Compsognathus

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Compsognathus (Template:IPAc-en;<ref>Template:Cite OED</ref> Greek kompsos/κομψός; "elegant", "refined" or "dainty", and gnathos/γνάθος; "jaw")<ref>Template:Cite book</ref> 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 French specimen was once thought to belong to a separate species named C. corallestris.

Many presentations still describe Compsognathus as "chicken-sized" dinosaurs because of the size of the German specimen, which is now believed to be a juvenile. Compsognathus longipes is one of the few dinosaur species whose diet is known with certainty: the remains of small, agile lizards are preserved in the bellies of both specimens. Teeth discovered in Portugal may be further fossil remains of the genus.

Although not recognized as such at the time of its discovery, Compsognathus is the first theropod dinosaur known from a reasonably complete fossil skeleton. Until the 1990s, it was the smallest-known non-avialan dinosaur, with the preceding centuries incorrectly labelling them as the closest relative of Archaeopteryx.

Discovery and speciesEdit

File:Compsognathus longipes cast 3.jpg
Joseph Oberndorfer acquired this fossil in Bavaria, Germany, in 1859. Shown here is a cast at the Bavarian State Institute for Paleontology and Historical Geology

Compsognathus is known from two almost complete skeletons.<ref name=GSP88>Template:Cite book</ref> The German specimen (specimen number BSP AS I 563) stems from limestone deposits in Bavaria and was part of the collection of physician and fossil collector Joseph Oberndorfer. Oberndorfer lent the specimen to paleontologist Johann A. Wagner, who published a brief discussion in 1859, where he coined the name Compsognathus longipes.<ref name=wagner1859>Template:Cite journal</ref> Wagner did not recognise Compsognathus as a dinosaur, but instead described it as one of the "most curious forms among the lizards".<ref name=wagner1859/><ref name=ostrom78/> He published a more detailed description in 1861.<ref>Template:Cite journal</ref> In 1866, Oberndorfer's collection, including the Compsognathus specimen, was acquired by the paleontological state collection in Munich.<ref name="goerlich2006"/>

The year of discovery and the exact locality of the German specimen are unknown, possibly because Oberndorfer did not reveal details of the discovery to prevent other collectors from exploiting the locality;<ref name="goerlich2006">Template:Cite journal</ref> later authors have suggested that the German specimen was probably discovered during the 1850s. Weathering of the slab on which the fossil is preserved indicates that it was collected from a pile of waste rock left behind by quarrying.<ref name=reisdorf2012/> The specimen either stems from Jachenhausen or the region Riedenburg–Kehlheim.<ref name=reisdorf2012/><ref name="mauser1983">Template:Cite journal</ref> All possible localities are part of lagoonal deposits of the Painten Formation, and date to the latest part of the late Kimmeridgian or the earlier part of the early Tithonian.<ref name=reisdorf2012/> In the Jurassic, the region was part of the Solnhofen archipelago. The limestone of the area, the Solnhofen limestone, had been quarried for centuries, and yielded such well-preserved fossils as Archaeopteryx with feather impressions and pterosaurs with imprints of their wing membranes.<ref name=Wellnhofer2008>Template:Cite book</ref>

In two publications in 1868 and 1870, Thomas Huxley, a major proponent of Charles Darwin's theory of evolution, compared Compsognathus with Archaeopteryx, which was considered the earliest known bird. Following earlier suggestions by Carl Gegenbaur<ref name="gegenbaur1863">Template:Cite journal</ref> and Edward Drinker Cope,<ref name="cope1867">Template:Cite journal</ref> Huxley found that Archaeopteryx was closely similar to Compsognathus, and referred to the latter as a "bird-like reptile". He concluded that birds must have evolved from dinosaurs, an assessment that established Compsognathus as one of the most widely known dinosaurs.<ref name="huxley1868">Template:Cite journal</ref><ref name="huxley1870b">Template:Cite wikisource</ref><ref name=ostrom78/> The specimen has since been studied by many prominent paleontologists, including Othniel Charles Marsh, who visited Munich in 1881. The German paleontologist J.G. Baur, who worked as an assistant of Marsh, removed the right ankle from the slab for illustration and study; this removed part got lost since.<ref name=ostrom78/><ref name="ostrom_discovery">Template:Cite journal</ref> Although Baur published a detailed study of the ankle in 1882, which is now the only available source of information of this part of the skeleton, his reconstruction was later found to be inconsistent with corresponding impressions on the slab.<ref name=ostrom78/> John Ostrom thoroughly described the German specimen as well as the newly discovered French specimen in 1978, making Compsognathus one of the best-known small theropods at that time.<ref name=ostrom78/> He also concluded that the German specimen likely belongs to an immature individual.<ref name=ostrom78/>

