Compsognathidae

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Compsognathidae
Temporal range: Late Jurassic – Early Cretaceous, 151.5–108  Ma
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Possible Late Cretaceous record
Compsognathidae.jpg
Compsognathid skeletons to scale
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: Neocoelurosauria
Family:† Compsognathidae
Cope, 1871
Type species
† Compsognathus longipes
Wagner, 1861
Genera [1] [2]
Possible compsognathids
Synonyms
  • SinosauropterygidaeJi & Ji, 1996

Compsognathidae is a family of coelurosaurian theropod dinosaurs. Compsognathids were small carnivores, generally conservative in form, hailing from the Jurassic and Cretaceous Periods. The bird-like features of these species, along with other dinosaurs such as Archaeopteryx inspired the idea for the connection between dinosaur reptiles and modern-day avian species. [4] Compsognathid fossils preserve diverse integument — skin impressions are known from four genera commonly placed in the group, Compsognathus , Sinosauropteryx , Sinocalliopteryx and Juravenator . [5] While the latter three show evidence of a covering of some of the earliest primitive feathers over much of the body, Juravenator and Compsognathus also show evidence of scales on the tail or hind legs. "Ubirajara jubatus", informally described in 2020, had elaborate integumentary structures on its back and shoulders superficially similar to the display feathers of a standardwing bird-of-paradise, and unlike any other non-avian dinosaur currently described.

Contents

The first member of the group, Compsognathus , was discovered in 1861, after Johann A. Wagner published his description of the taxon. [6] [7] [8] The family was created by Edward Drinker Cope in 1875. [9] This classification was accepted by Othniel Charles Marsh in 1882, and added to the Coelurosauria clade by Friedrich von Huene in 1914 after additional fossils had been found. [6] With further discoveries, fossils have been uncovered across three different continents, in Asia, [5] [6] [7] [8] [10] Europe, [11] and South America. [7] [9] Assignment to Compsognathidae is usually determined through examination of the metacarpal, which is used to separate Compsognathidae from other dinosaurs. [12] However, classification is still complicated due to similarities to the body of several other theropod dinosaurs, as well as the lack of unifying, diagnostic features that are shared by all compsognathids. [9] [13] Some authors have proposed that Compsognathidae is not a monophyletic group, and at least some compsognathids represent juvenile specimens of larger tetanuran theropods, such as carnosaurs and tyrannosaurs. [14] [15] [16]

History of discovery

The holotype of Compsognathus longipes Compsognathus longipes cast 3.jpg
The holotype of Compsognathus longipes

The first significant fossil specimen of Compsognathidae was found in the Bavaria region of Germany (BSP AS I 563) and given to collector Joseph Oberndorfer in 1859. [17] The finding was initially significant because of the small size of the specimen. In 1861, after an initial period of review, Johann A. Wagner presented his analysis of the specimen to the public and named the fossil Compsognathus longipes ("elegant jaw"). [18] In 1868, Thomas Henry Huxley, an early supporter of Charles Darwin and his theory of evolution, used Compsognathus in a comparison to similar feathered dinosaur Archaeopteryx in order to propose the origin of birds. Huxley noticed that these dinosaurs shared a similar layout to birds and proposed an exploration of the similarities. He is credited as being the first person to do so. [19] This initial comparison sparked the interest into the origin of birds and feathers. In 1882, Othniel Charles Marsh named a new family of dinosaurs for this species Compsognathidae and officially recognized the species as part of Dinosauria. [20]

Notable Specimens

Size comparison of the Juravenator specimen to human Juravenator Scale.svg
Size comparison of the Juravenator specimen to human

In 1971, a second nearly complete specimen of Composgnathus longipes was found in the area of Canjuers, which is located in the southeast of France near Nice. [6] This specimen was much larger than the original German specimen, but similarities led to experts categorizing the fossil as an adult Compsognathus longipes and leading to the further classification of the German specimen as a juvenile. [21] This specimen also contained a lizard in the digestive region, further solidifying the theory that compsognathids consumed small vertebrate species.

