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]
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. [5] Compsognathid fossils preserve diverse integument — skin impressions are known from four genera commonly placed in the group, Compsognathus , Sinosauropteryx , Sinocalliopteryx , and Juravenator . [6] 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.

The first member of the group, Compsognathus , was discovered in 1861, after Johann A. Wagner published his description of the taxon. [7] [8] [9] The family was created by Edward Drinker Cope in 1875. [10] 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. [7] With further discoveries, fossils have been uncovered across three different continents, in Asia, [11] [7] [8] [9] [6] Europe, [12] and South America. [8] [10] Assignment to Compsognathidae is usually determined through examination of the metacarpal, which is used to separate Compsognathidae from other dinosaurs. [13] 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. [10] [14] 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. [15] [16] [17]

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. [18] 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"). [19] 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. [20] 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. [21]

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. [7] 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. [22] 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 due to the fact that Juravenator could also be classified into a similar group within Coelurosauria, Maniraptoriformes. Maniraptoriforms share many similarities with compsognathids and due to the fact that 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. [23]

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). [24] 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. [25] 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, [26] while Sinocalliopteryx measures around 2.4 m (7 ft 10 in) long. [27] Sinosauropteryx is the most similar to Compsognathus, measuring at most 1.07 m (3 ft 6 in) long. [28]

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. [29] 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. [30] 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. [31] 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. [32] A 2020 study concluded the "scales" were actually adipocere, though the same study defended Juravenator was a megalosauroid and not a compsognathid. [33]

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. [13]

Classification

The Compsognathidae are a group of mostly small dinosaurs from the late Jurassic and early Cretaceous periods of China, Europe and South America. [34] For many years, Compsognathus was the only member known, but in recent decades paleontologists have discovered several related genera. The clade includes Aristosuchus , [35] Huaxiagnathus , [36] Mirischia , [37] Sinosauropteryx , [38] [39] and perhaps Juravenator [40] and Scipionyx . [41] At one time, Mononykus was proposed as a member of the family, but this was rejected by Chen and coauthors in a 1998 paper; they considered the similarities between Mononykus and the compsognathids to be an example of convergent evolution. [42] 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, [43] while others as part of the Maniraptora. [44] [45]

Below is a simplified cladogram showing Compsognathidae by Senter et al. in 2012. [46]

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. [15] [16] [17]

Here is a simplified version of Cau (2024), which does recovers 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. [47] However, further analysis found that the remains belong to a lizard with an elongated tail and stretched legs. [21] [48] Other compsognathids, such as Sinosauropteryx, have been shown to eat lizards. [28]

Related Research Articles

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

Theropoda, whose members are known as theropods, is a 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 10,500 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.

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

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.

<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, tyrannosauroids, ornithomimosaurs, compsognathids and maniraptorans. 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.

<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.

Coelurus is a genus of coelurosaurian dinosaur from the Late Jurassic period. The name means "hollow tail", referring to its hollow tail vertebrae. Although its name is linked to one of the main divisions of theropods (Coelurosauria), it has historically been poorly understood, and sometimes confused with its better-known contemporary Ornitholestes. Like many dinosaurs studied in the early years of paleontology, it has had a confusing taxonomic history, with several species being named and later transferred to other genera or abandoned. Only one species is currently recognized as valid: the type species, C. fragilis, described by Othniel Charles Marsh in 1879. It is known from one partial skeleton found in the Morrison Formation of Wyoming, United States. It was a small bipedal carnivore with elongate legs.

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

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

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

Maniraptoriformes is a clade of dinosaurs with pennaceous feathers and wings that contains ornithomimosaurs and maniraptorans. This group was named by Thomas Holtz, who defined it as "the most recent common ancestor of Ornithomimus and birds, and all descendants of that common ancestor."

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

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

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

Proceratosauridae is a family or clade of tyrannosauroid theropod dinosaurs from the Middle Jurassic to the Early Cretaceous.

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

Haplocheirus is an extinct genus of theropod dinosaur from the Middle Jurassic Shishugou Formation of Xinjiang in China. It is generally considered to be an alvarezsauroid, although some researchers have questioned this assignment. The genus contains a single species, H. sollers, which is known from a mostly complete skeleton including the skull.

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

Concavenator is an extinct carcharodontosaurid theropod dinosaur that lived approximately 130 million years ago during the Early Cretaceous period. The type species is C. corcovatus. Concavenator corcovatus means "Cuenca hunter with a hump". The fossil was discovered in the Las Hoyas fossil site of Spain by paleontologists José Luis Sanz, Francisco Ortega, and Fernando Escaso from the Autonomous University of Madrid and the National University of Distance Education.

<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.

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

Aorun is a genus of carnivorous theropod dinosaur first discovered in 2006, with its scientific description published in 2013. It is one of the oldest known coelurosaurian dinosaurs and is estimated to have lived ~161.6 million years ago during the Late Jurassic Period. It is the fifth extinct theropod discovered from Wucaiwan, China.

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.

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