File:Compsognathus corallestris = longipes.JPG
The fossil from Canjuers, France

The larger French specimen (Y85R M4M) was discovered in around 1971 in the Portlandian lithographic limestone of Canjuers near Nice.<ref name=peyer06/> It dates to the lower Tithonian, as indicated by ammonite index fossils. As Solnhofen, Canjures was famous for its limestone plates, which were quarried and sold under the name "dalles de Provence". The specimen was originally part of a large private fossil collection of Louis Ghirardi, the owner of the Canjures quarries. The collection, including the Compsognathus specimen, was sold to the National Museum of Natural History in Paris in 1983. Alain Bidar and Gérard Thomel, in a brief 1972 description, announced the new find under a separate species, Compsognathus corallestris.<ref name="bidar1972a">Template:Cite journal</ref> A more comprehensive description followed in the same year.<ref name="Bidar72">Template:Cite journal</ref> According to these authors, the new species differed from the German species in its larger size and modified, flipper-like hand. Ostrom, Jean-Guy Michard and others have since relabeled it as another example of Compsognathus longipes.<ref name=peyer06/><ref name="Michard91">Template:Cite journal</ref> In 1984, George Callison and Helen Quimby identified the smaller German specimen as a juvenile of the same species.<ref name="CallisonQuimby1984">Template:Cite journal</ref><ref name="Michard91"/>

Collector Heinrich Fischer had originally labeled a partial foot consisting of three metatarsals and a phalanx, from the Solnhofen area, as belonging to Compsognathus longipes. This identification was rejected by Wilhelm Dames, when he described the specimen for the first time in 1884.<ref name="dames1884">Template:Cite journal</ref> Friedrich von Huene, in 1925 and 1932, also found that the foot did probably not belong to Compsognathus itself but to a closely related genus.<ref name="huene1925">Template:Cite journal</ref><ref name="huene1932">Template:Cite journal</ref>Template:Rp Ostrom, in his 1978 monography, questioned the attribution of this fossil to Compsognathus once more.<ref name=ostrom78/> Jens Zinke, in 1998, assigned forty-nine isolated teeth from the Guimarota coal mine of Portugal to the genus. Zinke found that these teeth are not identical to those of Compsognathus longipes, having serrations on the front edge, and thus labeled the teeth as Compsognathus sp. (of unknown species).<ref>Template:Cite journal</ref>

DescriptionEdit

File:Compysizes1.png
Size comparison of the French (orange) and German (green) specimens, with a human

For decades, Compsognathus was known as the smallest known non-avian dinosaur,<ref name=ostrom78/> although some dinosaurs discovered later, such as Mahakala and Microraptor, were even smaller.<ref name="campione2014">Template:Cite journal</ref><ref name="agnolin2019">Template:Cite journal</ref> The German specimen was estimated to be Template:Convert<ref name=ostrom78/> and Template:Convert<ref name=GSP88/> in length by separate authors, while the larger French specimen was estimated at Template:Convert<ref name=GSP88/> and Template:Convert<ref name=peyer06/> in length. The height at the hip has been estimated at Template:Convert for the German specimen and at Template:Convert for the French specimen.<ref name=GSP88/> The German specimen was estimated to have weighed Template:Convert<ref name=TH07>Template:Cite journal</ref> and Template:Convert,<ref name=GSP88/> and the French specimen Template:Convert<ref name=GSP88/> and Template:Convert.<ref name="seebacher2001">Template:Cite journal</ref> Compared to other compsognathids, the larger French specimen would have been similar in size to larger Sinosauropteryx specimens, but smaller than Huaxiagnathus and Mirischia.<ref name=peyer06/>

Compsognathus were small, bipedal animals with long legs and longer tails, which they used for balance during locomotion. The forelimbs were smaller than the hindlimbs. The hand bore two large, clawed digits and a third, smaller digit that may have been non-functional.<ref name=peyer06/><ref name=gishlick07/> Their delicate skulls were narrow and long, with tapered snouts. The skull had five pairs of fenestrae (skull openings), the largest of which was for the orbit (eye socket),<ref name="Lambert">Template:Cite book</ref> with the eyes being larger in proportion to the rest of the skull. The lower jaw was slender and had no mandibular fenestra, a hole in the side of the lower jawbone commonly seen in archosaurs.