The holotype of Juravenator is the only known specimen of the species. Though Juravenator has previously been accepted as a member of Compsognathidae, recent research has led some experts to believe that Juravenator does not belong in this group. This is because Juravenator could also be classified into a similar group within Coelurosauria, Maniraptoriformes. Maniraptoriforms share many similarities with compsognathids and, because there has been only one verified specimen of Juravenator, experts have disagreed on exactly where to place this genus. Since 2013, Juravenator is still commonly classified as a coelurosaur, but near the group Maniraptoriformes instead of Compsognathidae. [22]

A compsognathid specimen consisting of a single finger bone has been described from Late Jurassic (Tithonian Age, about 150 million years ago) sediments at Port Waikato, New Zealand. It is the first and so far, only dinosaur specimen known from Jurassic New Zealand, as well as being the first New Zealand dinosaur fossil to have been found outside of the Cretaceous marine sediments at Mangahouanga Stream. Possible coprolites have been referred to this specimen, however it is still not an officially classified species.

Description

Compsognathids share a variety of characteristics. The genera in this family demonstrate traits that are characteristic of theropods, such as smaller forelimbs than hind legs. Size, feathers and metacarpal size are among the most important classifying common characteristics.

Size

Comparison of German (green) and French (orange) Compsognathus longipes specimen Compysizes1.png
Comparison of German (green) and French (orange) Compsognathus longipes specimen

Compsognathids are considered to be among the smallest dinosaurs ever discovered. Compsognathus longipes was formerly the smallest known dinosaur. It was around the size of a chicken when fully grown: around 1 m (3 ft 3 in) long and weighing 2.5 kg (5.5 lb). [23] However, recently discovered adult specimens of other dinosaurs are smaller than Compsognathus, including Caenagnathasia , Microraptor and Parvicursor , all of which are estimated to be less than 1 m long. [24] However, most of these specimens are incomplete, so these sizes remain estimates.

The other genera in this family are slightly larger than Compsognathus longipes, but generally similar in size. The largest compsognathid is Huaxiagnathus, which is estimated from its holotype to be around 1.8 m (5 ft 11 in) long, [25] while Sinocalliopteryx measures around 2.4 m (7 ft 10 in) long. [26] Sinosauropteryx is the most similar to Compsognathus, measuring at most 1.07 m (3 ft 6 in) long. [27]

Feathers

Artistic recreation of Sinosauropteryx with feathers Sinosauropteryx color.jpg
Artistic recreation of Sinosauropteryx with feathers

The phylogeny of Compsognathidae organizes this family near the development of feathers in dinosaurs. In 1998, evidence of filamentous protofeathers was presented in a study on Sinosauropteryx, marking the first time that any sort of feather structure was found outside of birds and their related species. [28] After this, more evidence of feather structure was found in other genera of Compsognathidae. Evidence of protofeathers bearing resemblance to Sinosauropteryx was found on Sinocalliopteryx specimens, including on the foot of the specimen. [29] There have been signs of basic feather structures on Juravenator, but evidence of this is still not definite. Samples of Juravenator skin show scales instead of feathers, leading into debates about Juravenator’s place within the family Compsognathidae. [30] However, a 2010 examination of Juravenator under UV light showed filaments similar to those seen on other compsognathid specimens, indicating that it is likely that these dinosaurs had some sort of feathering. [31] A 2020 study concluded the "scales" were actually adipocere, though the same study defended Juravenator was a megalosauroid and not a compsognathid. [32]

Metacarpals

Another way of classification of Compsognathidae is shared metacarpal morphology. A 2007 study found similarities between compsognathid genera in certain metacarpal I morphologies. The conclusion of this study found that Composgnathidae had a distinct manual morphology where, like theropods, the first digit of the manus is larger than the other digits, but with a distinct metacarpal I morphology where the metacarpal is stocky and short. Compsognathidae also has a projection from the manus that is on this metacarpal. [12]

Classification

The Compsognathidae are a group of mostly small dinosaurs from the late Jurassic and early Cretaceous periods of China, Europe and South America. [33] For many years, Compsognathus was the only member known, but in recent decades paleontologists have discovered several related genera. The clade includes Aristosuchus , [34] Huaxiagnathus , [35] Mirischia , [36] Sinosauropteryx , [37] [38] and perhaps Juravenator [39] and Scipionyx . [40] At one time, Mononykus was proposed as a member of the family, but this was rejected by Chen and co-authors in a 1998 paper; they considered the similarities between Mononykus and the compsognathids to be an example of convergent evolution. [41] The position of Compsognathidae within the coelurosaur group is uncertain. Some, such as theropod expert Thomas Holtz Jr. and co-authors Ralph Molnar and Phil Currie in the landmark 2004 text Dinosauria, hold the family as the most basal of the coelurosaurs, [42] while others as part of the Maniraptora. [43] [44]

Below is a simplified cladogram showing Compsognathidae by Senter et al. in 2012. [45]

Coelurosauria

A number of authors have suggested that Compsognathidae is not a monophyletic group as currently defined, and that at least some "compsognathids" represent the juveniles of other tetanurans, including carnosaurs and tyrannosaurs. [14] [15] [16]

Here is a simplified version of Cau (2024), which does recover Compsognathidae as a polyphyletic group. Putative compsognathid specimens are in bold.

