Ctenochasma Elegans
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''Ctenochasma'' (meaning "comb jaw") is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
ctenochasmatid Ctenochasmatidae is a group of pterosaurs within the suborder Pterodactyloidea. They are characterized by their distinctive teeth, which are thought to have been used for filter-feeding. Ctenochasmatids lived from the Late Jurassic to the Early ...
pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
belonging to the suborder
Pterodactyloidea Pterodactyloidea ( ; derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger") is one of the two traditional suborders of pterosaurs ("wing lizards"), and contai ...
. Three species are currently recognized: ''C. roemeri'' (named after
Friedrich Adolph Roemer Friedrich Adolph Roemer (15 April 1809 – 25 November 1869), German geologist, was born at Hildesheim, in the Kingdom of Westphalia. His father was a lawyer and councillor of the high court of justice. In 1845 he became professor of mineralogy ...
), ''C. taqueti'', and ''C. elegans''. Their fossilized remains have been found in the
Solnhofen Limestone The Solnhofen Limestone or Solnhofen Plattenkalk, formally known as the Altmühltal Formation, is a Jurassic Konservat-Lagerstätte that preserves a rare assemblage of fossilized organisms, including highly detailed imprints of soft bodied organi ...
of
Bavaria Bavaria, officially the Free State of Bavaria, is a States of Germany, state in the southeast of Germany. With an area of , it is the list of German states by area, largest German state by land area, comprising approximately 1/5 of the total l ...
,
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
, the "
Purbeck Group The Purbeck Group is an Upper Jurassic to Lower Cretaceous lithostratigraphy, lithostratigraphic group (stratigraphy), group (a sequence of rock strata) in south-east England. The name is derived from the district known as the Isle of Purbeck in ...
" of northeastern
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
, and the Calcaires tâchetés of eastern
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
.


History

The name ''Ctenochasma'' was coined by the German paleontologist
Christian Erich Hermann von Meyer Christian Erich Hermann von Meyer (3 September 1801 – 2 April 1869), known as Hermann von Meyer, was a German palaeontologist. He was awarded the 1858 Wollaston medal by the Geological Society of London. Life He was born in Frankfurt am ...
in 1851, based on a single lower jaw full of closely packed teeth which he gave the species name ''Ctenochasma roemeri''.von Meyer, C.E.H. (1851). "''Ctenochasma Roemeri''." ''Paläontographica'', 2: 82–84 & pl. 13. Von Meyer did not consider the specimen to belong to a pterosaur; instead, he compared it to ''
Gnathosaurus ''Gnathosaurus'' (meaning "jawed lizard") is a genus of ctenochasmatid pterosaur containing two species: ''G. subulatus'', named in 1833 from the Solnhofen Limestone of Germany, and ''G. macrurus'', known from the Purbeck Limestone of the UK. ...
'', which at that time was considered to be a crocodilian. A second species, ''C. gracile'', was named by Oppel in 1862 based on a fragmentary skull.Oppel A. (1862). "Über Fährten im lithographischen Schiefer. Paläontologische Mitteilungen aus dem Museum des Koenigl." Bayrischen Staates, ed. A. Oppel, vol. 1, pp. 121–125 & pl. 39. Stuttgart: Ebner & Sembek. In the mid-20th century, the possibility was raised that these multi-toothed skulls with distinctive awl-shaped teeth might belong to birds or pterosaurs rather than
gharial The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family (biology), family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males ...
-like crocodilians. However, it was not until the work of Peter Wellnhofer, who systematically reviewed known German pterosaur fossils in 1970, that ''Ctenochasma'' and ''Gnathosaurus'' became universally accepted as pterosaurs. At this time, many juvenile pterosaurs previously referred to as ''Pterodactylus micronyx'' and ''Pterodactylus elegans'' were referred to ''Ctenochasma''. This included the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
specimen of ''P. elegans''. Because the species name ''elegans'' was named before ''gracile'', the species is now known as ''Ctenochasma elegans''. Another specimen originally attributed to a small or juvenile ''Pterodactylus'', ''P. brevirostris'' (Wellnhofer's "example 29", which was later placed in its own genus ''Ptenodracon''), probably represents a young juvenile ''Ctenochasma'' based on similar wing bone proportions that differ from the similar ''
Aurorazhdarcho ''Aurorazhdarcho'' is an extinct genus of ctenochasmatoid pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous ...
''. An additional species, ''C. porocristata'', was named by
Paul de Buisonjé Paul may refer to: People * Paul (given name), a given name, including a list of people * Paul (surname), a list of people * Paul the Apostle, an apostle who wrote many of the books of the New Testament * Ray Hildebrand, half of the singing duo P ...
in 1981. However, it was differentiated mainly by the presence of a crest along the snout, which has since been shown to be a feature related to growth or sex, rather than species. A fourth species of ''Ctenochasma'' was first described (but not named) by
Philippe Taquet Philippe Taquet (born 25 April 1940 Saint-Quentin, Aisne) is a French paleontologist who specializes in dinosaur systematics of finds primarily in northern Africa. He is a member of the French Academy of Sciences since 30 November 2004, president ...
in 1972. A single specimen, consisting of a partial skull with complete brain case, was found in France and housed in the collections of the Saint-Dizier Museum. Detailed comparison to other ''Ctenochasma'' specimens in 2004 confirmed that it was a new species. In honor of Taquet's work on the specimen, Christopher Bennett named the species ''C. taqueti'' in 2007. Although researchers in the late 1990s and early 2000s, including Bennett and Jouve, suggested that all these species probably represented growth stages on just one kind of ''Ctenochasma'', in a more comprehensive 2007 study, Bennett demonstrated that three distinct species could be recognized based on the number and slenderness of the teeth, which varied across all growth stages. ''C. roemeri'', the earliest species, is characterized by its relatively low number of only moderately slender teeth. The later ''C. taqueti'' had a moderate number of slender teeth, and the last species, ''C. elegans'', had a high number of very slender teeth. These three species, arranged chronologically, probably represent a single evolutionary lineage in which the filter-feeding apparatus was gradually refined.


