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''Fodonyx'' (meaning "digging claw") is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
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
rhynchosaur Rhynchosaurs are a group of extinct herbivorous Triassic archosauromorph reptiles, belonging to the order Rhynchosauria. Members of the group are distinguished by their triangular skulls and elongated, beak like premaxillary bones. Rhynchosaurs ...
from the middle
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
epoch of
Devon Devon ( ; historically also known as Devonshire , ) is a ceremonial county in South West England. It is bordered by the Bristol Channel to the north, Somerset and Dorset to the east, the English Channel to the south, and Cornwall to the west ...
in
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
. Its fossils (25 specimens) were discovered in
Otter Sandstone Formation The Otter Sandstone Formation is an obsolete geological unit in England, now referred to as the Helsby Sandstone Formation.http://www.bgs.ac.uk/Lexicon/lexicon.cfm?pub=HEY (BGS on-line lexicon of rock units) It preserves fossils dated to the Tr ...
(late
Anisian In the geologic timescale, the Anisian is the lower stage (stratigraphy), stage or earliest geologic age, age of the Middle Triassic series (stratigraphy), series or geologic epoch, epoch and lasted from million years ago until million years ag ...
age) and were first assigned to '' Rhynchosaurus spenceri''. This species was reassigned to its own genus, ''Fodonyx'' (the type and only species is ''Fodonyx spenceri'') the holotype of which i
EXEMS 60/1985/292
that described by David W. E. Hone and Michael J. Benton in
2008 2008 was designated as: *International Year of Languages *International Year of Planet Earth *International Year of the Potato *International Year of Sanitation The Great Recession, a worldwide recession which began in 2007, continued throu ...
. In 2010, one skull was reassigned to the new genus '' Bentonyx''. It is distinguished from other rhynchosaurs by a single
autapomorphy In phylogenetics, an autapomorphy is a distinctive feature, known as a Synapomorphy, derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or Outgroup (cladistics), outgroup taxa, not ...
, the ventral angling of the paraoccipital processes. In all other rhynchosaurs these processes angle dorsally or are horizontal. It is not known if this conferred any advantage to ''Fodonyx. Fodonyx'' was between 40 and 50 cm long.


Features


Skull and lower jaw

The two
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
e are very long and run up over the snout to meet the prefrontals at the orbit. At the anterior tip they are narrow and triangular in cross-section. They form the classic rhynchosaurian beak, and there is evidence on the fossil showing that it was probably covered by a keratinous sheath. The
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
carries a massive tooth plate and has numerous
foramina In anatomy and osteology, a foramen (; : foramina, or foramens ; ) is an opening or enclosed gap within the dense connective tissue (bones and deep fasciae) of extant and extinct amniote animals, typically to allow passage of nerves, arter ...
for
nerve A nerve is an enclosed, cable-like bundle of nerve fibers (called axons). Nerves have historically been considered the basic units of the peripheral nervous system. A nerve provides a common pathway for the Electrochemistry, electrochemical nerv ...
s and
blood vessels Blood vessels are the tubular structures of a circulatory system that transport blood throughout many animals’ bodies. Blood vessels transport blood cells, nutrients, and oxygen to most of the tissues of a body. They also take waste an ...
to reach the
gums The gums or gingiva (: gingivae) consist of the mucosal tissue that lies over the mandible and maxilla inside the mouth. Gum health and disease can have an effect on general health. Structure The gums are part of the soft tissue lining of the ...
through. Many of the posterior and lateral teeth are unworn from use, unlike the more anterior teeth which have been worn smooth. The
nasal bone The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
s are large, but no wider than the frontals. They form a pointed posterior tip with a strong zigzag suture. The lacrimal ducts are clearly visible next to the
orbit In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...
, while the
lacrimal bone The lacrimal bones are two small and fragile bones of the facial skeleton; they are roughly the size of the little fingernail and situated at the front part of the medial wall of the orbit. They each have two surfaces and four borders. Several bon ...
s form much of the interior surface of the orbit. The prefrontal forms a thick eyebrow ridge, possibly as protection from predators. The
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
is complex, with four branches, and forms the anterior and ventral margins of the lower
temporal fenestra Temporal fenestrae are openings in the temporal region of the skull of some amniotes, behind the orbit (eye socket). These openings have historically been used to track the evolution and affinities of reptiles. Temporal fenestrae are commonly (al ...
. The dorsal branch forms a strong pillar behind the orbit, which has a more pronounced crest than other rhynchosaurs. The frontals are very long, and form a dish shape posteriorly. The
postfrontal The postfrontal is a paired cranial bone found in many tetrapods. It occupies an area of the skull roof between and behind the orbits (eye sockets), lateral to the frontal and parietal bones, and anterior to the postorbital bone. The postfrontal ...
is triangular and forms part of the back of the orbit. The parietals are fused and have a high narrow ridge dorsally, with lateral wings extending across the upper temporal fenestrae. The postorbitals are roughly T-shaped, with three branches. Unlike Late Triassic forms, ''Fodonyx'' has a
supratemporal The supratemporal bone is a paired Skull, cranial bone present in many Tetrapod, tetrapods and Tetrapodomorpha, tetrapodomorph fish. It is part of the temporal region (the portion of the skull roof behind the eyes), usually lying medial (inwards) re ...
bone. The
quadratojugal The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians. Anatomy and function In animals with a quadratojugal bone, it is typically found connected to the jugal (cheek) bone from the front and ...
and quadrate are mainly missing. One
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestra ...
is preserved, forming much of the posterior margin of the skull. Much of the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sep ...
is intact, although the vomeronasals are quite degraded due to their length and thinness. The
palatines Palatines () were the citizens and princes of the Palatinates, Holy Roman States that served as capitals for the Holy Roman Emperor. After the fall of the Holy Roman Empire in 1806, the nationality referred more specifically to residents of the ...
form most of the borders of the
choana The choanae (: choana), posterior nasal apertures or internal nostrils are two openings found at the back of the nasal passage between the nasal cavity and the pharynx, in humans and other mammals (as well as crocodilians and most skinks). They ...
. The pterygoids are very large and have three main processes, all broad and flat. The ectopterygoids are very small and hidden in palatal view. Small fragments of the hyoids are preserved, with a circular cross section and lateral striations. The basioccipital is short and attached to the narrow basisphenoid. The occipital condyle is hemispherical. Much of the detail on the paraoccipital is hard to make out due to difficulties of preparation. Only the anterior portions of the
lower jaw In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
are well preserved, but it has the typical rhynchosaur shape, curving up to the anterior tip. Teeth are mainly obscured as the jaws are tightly shut. The
splenial The splenial is a small bone in the lower jaw of reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology ...
is narrow except at the tip where it supports the symphyseal plate.