Tetanurae

Palaeobiology

Diet

Compsognathids were carnivores and certain specimens have contained the remains of their diet. The German specimen of Compsognathus included remains in the digestive region, which was initially thought to be an unborn embryo. [46] However, further analysis found that the remains belong to a lizard with an elongated tail and stretched legs. [20] [47] Other compsognathids, such as Sinosauropteryx, have been shown to eat lizards. [27]

Related Research Articles

<span class="mw-page-title-main">Theropoda</span> Clade of dinosaurs

Theropoda, whose members are known as theropods, is an extant dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally carnivorous, although a number of theropod groups evolved to become herbivores and omnivores. Theropods first appeared during the Carnian age of the late Triassic period 231.4 million years ago (Ma) and included the majority of large terrestrial carnivores from the Early Jurassic until at least the close of the Cretaceous, about 66 Ma. In the Jurassic, birds evolved from small specialized coelurosaurian theropods, and are today represented by about 11,000 living species.

<i>Compsognathus</i> Genus of dinosaurs

Compsognathus is a genus of small, bipedal, carnivorous theropod dinosaur. Members of its single species Compsognathus longipes could grow to around the size of a chicken. They lived about 150 million years ago, during the Tithonian age of the late Jurassic period, in what is now Europe. Paleontologists have found two well-preserved fossils, one in Germany in the 1850s and the second in France more than a century later. Today, C. longipes is the only recognized species, although the larger specimen discovered in France in the 1970s was once thought to belong to a separate species and named C. corallestris.

Coeluridae is a historically unnatural group of generally small, carnivorous dinosaurs from the late Jurassic Period. For many years, any small Jurassic or Cretaceous theropod that did not belong to one of the more specialized families recognized at the time was classified with the coelurids, creating a confusing array of 'coelurid' theropods that were not closely related. Although they have been traditionally included in this family, there is no evidence that any of these primitive coelurosaurs form a natural group with Coelurus, the namesake of Coeluridae, to the exclusion of other traditional coelurosaur groups.

<i>Sinosauropteryx</i> Extinct genus of dinosaurs

Sinosauropteryx is a compsognathid dinosaur. Described in 1996, it was the first dinosaur taxon outside of Avialae to be found with evidence of feathers. It was covered with a coat of very simple filament-like feathers. Structures that indicate colouration have also been preserved in some of its feathers, which makes Sinosauropteryx the first non-avialian dinosaurs where colouration has been determined. The colouration includes a reddish and light banded tail. Some contention has arisen with an alternative interpretation of the filamentous impression as remains of collagen fibres, but this has not been widely accepted.

<span class="mw-page-title-main">Coelurosauria</span> Clade of dinosaurs

Coelurosauria is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs.

<i>Beipiaosaurus</i> Extinct genus of dinosaurs

Beipiaosaurus is a genus of therizinosauroid theropod dinosaurs that lived in China during the Early Cretaceous in the Yixian Formation. The first remains were found in 1996 and formally described in 1999. Before the discovery of Yutyrannus, Beipiaosaurus were among the heaviest dinosaurs known from direct evidence to be feathered. Beipiaosaurus is known from three reported specimens. Numerous impressions of feather structures were preserved that allowed researchers to determine the feathering color which turned out to be brownish.

<span class="mw-page-title-main">Tetanurae</span> Clade containing most theropod dinosaurs

Tetanurae is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, and coelurosaurs. Tetanurans are defined as all theropods more closely related to modern birds than to Ceratosaurus and contain the majority of predatory dinosaur diversity. Tetanurae likely diverged from its sister group, Ceratosauria, during the late Triassic. Tetanurae first appeared in the fossil record by the Early Jurassic about 190 mya and by the Middle Jurassic had become globally distributed.