Description

''Ctenochasma'' is distinguished mainly by its numerous (over 400 in adults) long, thin, curved and closely packed teeth, which lined its elongated and narrow snout. The teeth were so closely packed that they formed a comb, and in adults they projected outward away from the jaws, forming a basket; traditionally, these are thought to indicate a
filter feeding Filter feeders are aquatic animals that acquire nutrients by feeding on organic matters, food particles or smaller organisms (bacteria, microalgae and zooplanktons) suspended in water, typically by having the water pass over or through a spe ...
lifestyle, straining water through the teeth in order to capture and eat small invertebrates, but unlike the related ''
Pterodaustro ''Pterodaustro'' (from Greek language, Greek , and Latin , ) is a genus of ctenochasmatid Pterodactyloidea, pterodactyloid pterosaur from South America. Its fossil remains dated back to the Early Cretaceous period, about 105 million years ago. ...
'' it lacks adaptations that would form a pumping mechanism. Instead, the spatulate profile formed by the teeth probably indicate a
spoonbill Spoonbills are a genus, ''Platalea'', of large, long-legged wading birds. The spoonbills have a global distribution, being found on every continent except Antarctica. The genus name ''Platalea'' derives from Ancient Greek and means "broad", refe ...
-like lifestyle, increasing the surface area of the jaws in order to catch small prey. The snout curved slightly upward and was rounded at the tip, and the teeth were restricted to the front half of the jaws. The smallest species, ''Ctenochasma elegans'', had a maximum wingspan of as an adult. ''Ctenochasma'' is distinguished by its mouth which contained as many as 260 long, thin comb-like teeth. Adult ''Ctenochasma'' had a bony crest along the skull, though this is not found in juveniles. Comparisons between the
scleral ring The scleral ring or sclerotic ring is a hardened ring of plates, often derived from bone, that is found in the eyes of many animals in several groups of vertebrates. Some species of mammals, amphibians, and crocodilians lack scleral rings. The ring ...
s of both ''Ctenochasma elegans'' and ''Ctenochasma taqueti'' and modern birds and reptiles suggest that these taxa may have been
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
, and may have had activity patterns similar to modern nocturnal seabirds. This may also indicate
niche partitioning In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for e ...
with contemporary pterosaurs inferred to be diurnal, such as ''Pterodactylus'' and ''
Scaphognathus ''Scaphognathus'' was a pterosaur that lived around Germany during the Late Jurassic. It had a wingspan of 0.9 m (3 ft). Naming The first known ''Scaphognathus'' specimen was described in 1831 by August Goldfuss who mistook the taille ...
''. Compared to other ctenochasmatoids, it had larger wings, and may have displayed a flying style comparable to that of modern
skua The skuas are a group of predatory seabirds with seven species forming the genus ''Stercorarius'', the only genus in the family Stercorariidae. The three smaller skuas, the Arctic skua, the long-tailed skua, and the pomarine skua, are called ...
s.


Classification

Below is
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
following a topology by Andres, Clark and Xu (2014). In the analysis, they recovered ''Ctenochasma'' within the family Ctenochasmatidae, more precisely within the subfamily Ctenochasmatinae, sister taxon to the tribe Pterodaustrini.


See also

*
List of pterosaur genera This list of pterosaurs is a comprehensive listing of all Genus, genera that have ever been included in the order Pterosauria, excluding purely vernacular terms. The list includes all commonly accepted genera, but also genera that are now considere ...
*
Timeline of pterosaur research This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and Biological taxonomy, taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic Era (ge ...


References

{{Portal bar, Paleontology, Germany, France Late Jurassic pterosaurs of Europe Early Cretaceous pterosaurs of Europe Ctenochasmatoids Solnhofen fauna Taxa named by Christian Erich Hermann von Meyer Fossil taxa described in 1851