The teeth were small and pointed, suited for its diet of small vertebrates and possibly other small animals, such as insects. The German specimen had three teeth in each premaxilla (front bone of the lower jaw), 15 or 16 teeth in each maxilla, and 18 teeth in the lower jaw.<ref name="stromer1934">Template:Cite journal</ref><ref name=ostrom78/> The French specimen had more teeth, including four in each premaxilla, 17 or 18 in the maxilla, and at least 21 teeth in the dentary.<ref name=peyer06/> Compsognathids were unique among theropods in having tooth crowns that curved backwards at two thirds of their height, while their mid-parts were straight; also, the crowns had expanded bases.<ref name=peyer06/> In Compsognathus, the frontmost teeth of the upper and lower jaws were unserrated, while those further back had fine serrations on their rear edges. In the German specimen, the crowns were around two times higher than wide in the front of the jaws but diminished in height further back, with the last tooth about as high as wide.<ref name="stromer1934"/> The German specimen also shows a diastema (tooth gap) behind the first three teeth of the premaxilla.<ref name=ostrom78/> As such a gap was not present in the French specimen, Peyer suggested that additional teeth were possibly present in this region the German specimen.<ref name=peyer06/>

The number of digits on the hand of Compsognathus has been a source of debate.<ref name=gishlick07/> For much of its history, Compsognathus was typically depicted with three digits, as is typical for theropods.<ref name=gishlick07/><ref name=ostrom78/> However, the type specimen only preserved phalanges from the first two digits, leading to the suggestion that Compsognathus had only two functional digits, with the third metacarpal being smaller and thinner.<ref name=ostrom78/> Study of the French specimen indicated that the third digit bore at least one or two small phalanges.<ref name=peyer06/><ref name=gishlick07/> However, there remains no evidence for an ungual phalanx on the third digit, so the digit may have been reduced and non-functional.<ref name=gishlick07/>

IntegumentEdit

File:Compsognathus BW.jpg
Evidence from related species suggests that the body might have been covered with feather-like structures.

Some relatives of Compsognathus, namely Sinosauropteryx and Sinocalliopteryx, have been preserved with the remains of simple feathers covering the body like fur,<ref name="Currie2001"/> prompting some scientists to suggest that Compsognathus might have been feathered in a similar way.<ref name="jietal2007">Template:Cite journal</ref> Consequently, many depictions of Compsognathus show them with coverings of downy proto-feathers. However, no feathers or feather-like covering have been preserved with Compsognathus fossils, in contrast to Archaeopteryx, which are found in the same sediments. Karin Peyer, in 2006, reported skin impressions preserved on the side of the tail starting at the 13th tail vertebra. The impressions showed small bumpy tubercles, similar to the scales found on the tail and hind legs of Juravenator.<ref name=peyer2006>Template:Cite journal</ref> Additional scales had in 1901 been reported by Von Huene, in the abdominal region of the German Compsognathus, but Ostrom subsequently disproved this interpretation;<ref name=ostrom78/><ref name="Huene01">Template:Cite journal</ref> in 2012, Achim Reisdorf postulated that they are plaques of adipocere, corpse wax.<ref name=reisdorf2012>Template:Cite journal</ref>

Like Compsognathus, and unlike Sinosauropteryx, a patch of fossilized skin from the tail and hindlimb of the possible relative Juravenator starki shows mainly scales, though there is some indication that simple feathers were also present in the preserved areas.<ref name=Goehlich2006>Template:Cite journal</ref> This may mean that a feather covering was not ubiquitous in this group of dinosaurs, or maybe that some species had fewer feathers than others.<ref>Template:Cite journal</ref>