<span class="mw-page-title-main">Ornithomimosauria</span> Extinct clade of theropod dinosaurs

Ornithomimosauria are theropod dinosaurs which bore a superficial resemblance to the modern-day ostrich. They were fast, omnivorous or herbivorous dinosaurs from the Cretaceous Period of Laurasia, as well as Africa and possibly Australia. The group first appeared in the Early Cretaceous and persisted until the Late Cretaceous. Primitive members of the group include Nqwebasaurus, Pelecanimimus, Shenzhousaurus, Hexing and Deinocheirus, the arms of which reached 2.4 m (8 feet) in length. More advanced species, members of the family Ornithomimidae, include Gallimimus, Struthiomimus, and Ornithomimus. Some paleontologists, like Paul Sereno, consider the enigmatic alvarezsaurids to be close relatives of the ornithomimosaurs and place them together in the superfamily Ornithomimoidea.

<i>Ornithomimus</i> Ornithomimid dinosaur genus from the Late Cretaceous Period

Ornithomimus is a genus of ornithomimid theropod dinosaurs from the Campanian and Maastrichtian ages of Late Cretaceous Western North America. Ornithomimus was a swift, bipedal dinosaur which fossil evidence indicates was covered in feathers and equipped with a small toothless beak that may indicate an omnivorous diet. It is usually classified into two species: the type species, Ornithomimus velox, and a referred species, Ornithomimus edmontonicus. O. velox was named in 1890 by Othniel Charles Marsh on the basis of a foot and partial hand from the Denver Formation of Colorado. Another seventeen species have been named since then, though almost all of them have been subsequently assigned to new genera or shown to be not directly related to Ornithomimus velox. The best material of species still considered part of the genus has been found in Alberta, representing the species O. edmontonicus, known from several skeletons from the Horseshoe Canyon Formation. Additional species and specimens from other formations are sometimes classified as Ornithomimus, such as Ornithomimus samueli from the earlier Dinosaur Park Formation.

<i>Yixianosaurus</i> Extinct genus of dinosaurs

Yixianosaurus is a maniraptoran theropod dinosaur genus from the Early Cretaceous of China.

<span class="mw-page-title-main">Tyrannosauroidea</span> Extinct superfamily of dinosaurs

Tyrannosauroidea is a superfamily of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent beginning in the Jurassic Period. By the end of the Cretaceous Period, tyrannosauroids were the dominant large predators in the Northern Hemisphere, culminating in the gigantic Tyrannosaurus. Fossils of tyrannosauroids have been recovered on what are now the continents of North America, Europe and Asia, with fragmentary remains possibly attributable to tyrannosaurs also known from South America and Australia.

<i>Juravenator</i> Extinct genus of dinosaurs

Juravenator is a genus of small coelurosaurian theropod dinosaur, which lived about 151 or 152 million years ago in the area that would someday become the top of the Franconian Jura of Germany. It is known from a single, juvenile specimen.

<i>Huaxiagnathus</i> Extinct genus of dinosaurs

Huaxiagnathus is a genus of theropod dinosaur from the Lower Cretaceous of China. It was a compsognathid, large for that group at about half a meter longer than Compsognathus and larger specimens of Sinosauropteryx, with the largest specimen about 1.8 meters in length.

<i>Sinocalliopteryx</i> Extinct genus of dinosaurs

Sinocalliopteryx is a genus of carnivorous compsognathid theropod dinosaurs from the Lower Cretaceous Yixian Formation of China.

<i>Concavenator</i> Carcharodontosaurid dinosaur genus from the Early Cretaceous period

Concavenator is a genus of carcharodontosaurid dinosaur that lived in Spain during the Early Cretaceous epoch, about 125 million years ago. The genus contains a single species, Concavenator corcovatus named and described in 2010 from a nearly complete skeleton collected from Las Hoyas fossil site of La Huérguina Formation.

<i>Sciurumimus</i> Extinct species of reptile

Sciurumimus is an extinct genus of tetanuran theropod from the Late Jurassic Torleite Formation of Germany. It is known from a single juvenile specimen representing the type species, Sciurumimus albersdoerferi, which was found in a limestone quarry close to Painten in Lower Bavaria. The specimen was preserved with traces of feather-like filaments.

Aorun is a genus of carnivorous theropod dinosaur first discovered in 2006, with its scientific description published in 2013. It is possibly one of the oldest known coelurosaurian dinosaurs and is estimated to have lived ~161.6 million years ago during the Late Jurassic Period, though some researchers consider it to be a carnosaurian instead.

The Painten Formation is a geologic formation in Germany. It preserves fossils dating back to the Tithonian stage of the Late Jurassic period.

<span class="mw-page-title-main">Anchiornithidae</span> Extinct family of dinosaurs

Anchiornithidae is a family of small paravian dinosaurs. Anchiornithids have been classified at varying positions in the paravian tree, with some scientists classifying them as a distinct family, a basal subfamily of Troodontidae, members of Archaeopterygidae, or an assemblage of dinosaurs that are an evolutionary grade within Avialae or Paraves.

