysoidea, which in turn is a subset of the larger Neotheropoda clade. As part of Coelophysoidea, Coelophysidae is often placed as sister to the Dilophosauridae family, however, the monophyly of this clade has often been disputed. The older term "Podokesauridae", named 14 years prior to Coelophysidae (which would normally grant it priority), is now usually ignored, since its type specimen was destroyed in a fire and can no longer be compared to new finds.

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.

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.

Jeholosauridae

Jeholosaurids were herbivorous neornithischian dinosaurs from the Cretaceous Period (Aptian - Santonian, with a possible Campanian record) of Asia. The family was first proposed by Han et al. in 2012. The jeholosaurids were defined as those ornithischians more closely related to Jeholosaurus shangyuanensis than to Hypsilophodon foxii, Iguanodon bernissartensis, Protoceratops andrewsi, Pachycephalosaurus wyomingensis, or Thescelosaurus neglectus. The Jeholosauridae includes the type genus Jeholosaurus and Yueosaurus.

Jingshanosaurus

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

Melanorosauridae

The Melanorosauridae were a family of sauropodomorph dinosaurs which lived during the Late Triassic and Early Jurassic. The name Melanorosauridae was first coined by Friedrich von Huene in 1929. Huene assigned several families of dinosaurs to the infraorder "Prosauropoda": the Anchisauridae, the Plateosauridae, the Thecodontosauridae, and the Melanorosauridae. Since then, these families have undergone numerous revisions. Galton and Upchurch (2004) considered Camelotia, Lessemsaurus, and Melanorosaurus members of the family Melanorosauridae. A more recent study by Yates (2007) indicates that the melanorosaurids were instead early sauropods.

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.

Orionides

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

Orodrominae

Orodrominae is a subfamily of parksosaurid dinosaurs from the Cretaceous of North America and Asia.

Pterospondylus

Pterospondylus (meaning "winged vertebra") is a dubious genus of theropod dinosaur from the Late Triassic. It was a coelophysoid theropod which lived in what is now Germany (Trossingen Formation). The type species, Pterospondylus trielbae, was described by Jaekel in 1913-14 for a single back vertebra. Sometimes, it is aligned with Procompsognathus, or even considered to be synonymous with it, despite being based on a vertebra that is twice the size of the corresponding bone in Procompsognathus.The specific name trielbae is derived from "Tri", in reference to the Triassic period, and "Elba", which points to the Elbe River area.

Riojasauridae

Riojasauridae is a family of sauropod-like dinosaurs from the Upper Triassic. It is known primarily from the genera Riojasaurus and Eucnemesaurus. Sites containing Riojasauridae include the Lower Elliot Formation of Orange Free State, South Africa (where fossils of Eucnemesaurus have been found), and Ischigualasto, in La Rioja Province, Argentina ( where fossils of Riojasaurus have been recovered).

Saltopus

Saltopus ("hopping foot") is a genus of very small bipedal dinosauriform containing the single species S. elginensis from the late Triassic period of Scotland. It is one of the most famous Elgin Reptiles.

Segisaurus

Segisaurus (meaning "Segi canyon lizard") is a genus of small coelophysoid theropod dinosaur, that measured approximately 1 metre (3.3 feet) in length. The only known specimen was discovered in early Jurassic strata in Tsegi Canyon, Arizona, for which it was named. Segisaurus is the only dinosaur to have ever been excavated from the area.

Timeline of coelophysoid research

This timeline of coelophysoid research is a chronological listing of events in the history of paleontology focused on the coelophysoids, a group of primitive theropod dinosaurs that were among Earth's dominant predators during the Late Triassic and Early Jurassic epochs. Although formally trained scientists didn't discover coelophysoid fossils until the late 19th century, Native Americans of the modern southwestern United States may have already encountered their fossils. Navajo creation mythology describes the early Earth as being inhabited by a variety of different kinds of monsters who hunted humans for food. These monsters were killed by storms and the heroic Monster Slayers, leaving behind their bones. As these tales were told in New Mexico not far from bonebeds of Coelophysis, this dinosaur's remains may have been among the fossil remains that inspired the story.The first scientifically documented coelophysoid taxon was Coelophysis bauri itself. However, when the species was first described by Edward Drinker Cope in 1887, it was thought to belong to a genus of small carnivorous dinosaurs called Coelurus. Later that same year Cope changed his mind and transferred it to the genus Tanystropheus. Tanystropheus turned out to be a long-necked reptile not regarded by scientists as a true dinosaur. As such, "Tanystrophaeus" bauri was soon given its own genus, Coelophysis in 1889. Over the ensuing decades, many new coelophysoids would be discovered, like Podokesaurus, Procompsognathus, and Segisaurus.In 1947, a paleontological team led by Edwin Colbert made a major discovery in New Mexico. While on an expedition to Petrified Forest National Park in Arizona, he made a detour to Ghost Ranch, New Mexico where many phytosaur fossils had been found. There they discovered a massive bonebed preserving hundreds of Coelophysis, many of which were complete and articulated. The find has been considered the most significant Triassic fossil discovery in North America. Later, other coelophysoids and even other bonebeds would be discovered. Notable coelophysoids discovered during the mid to late 20th century include Syntarsus (now Megapnosaurus) and Gojirasaurus. Despite this extensive history of research, the formal recognition of the Coelophysoidea as a distinct group of dinosaurs is relatively recent and the group would not be formally named until a 1994 by Thomas Holtz.

Xixiposaurus

Xixiposaurus is a genus of prosauropod dinosaur which existed in what is now Lower Lufeng Formation, China during the lower Jurassic period. It was first named by Sekiya Toru in 2010 and the type species is Xixiposaurus suni.

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