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''Dimorphodon'' was a genus of medium-sized pterosaur from the early Jurassic Period. It was named by paleontologist
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Owe ...
in 1859. ''Dimorphodon'' means "two-form tooth", derived from the Greek (') meaning "two", (') meaning "shape" and (') meaning "tooth", referring to the fact that it had two distinct types of teeth in its jaws – which is comparatively rare among reptiles. ''Dimorphodon'' inhabited Europe.


Description

The body structure of ''Dimorphodon'' displays many "primitive" characteristics, such as, according to Owen, a very small brain-pan and proportionally short wings. The first phalanx in its flight finger is only slightly longer than its lower arm. The neck was short but strong and flexible and may have had a membranous pouch on the underside. The vertebrae had pneumatic foramina, openings through which the air sacs could reach the hollow interior. ''Dimorphodon'' had an adult body length of long, with a 1.45 metre (4.6 ft) wingspan. The tail of ''Dimorphodon'' was long and consisted of thirty vertebrae. The first five or six were short and flexible, but the remainder gradually increased in length and were stiffened by elongated vertebral processes. The terminal end of the tail may have borne a '' Rhamphorhynchus''-like tail vane, although no impressions have yet been found in ''Dimorphodon'' fossils to confirm this speculation.


Skull

''Dimorphodon'' had a large, bulky skull approximately in length, whose weight was reduced by large openings separated from each other by thin bony partitions. Its structure, reminiscent of the supporting arches of a bridge, prompted Richard Owen to declare that, as far as achieving great strength from lightweight materials was concerned, no vertebra was more economically constructed; Owen saw the vertebrate skull as a combination of four vertebrae modified from the
ideal type Ideal type (german: Idealtypus), also known as pure type, is a typological term most closely associated with sociologist Max Weber (1864–1920). For Weber, the conduct of social science depends upon the construction of abstract, hypothetical co ...
of the vertebra. The front of the upper jaw had four or five fang-like teeth followed by an indeterminate number of smaller teeth; the maxilla of all exemplars is damaged at the back. The lower jaw had five longer teeth and thirty to forty tiny, flattened pointed teeth, shaped like lancets."Dimorphodon." In: Cranfield, Ingrid (ed.). ''The Illustrated Directory of Dinosaurs and Other Prehistoric Creatures''. London: Salamander Books, Ltd. Pp. 288-291. Many depictions give it a speculative
puffin Puffins are any of three species of small alcids (auks) in the bird genus ''Fratercula''. These are pelagic seabirds that feed primarily by diving in the water. They breed in large colonies on coastal cliffs or offshore islands, nesting in crev ...
-like 'beak' because of similarities between the two animals' skulls.


