Dasygnathoides
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''Dasygnathoides'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
pseudosuchia Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
n from the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...
(
Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by t ...
)
Lossiemouth Sandstone The Lossiemouth Sandstone is a Middle to Late Triassic (Ladinian to Norian) age geological formation. It is exposed on the south side of the Moray Firth near Lossiemouth and near Golspie in Sutherland. Dinosaur remains are among the fossils tha ...
of
Scotland Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to the ...
.


Classification

"''Dasygnathus''" ''longidens'' was erected by
Thomas Huxley Thomas Henry Huxley (4 May 1825 – 29 June 1895) was an English biologist and anthropologist specialising in comparative anatomy. He has become known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution. The storie ...
for a
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
from the Lossiemouth Sandstone in 1877. The genus name '' Dasygnathus'' had already been used for a
coleoptera Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
n insect, so
Oskar Kuhn Oskar Kuhn (7 March 1908, Munich – 1990) was a German palaeontologist. Life and career Kuhn was educated in Dinkelsbühl and Bamberg and then studied natural science, specialising in geology and paleontology, at the University of Munich, fr ...
renamed it ''Dasygnathoides''.Kukn, O., 1961, Fossilium Catalogus I: Animalia, Reptila, supplementum 1, n. 2, p. 1-163. Although synonymized with ''
Ornithosuchus ''Ornithosuchus'' (meaning "bird crocodile") is an extinct genus of pseudosuchians from the Late Triassic (Carnian) Lossiemouth Sandstone of Scotland. It was originally thought to be the ancestor to the carnosaurian dinosaurs (such as ''Allosauru ...
'' by Walker (1964), a 2016 study found ''Dasygnathoides'' indeterminate beyond Pseudosuchia. For this reason, many paleontologists consider '''Dasygnathoides a nomem dubium. The lack of
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
shared by
ornithosuchids Ornithosuchidae is an extinct family of pseudosuchian archosaurs (distant relatives of modern crocodilians) from the Triassic period. Ornithosuchids were quadrupedal and facultatively bipedal (e.g. like chimpanzees), meaning that they had the a ...
indicates that ''Dasygnathoides'' was not an ornithosuchid as previously thought, but there are no links between it and any other family of pseudosuchians. It has remained impossible to resolve further. The morphology of the maxilla is quite different from
aetosaurs Aetosaurs () are heavily armored reptiles belonging to the extinct order Aetosauria (; from Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs ...
,
silesaurids Silesauridae is an extinct family of Triassic dinosauriforms. It is most commonly considered to be a clade of non-dinosaur dinosauriforms, and the sister group of dinosaurs. Some studies have instead suggested that most or all silesaurids compri ...
, erpetosuchians,
saurischians Saurischia ( , meaning "reptile-hipped" from the Ancient Greek, Greek ' () meaning 'lizard' and ' () meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia), classified by their hip structure. Saurisch ...
or
ornithischians Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek s ...
. The interdental plates also differ greatly from
erythrosuchids Erythrosuchidae (meaning "red crocodiles" in Greek) are a family of large basal archosauriform carnivores that lived from the later Early Triassic (Olenekian) to the early Middle Triassic (Anisian). Naming The family Erythrosuchidae was named b ...
and proterosuchids. ''Dasygnathoides'' cannot be assigned to any particular group of pseudosuchians at present. It is probably the first or only specimen of an entirely new group.


History

Thomas Huxley first found the maxilla in the Lossiemouth Sandstone in 1859, and originally identified it as ''
Stagonolepis ''Stagonolepis'' is an extinct genus of stagonolepidid aetosaur known from the Late Triassic (Carnian stage) Hassberge Formation of Germany, the Drawno Beds of Poland, and the Lossiemouth Sandstone of Scotland. Supposed fossils from North and ...
,'' due to its very similar tooth implantation pattern. He later changed his mind and reidentified it as a new species, "''Dasygnathus".'' This name later had to be changed in 1961 as it had already been used for a beetle. In 1964, the specimen of ''Dasygnathoides'' was compared with all the known ''Ornithosuchus'' specimens and then synonymized by Walker. This was due to the shared presence of 'rear forking' at the posterior end of the maxillae, only nine maxillary teeth, and the similarity between their right
pterygoid bone The pterygoid is a paired bone forming part of the palate of many vertebrates, behind the palatine bone In anatomy, the palatine bones () are two irregular bones of the facial skeleton in many animal species, located above the uvula in the th ...
s. In 2016, M. Belén von Baczko and M. Ezcurra reassessed this synonymization and found that ''Dasygnathoides'' was in fact a much larger and quite different species to ''Ornithosuchus,'' and not even an ornithosuchid (see above).