Spine and vertebrae

Cervical and
sacral vertebrae The sacrum (: sacra or sacrums), in human anatomy, is a triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part of the pelvic cavity, ...
are not known, only
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...
and caudal. The dorsal vertebrae have round
centra Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the individual shops are all owned by individual franchisees. The chain has three ...
which narrow noticeably towards the centre of the bone, and are deeply amphicoelous. They lack a keel, but have a deep excavation in the floor of the
neural canal In the developing chordate (including vertebrates), the neural tube is the embryonic precursor to the central nervous system, which is made up of the brain and spinal cord. The neural groove gradually deepens as the neural folds become elevated, ...
. Neural arches are around 15 mm tall, with narrow pillars supporting the zygapophyses, and are attached by broad flat facets. The zygapophyses are almost circular and very flat, which would allow side-to-side movement but little up-and-down movement. Three
chevrons Chevron (often relating to V-shaped patterns) may refer to: Science and technology * Chevron (aerospace), sawtooth patterns on some jet engines * Chevron (anatomy), a bone * '' Eulithis testata'', a moth * Chevron (geology), a fold in rock la ...
are preserved. The dorsal elements have fused to create a triangular opening.


Ribs and gastralia

Many
rib In vertebrate anatomy, ribs () are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the thoracic cavity, enabling the lungs to expand and thus facilitate breathing by expanding the ...
s are present, although partially fragmented, and show that the ribs were robust and the
rib cage The rib cage or thoracic cage is an endoskeletal enclosure in the thorax of most vertebrates that comprises the ribs, vertebral column and sternum, which protect the vital organs of the thoracic cavity, such as the heart, lungs and great ve ...
deep. They were also remarkably straight, at least at the anterior. There are also many
gastralia Gastralia (: gastralium) are dermal bones found in the ventral body wall of modern crocodilians and tuatara, and many prehistoric tetrapods. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In these reptil ...
, although these are quite jumbled together, and appear to be made of three segments. The gastral basket is almost entirely disarticulated.


Forelimbs and pectoral girdle

The
scapula The scapula (: scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either side ...
e have broad blades, and prominent bosses where the
clavicle The clavicle, collarbone, or keybone is a slender, S-shaped long bone approximately long that serves as a strut between the scapula, shoulder blade and the sternum (breastbone). There are two clavicles, one on each side of the body. The clavic ...
s may have attached. Much of the front limbs are missing, but the
humeri The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of ...
were broad and not very long, with an oval cross-section.


Hindlimbs and pelvic girdle

The
pelvis The pelvis (: pelves or pelvises) is the lower part of an Anatomy, anatomical Trunk (anatomy), trunk, between the human abdomen, abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also c ...
is incomplete, with the ilium clearly showing growth lines. The
ischium The ischium (; : is ...
has a thick round dorsal margin and a curved blade. The
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
is missing, but the remainder of the hindlimb is present. The
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
is fairly long but quite thin, with a compressed oval cross-section. It shows heavy wear, and has a distinct twist where it probably attached to the
fibula The fibula (: fibulae or fibulas) or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. ...
. This is more slender than the tibia, but still relatively robust. All the proximal tarsals and three of the distal tarsals are present, but are worn and broken. In general, they are quite rounded. The first
metatarsal The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
is short and broad but the other four are long and flat, although they are broken so it is hard to say exactly how long. The toes have, respectively, 2,3,4,5 and 4
phalanges The phalanges (: phalanx ) are digit (anatomy), digital bones in the hands and foot, feet of most vertebrates. In primates, the Thumb, thumbs and Hallux, big toes have two phalanges while the other Digit (anatomy), digits have three phalanges. ...
. These all narrow as they head towards the claws. The
ungual An ungual (from Latin ''unguis'', i.e. ''nail'') is a highly modified distal toe bone which ends in a hoof, claw, or nail. Elephants and ungulates have ungual phalanges, as did the sauropod Sauropoda (), whose members are known as sauropods (; ...
s are all very large and broad, and have rounded ends without a recurve. Each ungual bears a shallow groove along the sides, probably for locking the keratin sheath.


Taxonomy

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 ...
based on Ezcurra ''et al.'' (2016):


Links

* *


References

{{Taxonbar, from=Q5463983 Rhynchosauria Middle Triassic reptiles of Europe Fossil taxa described in 2008 Anisian life Prehistoric reptile genera