ClassificationEdit

Originally classified as a lizard, the dinosaurian affinities of Compsognathus were first noted by Gegenbaur, Cope, and Huxley between 1863 and 1868.<ref name="gegenbaur1863"/><ref name="cope1867"/><ref name="huxley1868"/> Cope, in 1870, classified Compsognathus within a new clade of dinosaurs, the Symphypoda, which also contained Ornithotarsus (today classified as Hadrosaurus).<ref name="cope1870">Template:Cite journal</ref><ref name="baur1891">Template:Cite journal</ref> Later, both genera were found to belong to other groups of Cope's classification of dinosaurs: Compsognathus to the Gonipoda (equivalent to Theropoda, in which it is now classified), and Ornithotarsus to the Orthopoda (equivalent to Ornithischia).<ref name="baur1891"/> Huxley, in 1870, rejected Cope's dinosaur classification scheme, and instead proposed the new clade Ornithoscelida, in which he included the Dinosauria (comprising several forms now considered as ornithischians) and another new clade, the Compsognatha, which contained Compsognathus as the only member.<ref name="huxley1870">Template:Cite wikisource</ref><ref name="seeley1888">Template:Cite journal</ref> Later, these groups fell into disuse, although a resurrection of the Ornithoscelida was proposed in 2017.<ref name="baron2017">Template:Cite journal</ref> The group Compsognatha was used for the last time by Marsh in a 1896 publication, where it was treated as a suborder of Theropoda.<ref name="marsh1996">Template:Cite journal</ref><ref name=peyer06/> In the same publication, Marsh erected the new family Compsognathidae.<ref name="marsh1996"/><ref name=peyer06/> Friedrich von Huene, in 1914, erected the new infraorder Coelurosauria, which includes the Compsognathidae amongst other families of small theropods; this classification remained in use since.<ref name="huene1914">Template:Cite journal</ref><ref name=peyer06/>

The Compsognathidae are a group of mostly small dinosaurs from the late Jurassic and early Cretaceous periods of China, Europe and South America.<ref name=peyer06/> For many years, Compsognathus was the only member known, but in recent decades paleontologists have discovered several related genera. The clade includes Aristosuchus,<ref>Template:Cite journal</ref> Huaxiagnathus,<ref>Template:Cite journal</ref> Mirischia,<ref>Template:Cite journal</ref> Sinosauropteryx,<ref name="Currie2001">Template:Cite journal</ref><ref name="Ji1996">Template:Cite journal</ref> and perhaps Juravenator and Scipionyx.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> 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.<ref name="Chen">Template:Cite journal</ref> The position of Compsognathus and its relatives 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,<ref name="dinosauria04">Template:Cite book</ref> while others as part of the Maniraptora.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>

For almost a century, Compsognathus longipes was the only well-known small theropod species. This led to comparisons with Archaeopteryx and to suggestions of an especially close relationship with birds. In fact, Compsognathus, rather than Archaeopteryx, piqued Huxley's interest in the origin of birds.<ref name = "FasWeis04">Template:Cite book</ref> The two animals share similarities in shape and proportions, so many in fact that two specimens of Archaeopteryx, the "Eichstätt" and the "Solnhofen", were for a time misidentified as those of Compsognathus.<ref name="Lambert"/> Many other types of theropod dinosaurs, such as maniraptorans, are now known to have been more closely related to birds.<ref name=weishampel04/>

File:Compsognathus longipes.jpg
Skeletal reconstruction by Marsh, 1896
File:Compsognathidae.jpg
Diagrams showing known elements of the two specimens (middle) and other compsognathids

Below is a simplified cladogram placing Compsognathus in Compsognathidae by Senter et al. in 2012.<ref name=senter10>Template:Cite journal</ref>

Template:Clade

Here is an alternative phylogeny, published by Cau in 2024, with both specimens in bold. Template:Clade

PaleobiologyEdit

In a 2001 study conducted by Bruce Rothschild and other paleontologists, nine foot bones referred to Compsognathus were examined for signs of stress fracture, but none were found.<ref name="rothschild-dino">Template:Cite journal</ref>

HabitatEdit

Bidar and colleagues, in their 1972 description of the French specimen, argued that this specimen had webbed hands which would look like flippers in life. This interpretation was based on a supposed impression of the flipper that consists of several undulating wrinkles running parallel to the forelimb on the surface of the slab.<ref name="Bidar72"/> In a 1975 popular book, L. Beverly Halstead depicts the animal as an amphibious dinosaur capable of feeding on aquatic prey and swimming out of reach of larger predators.<ref>Template:Cite book</ref> Ostrom debunked this hypothesis, noting that the forelimb of the French specimen is poorly preserved, and that the wrinkles extend well beyond the skeleton and thus are likely sedimentary structures unrelated to the fossil.<ref name=ostrom78/><ref name=peyer06/>

DietEdit

File:Compsognathus by Nopcsa, 1903.jpg
This 1903 illustration by Franz Nopcsa von Felső-Szilvás shows the gastric content of the German specimen