History of discovery

The first fossil remains now attributed to ''Dimorphodon'' were found in England by fossil collector Mary Anning, at Lyme Regis in Dorset, UK in December 1828. This region of Britain is now a World Heritage Site, dubbed the Jurassic Coast; in it layers of the Blue Lias are exposed, dating from the
Hettangian The Hettangian is the earliest age and lowest stage of the Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (My ...
- Sinemurian. The specimen was acquired by William Buckland and reported in a meeting of the
Geological Society The Geological Society of London, known commonly as the Geological Society, is a learned society based in the United Kingdom. It is the oldest national geological society in the world and the largest in Europe with more than 12,000 Fellows. Fe ...
on 5 February 1829. In 1835, after a thorough study by William Clift and William John Broderip, this report, strongly expanded, was published in the Transactions of the Geological Society, describing and naming the fossil as a new species. As was the case with most early pterosaur finds, Buckland classified the remains in the genus '' Pterodactylus'', coining the new species ''Pterodactylus macronyx''. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
is derived from Greek ''makros'', "large" and ''onyx'', "claw", in reference to the large claws of the hand. The specimen, presently NHMUK PV R 1034, consisted of a partial and disarticulated skeleton on a slab, lacking the skull. Buckland in 1835 also assigned a piece of jaw from the collection of
Elizabeth Philpot Elizabeth Philpot (1780–1857) was an early 19th-century British fossil collector, amateur palaeontologist and artist who collected fossils from the cliffs around Lyme Regis in Dorset on the southern coast of England. She is best known today fo ...
to ''P. macronyx''. Later, the many putative species assigned to ''Pterodactylus'' had become so anatomically diverse that they began to be broken into separate genera. In 1858,
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Owe ...
reported finding two new specimens, NHMUK PV OR 41212 and NHMUK PV R 1035, again partial skeletons but this time including the skulls. Having found the skull to be very different from that of ''Pterodactylus'', Owen assigned ''Pterodactylus macronyx'' its own genus, which he named ''Dimorphodon''. His first report contained no description and the name remained a ''
nomen nudum In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate descr ...
''. In 1859, however, a subsequent publication by Owen provided a description. After several studies highlighting aspects of ''Dimorphodon''s anatomy, Owen in 1874 made NHMUK PV R 1034 the holotype. Meanwhile, though ''Dimorphodon'' is not a very common fossil, other fragmentary specimens were found. Some of these were acquired by Othniel Charles Marsh between 1873 and 1881 from the London
fossil dealer The fossil trade is the purchase and sale of fossils. This is many times done illegally with stolen fossils, and many important scientific specimens are lost each year. The trade is lucrative, and many celebrities collect fossils. The fossil tr ...
Bryce McMurdo Wright Bryce may refer to: People * Bryce (given name) * Bryce (surname) Places * Bryce Canyon National Park * Mount Bryce * Bryce, Utah * Bryce, Arizona Other * Bryce (software) * Bryce Hospital See also *Brice (disambiguation) Bricius most often refe ...
. One of these had been recovered from early Jurassic strata at the south bank of the Severn river, at the
Aust Cliff Aust Cliff () is a 5.3 hectare geological Site of Special Scientific Interest adjacent to the Severn Estuary, near the village of Aust, South Gloucestershire, notified in 1954. The Severn Bridge crosses the cliff. Its SSSI designation is due to ...
. An additional species of ''Dimorphodon'', ''D. weintraubi'', was named by James Clark ''et al'' in 1998 from a partial skeleton recovered in siltstones from the site Huizachal Canyon in
La Boca Formation The La Boca Formation is a geological formation in Tamaulipas state, northeast Mexico. It was thought to date back to the Early Jurassic, concretely the Pliensbachian stage epoch. Although, the latest studies had proven that the local Vulcanism, r ...
in Tamaulipas, Mexico, from the Early Jurassic ( Pliensbachian), where remains of
sphenodontian Rhynchocephalia (; ) is an order of lizard-like reptiles that includes only one living species, the tuatara (''Sphenodon punctatus'') of New Zealand. Despite its current lack of diversity, during the Mesozoic rhynchocephalians were a diverse g ...
s, dinosaurs and mammaliaforms have also been found. It is known from the type specimen, IGM 3494 (Instituto Geológico de México, of the Universidad Nacional Autónoma de México), that comprises articulated pieces of the skeleton including the posterior part of skull, four cervical vertebrae, the scapulocoracoids, left humerus, partial right wing and right leg distal to mid tibiotarsus. This specimen is larger than ''D. macronyx'' and the well preserved foot of it shows that pterosaurs do not have a
digitigrade In terrestrial vertebrates, digitigrade () locomotion is walking or running on the toes (from the Latin ''digitus'', 'finger', and ''gradior'', 'walk'). A digitigrade animal is one that stands or walks with its toes (metatarsals) touching the groun ...
posture in their hindlimbs, but that it have a plantigrade gait, as has been inferred from footprints. The name of the species is a homage to Dr. Robert L. Weintraub. Later studies considered this species not closely related to ''Dimorphodon macronyx'', but an early relative of Anurognathidae.