References

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  2. ↑ Hartman, Scott; Mortimer, Mickey; Wahl, William R.; Lomax, Dean R.; Lippincott, Jessica; Lovelace, David M. (2019). "A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight". PeerJ. 7: e7247. doi: 10.7717/peerj.7247 . PMC   6626525 . PMID   31333906.
  3. ↑ J. N. Choiniere, J. M. Clark, C. A. Forster and X. Xu. 2010. A basal coelurosaur (Dinosauria: Theropoda) from the Late Jurassic (Oxfordian) of the Shishugou Formation in Wucaiwan, People's Republic of China. Journal of Vertebrate Paleontology 30(6):1773-1796
  4. ↑ Etnier, Michael A. "Neptune's Ark: From Ichthyosaurs to Orcas." (2008): 25.
  5. 1 2 Xu, Xing. "Palaeontology: Scales, feathers and dinosaurs." Nature 440.7082 (2006): 287-288.
  6. 1 2 3 4 Peyer, Karin. "A reconsideration of Compsognathus from the Upper Tithonian of Canjuers, southeastern France." Journal of vertebrate Paleontology 26.4 (2006): 879-896.
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  9. 1 2 3 Naish, D.; Martill, David M.; Frey, Eberhard (2004). "Ecology, systematics and biogeographical relationships of dinosaurs, including a new theropod, from the Santana Formation (?Albian, Early Cretaceous) of Brazil" (PDF). Historical Biology. 16 (2–4): 57–70. CiteSeerX   10.1.1.394.9219 . doi:10.1080/08912960410001674200. S2CID   18592288.
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  11. ↑ Dal Sasso C., Maganuco S. "Scipionyx samniticus (Theropoda: Compsognathidae) from the Lower Cretaceous of Italy: osteology, ontogenetic assessment, phylogeny, soft tissue anatomy, taphonomy and palaeobiology." Memorie, XXXVI-I (2011): 1-283.
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  30. ↑ Goehlich, U.B.; Tischlinger, H.; Chiappe, L.M. (2006). "Juravenator starki (Reptilia, Theropoda) ein neuer Raubdinosaurier aus dem Oberjura der Suedlichen Frankenalb (Sueddeutschland): Skelettanatomie und Weichteilbefunde". Archaeopteryx. 24: 1–26.
  31. ↑ Chiappe, L.M.; Göhlich, U.B. (2010). "Anatomy of Juravenator starki (Theropoda: Coelurosauria) from the Late Jurassic of Germany". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 258 (3): 257–296. doi : 10.1127/0077-7749/2010/0125.
  32. ↑ Foth, Christian; Haug, Carolin; Haug, Joachim T.; Tischlinger, Helmut; Rauhut, Oliver W. M. (2020), Foth, Christian; Rauhut, Oliver W. M. (eds.), "Two of a Feather: A Comparison of the Preserved Integument in the Juvenile Theropod Dinosaurs Sciurumimus and Juravenator from the Kimmeridgian Torleite Formation of Southern Germany", The Evolution of Feathers: From Their Origin to the Present, Fascinating Life Sciences, Cham: Springer International Publishing, pp. 79–101, doi:10.1007/978-3-030-27223-4_6, ISBN   978-3-030-27223-4, S2CID   216245045 , retrieved 2021-03-08
  33. ↑ Peyer, K. (2006). "A reconsideration of Compsognathus from the Upper Tithonian of Canjuers, southeastern France". Journal of Vertebrate Paleontology. 26 (4): 879–896. doi:10.1671/0272-4634(2006)26[879:AROCFT]2.0.CO;2. S2CID   86338296.
  34. ↑ Seeley, H.G. (1887). "On Aristosuchus pusillus (Owen), being further notes on the fossils described by Sir. R. Owen as Poikilopleuron pusillus, Owen". Quarterly Journal of the Geological Society of London. 43 (1–4): 221–228. doi:10.1144/GSL.JGS.1887.043.01-04.22. S2CID   131237500.
  35. ↑ Hwang, S. H.; Norell, M. A.; Qiang, J.; Keqin, G. (2004). "A large compsognathid from the Early Cretaceous Yixian Formation of China" (PDF). Journal of Systematic Palaeontology. 2: 13–39. doi:10.1017/S1477201903001081. S2CID   86321886.
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