The remains of a lizard in the German specimen's thoracic cavity show that Compsognathus preyed on small vertebrates.<ref name="Chen"/> Marsh, who examined the specimen in 1881, thought that this small skeleton in the Compsognathus belly was an embryo, but in 1903, Franz Nopcsa concluded that it was a lizard.<ref name="nopcsa1903">Template:Cite journal</ref> Ostrom identified the remains as belonging to a lizard of the genus Bavarisaurus,<ref>Template:Cite journal</ref> which he concluded was a fast and agile runner owing to its long tail and limb proportions. This in turn led to the conclusion that its predators, Compsognathus, must have had sharp vision and the ability to rapidly accelerate and outrun the lizard.<ref name=ostrom78/> Conrad made the lizard found in the thoracic cavity of the German specimen of Compsognathus the holotype of a new species Schoenesmahl dyspepsia.<ref name=":0">Template:Cite journal</ref> The lizard is in several pieces, indicating that the Compsognathus must have dismembered it while restraining it with its hands and teeth, and then swallowed the remains whole; a similar strategy is used by modern predatory birds.<ref name=":0" /> The French specimen's gastric contents consist of unidentified lizards or sphenodontids.<ref name=peyer06/>

Possible eggsEdit

The plate of the German Compsognathus shows several circular irregularities Template:Convert in diameter near the skeletal remains. Peter Griffiths interpreted them as immature eggs in 1993.<ref>Template:Cite journal</ref> However, later researchers have doubted their connection to the genus because they were found outside the body cavity of the animal. A well-preserved fossil of a Sinosauropteryx, a genus related to Compsognathus, shows two oviducts bearing two unlaid eggs. These proportionally larger and less numerous eggs of Sinosauropteryx cast further doubt on the original identification of the purported Compsognathus eggs.<ref name="Chen"/> In 1964 German geologist Karl Werner Barthel had explained the discs as gas bubbles formed in the sediment because of the putrefaction of the carcass.<ref>Template:Cite journal</ref>

SpeedEdit

In 2007, William Sellers and Phillip Manning estimated a maximum speed of Template:Convert based on a computer model of the skeleton and muscles.<ref>Template:Cite journal</ref> This estimate has been criticized by other scholars.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

PaleoenvironmentEdit

File:Archaeopteryx lithographica by durbed.jpg
Restoration of an Archaeopteryx chasing a juvenile Compsognathus

During the late Jurassic, Europe was a dry, tropical archipelago at the edge of the Tethys Sea. The fine limestone in which the skeletons of Compsognathus have been found originated in calcite from the shells of marine organisms. Both the German and French areas where Compsognathus specimens have been preserved were lagoons situated between the beaches and coral reefs of the Jurassic European islands in the Tethys Sea.<ref>Template:Cite book</ref> Contemporaries of Compsognathus longipes include the early avialan Archaeopteryx lithographica and the pterosaurs Rhamphorhynchus muensteri and Pterodactylus antiquus. The same sediments in which Compsognathus have been preserved also contain fossils of a number of marine animals such as fish, crustaceans, echinoderms and marine mollusks, confirming the coastal habitat of this theropod. No other dinosaur has been found in association with Compsognathus, indicating that these little dinosaurs might in fact have been the top land predator in these islands.<ref name=weishampel04>Template:Cite book</ref>

TaphonomyEdit

Much discussion revolved around the taphonomy of the German specimen, i.e. how the individual died and became fossilized. Reisdorf and Wuttke, in 2012, speculated about the events that lead to the death and transportation of the specimen to its place of burial. First, the individual must have been brought into the lagoon from its habitat, which probably was on the surrounding islands. It is possible that a flash flood swept the animal into the sea, in which case it likely died by drowning. It is also possible that the animal swam or drifted onto the sea, or that it rafted on plants, and was then transported by surface currents to its place of burial. In any case, the specimen would have arrived on the sea floor within a few hours after its death, as otherwise gases forming in its body cavity would have prevented it from sinking in one piece. Water depth at the burial site would have been large enough to prevent refloating of the carcass after such gases were produced. Rounded structures on the slab might have been formed by the release of these gases.<ref name=reisdorf2012/>