Classification

In 1870, Seeley assigned ''Dimorphodon'' to its own family,
Dimorphodontidae Dimorphodontidae (or dimorphodontids) is a group of early "rhamphorhynchoid" pterosaurs named after '' Dimorphodon'', that lived in the Late Triassic to Early Jurassic. While fossils that can be definitively referred to the group are rare, dimorp ...
, with ''Dimorphodon'' as the only member. It was suggested in 1991 by the German paleontologist Peter Wellnhofer that ''Dimorphodon'' might be descended from the earlier European pterosaur '' Peteinosaurus''. Later exact cladistic analyses are not in agreement. According to Unwin, ''Dimorphodon'' was related to, though probably not a descendant of, ''Peteinosaurus'', both forming the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Dimorphodontidae, the most basal group of the Macronychoptera and within it the sister group of the
Caelidracones The Caelidracones is a group of pterosaurs. The clade Caelidracones was defined in 2003 by David Unwin as the group consisting of the last common ancestor of ''Anurognathus ammoni'' and '' Quetzalcoatlus northropi'', and all its descendants. ...
. This would mean that both dimorphodontid species would be the most basal pterosaurs known with the exception of '' Preondactylus''. According to Alexander Kellner, however, ''Dimorphodon'' is far less basal and not a close relative of ''Peteinosaurus''. The cladogram recovered by Andres and Myers in 2013 is reproduced below.


Palaeobiology


Diet

The knowledge of how ''Dimorphodon'' lived is limited. It perhaps mainly inhabited coastal regions and might have had a very varied diet. Buckland suggested it ate insects. Later, it became common to depict it as a piscivore (fish eater), though biomechanical studies support Buckland's original insectivore idea better, and inconsistent with the animal's habits (see flight below). ''Dimorphodon'' had an advanced jaw musculature specialized for a "snap and hold" method of feeding. The jaw could close extremely quickly, but with relatively little force or tooth penetration. This, along with the short and high skull and longer, pointed front teeth suggest that ''Dimorphodon'' was an
insectivore A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores wer ...
, though it may have occasionally eaten small vertebrates and carrion as well. Mark Witton has argued that the animal was a specialised carnivore, being too large for an insectivorous diet and therefore specialised to hunt small lizards, sphenodonts and mammals, though its relatively weak jaw musculature probably meant that it ate proportionally small prey.
Dental microwear Dental microwear analysis is a method to infer diet and behavior in extinct animals, especially in fossil specimens. Typically, the patterns of pits and scratches on the occlusal or buccal surface of the enamel are compared with patterns observed ...
examinations confirm its status as a vertebrate predator, as opposed to several other insectivore or piscivore early pterosaurs.


Locomotion

Like many pterosaurs, ''Dimorphodon'' has been perceived as a soarer in the past, correlating to historical perceptions of pterosaurs as seabird analogues. However, more recent studies show that the animal was actually a rather poor flyer: its wings are proportionally short in relation to the body and its skeleton rather robust, offering very little gliding potential. In life, ''Dimorphodon'' probably relied on frantic short flights in the same manner as modern fowl, tinamous and woodpeckers, being unable to fly for long distances and probably only taking to the air as a last resort. Its derived position amidst primitive pterosaurs implies that this ineptitude is a developed trait, not an ancestral characteristic, as earlier pterosaurs like '' Preondactylus'' were capable aeronauts. Owen saw ''Dimorphodon'' as a quadruped. He speculated that the fifth toe supported a membrane between the tail and the legs and that the animal was therefore very ungainly on the ground. However, his rival Harry Govier Seeley, propagating the view that pterosaurs were warm-blooded and active, argued that ''Dimorphodon'' was either an agile quadruped or even a running biped due to its relatively well developed hindlimbs and characteristics of its pelvis. This hypothesis was revived by Kevin Padian in 1983. However, fossilised track remains of other pterosaurs ( ichnites) show a quadrupedal gait while on the ground and these traces are all attributed to derived pterosaurs with a short fifth toe. ''Dimorphodon'''s was elongated, clawless, and oriented to the side. David Unwin has therefore argued that even ''Dimorphodon'' was a quadruped, a view confirmed by computer modelling by Sarah Sangster. Like most non-pterodactyloid pterosaurs, ''Dimorphodon'' was a competent climber, possessing proportionally large and curved ungals and a low center of gravity. Like modern squirrels, it probably moved in a saltatorial manner as it climbed.


See also

* List of pterosaur genera * Timeline of pterosaur research * Mary Anning


References

{{Taxonbar, from=Q131245 Pterosaurs Early Jurassic pterosaurs Jurassic reptiles of Europe Early Jurassic reptiles of North America Taxa named by Richard Owen Fossil taxa described in 1859