Features

''Dasygnathoides'' is known from a right maxilla and pterygoid, a partial
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
, a
haemal arch A haemal arch also known as a chevron, is a bony arch on the ventral side of a tail vertebra of a vertebrate. The canal formed by the space between the arch and the vertebral body is the haemal canal. A spinous ventral process emerging from the hae ...
and a
phalanx The phalanx ( grc, φάλαγξ; plural phalanxes or phalanges, , ) was a rectangular mass military formation, usually composed entirely of heavy infantry armed with spears, pikes, sarissas, or similar pole weapons. The term is particularly use ...
, an
articular The articular bone is part of the lower jaw of most vertebrates, including most jawed fish, amphibians, birds and various kinds of reptiles, as well as ancestral mammals. Anatomy In most vertebrates, the articular bone is connected to two oth ...
and an
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amp ...
. They all come from the same animal. There was also a small radius and ulna found in the slab, but this was far too small to have been from the same specimen and can probably be discounted as stomach contents. The total skull length is estimated to have been about 450 mm, almost twice as long as the largest ''Ornithosuchus'' skull found, and the entire animal was probably between 3 and 4 metres long. It is the largest predatory tetrapod known from Triassic Scotland.


Maxilla (ELGNM 1)

The maxilla found has four
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
remaining in it and nine
interdental plates The interdental plate refers to the bone-filled mesial-distal region between the teeth. The word "''interdental''" is a combination of "''inter''" + "''dental''" (meaning "''between the teeth''") which originated in approximately 1870. In paleobi ...
, showing that five teeth are missing. The total length of the maxilla is 195 mm and the teeth range from 29.5 mm to 53.5 mm long. The teeth are posteriorly curved and have sharp edges, indicating that they might have been used as blades to slice flesh rather than merely to impale and grip. The interdental plates are
pentagonal In geometry, a pentagon (from the Greek πέντε ''pente'' meaning ''five'' and γωνία ''gonia'' meaning ''angle'') is any five-sided polygon or 5-gon. The sum of the internal angles in a simple pentagon is 540°. A pentagon may be simpl ...
, would have formed the medial walls of the
tooth sockets Dental alveoli (singular ''alveolus'') are sockets in the jaws in which the roots of teeth are held in the alveolar process with the periodontal ligament. The lay term for dental alveoli is tooth sockets. A joint that connects the roots of the ...
, and in life were richly supplied with
blood vessels The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away f ...
, giving them a rather wrinkled look. No ascending or palatal process can be found on the specimen, and nor can the 'rear forking prong' mentioned above. Indeed, the prong mentioned can only be found on a single specimen of ''Ornithosuchus.''


Pterygoid (ELGNM 15)

This specimen shows the dorsal surface of the right pterygoid bone, and a small part of either a
palatine A palatine or palatinus (in Latin; plural ''palatini''; cf. derivative spellings below) is a high-level official attached to imperial or royal courts in Europe since Roman times.
or
vomer bone The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxil ...
. It is 164 mm long. The anterior ramus is narrow and tapering, but the medial margin is thick and the quadrate ramus is almost as wide as it is long. Th quadrate ramus is dorsally curved and bears a groove extending anterolaterally. Unlike ''Ornithosuchus, Dasygnathoides'' did not appear to have a palatine-pterygoid fenestra as the homologous area of bone is curved outwards. This indicates that, in fact, it is not an ornithosuchan at all. The partial vertebra is found in the same piece of stone as the maxilla. It has a well-developed transverse process and is probably a dorsal or anterior caudal vertebra.


Haemal arch & phalanx (ELGNM 15) and possible articular (ELGNM 29)

These are small and fragmentary, and little information can be gained from them. It is possible that they are, in fact, stomach or throat contents of the actual ''Dasygnathoides'' fossil.


Osteoderm (ELGNM unnumbered)

This was previously identified as a possible nasal bone, but is far more likely to be an osteoderm indicating that ''Dasygnathoides'' had skin armour. It resembles the elongated osteoderms of ''Ornithosuchus.''


References


Sources

* {{Taxonbar, from=Q27989447 Prehistoric pseudosuchian genera Triassic archosaurs Late Triassic reptiles of Europe