Taphonomic reconstructions are complicated as the exact locality and the position and orientation of the fossil within the sediments is no longer known.<ref name=reisdorf2012/> As a compression fossil, the specimen would originally have been preserved on both the upper surface of a layer and the lower surface of the subsequent layer (i.e., on a slab and its counter-slab); the counter-slab is now lost. Reisdorf and Wuttke, in 2012, argued that the front and hind limbs of the left side of the body were better Template:Dinogloss (still connected together) than those of the right side. This suggests that the specimen is located on the bottom side of the upper slab, and was lying on its left side.<ref name=reisdorf2012/> The German specimen was preserved with a high degree of articulation – only the skull, hands, cervical ribs and gastralia show disarticulation. The braincase was displaced behind the skull, the first tail vertebra was rotated by 90°, and the tail shows a break between the seventh and eighth tail vertebra.<ref name=reisdorf2012/>

File:German Compsognathus.jpg
Illustration of the German specimen

In both Compsognathus specimens, the neck is strongly curved, with the head coming to rest above the pelvis; the spine of the tail was likewise curved.<ref name="faux2007"/> This posture, known as the death pose, is found in many vertebrate fossils, and the German Compsognathus specimen was central in several studies that sought to explain this phenomenon. The physician Moodie, in 1918, suggested that the death pose in Compsognathus and similar fossils was the result of an opisthotonus – death throes causing spastic stiffening of the back musculature – while the animal was dying. This hypothesis was soon challenged by paleontologist Friedrich von Huene, who argued that the death pose was the result of desiccation and therefore occurred only after the death. Peter Wellnhofer, in 1991, argued that death poses resulted from the elastic pull of the ligaments, which are released after death.<ref name=reisdorf2012/> The veterinarian Cynthia Faux and the paleontologist Kevin Padian, in a 2007 study that gained much attention, supported the original opisthotonus hypothesis of Moodie. These authors furthermore argued that upon death, muscles are relaxed and body parts can be easily moved relative to each other. Since opisthotonic postures are already established during death, they may only be preserved if the animal dies in place and becomes buried rapidly. This contradicts previous interpretations on the environment and taphonomy of Compsognathus and other fossils from the Solnhofen limestones, which assumed very slow burial at the bottom of lagoons into which the carcasses were transported from nearby islands.<ref name="faux2007">Template:Cite journal</ref><ref name=reisdorf2012/> Reisdorf and Wuttke concluded that the death posture indeed resulted from the release of ligaments, more specifically the Template:Dinogloss, which spans the spine from the neck to tail in modern birds. The release of this ligament would have occurred gradually while the surrounding muscle tissue decayed, and only after the carcass was transported to its final site of deposition.<ref name=reisdorf2012/>

The bottom water of the lagoon was likely anaerobic (devoid in oxygen), resulting in a sea floor devoid of life except for microbial mats, and therefore preventing scavenging of the carcass.<ref name=reisdorf2012/> In the trunk region of the German specimen, the surface of the slab is markedly different in texture to the surrounding areas of the slab, showing irregular, nodular surfaces within depressions. Ostrom, in 1978, interpreted these structures as traces of weathering that took place just before the fossil was collected.<ref name=ostrom78/><ref name=reisdorf2012/> Nopcsa, in 1903, instead suggested that these structures resulted from decomposing tissue of the carcass.<ref name="nopcsa1903"/><ref name=reisdorf2012/> Reisdorf and Wuttke, in their 2012 study, suggested that the structures are the remains of adipocere (corpse wax formed by bacteria) that formed around the carcass before burial. Such adipocere would have helped in conserving the state of articulation of the fossil for years when burial was very slow. The presence of adipocere would possibly rule out hypersalinity (very high salt contents) of the bottom water, because such conditions appear to be unfavorable for the adipocere producing bacteria.<ref name=reisdorf2012/>

In popular cultureEdit

Compsognathus is one of the more popular dinosaurs.<ref name=ostrom78/> For a long time, it was considered unique in its small size, which is commonly compared to that of a chicken.<ref name=ostrom78/><ref name=RW86>Template:Cite book</ref><ref name=SA88>Template:Cite book</ref> The animals have appeared in the Jurassic Park franchise: in the films The Lost World: Jurassic Park, Jurassic Park III, Jurassic World: Fallen Kingdom and Jurassic World Dominion, and the television series Camp Cretaceous. In The Lost World: Jurassic Park, the species is incorrectly identified as Compsognathus triassicus, combining the genus name of Compsognathus longipes with the specific name of Procompsognathus triassicus, a distantly related small carnivore that also appears in the Jurassic Park series.<ref>Template:Cite book</ref>

ReferencesEdit

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External linksEdit

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