Ankylosphenodon
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''Ankylosphenodon'' is an extinct
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
sphenodontian Rhynchocephalia (; ) is an order (biology), 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 ...
known from multiple specimens recovered from the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
deposits of the
Tlayúa Formation The Tlayúa Formation is an Cretaceous Konservat-Lagerstätte near Tepexi de Rodríguez, Puebla, Central Mexico. It consists of three members (Lower, Middle and Upper), spanning the lower Albian of the Early Cretaceous to the lower Cenomanian of ...
, near Tepexi de Rodriguez,
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
. It is likely part of a sphenodontid lineage that also includes ''
Eilenodon ''Eilenodon'' is an extinct genus of rhynchocephalian reptile from the Late Jurassic Morrison Formation of western North America,Foster, J. (2007). "Table 2.1: Fossil Vertebrates of the Morrison Formation." ''Jurassic West: The Dinosaurs of the M ...
'' and ''
Toxolophosaurus ''Toxolophosaurus'' was a sphenodont which lived in North America during the Early Cretaceous. The first specimen, consisting of a pair of lower jaws, was found by George Cloud and described by Everett C. Olson in 1960 in the Kootenai Formatio ...
'', though has similarities to the
sapheosaur Sapheosaurs are an Extinction, extinct group of rhynchocephalian reptiles from the Late Jurassic period. "Sapheosaurs" is an informal name for a group of rhynchocephalians closely related to the genus ''Sapheosaurus''. It was first recognized as ...
clade. Its skeleton is pachyostotic (thickened) skeleton, believed to be an adaptation for a semi-aquatic lifestyle. Its teeth appear to have grown continuously, like in
rodents Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
.


Taxonomy


Discovery and formal description

''Ankylosphenodon'' is known from multiple specimens recovered from the
Tlayúa Formation The Tlayúa Formation is an Cretaceous Konservat-Lagerstätte near Tepexi de Rodríguez, Puebla, Central Mexico. It consists of three members (Lower, Middle and Upper), spanning the lower Albian of the Early Cretaceous to the lower Cenomanian of ...
, a
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
-rich
Lagerstätte A Fossil-Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that preserves an exceptionally high amount of palaeontological information. ''Konzentrat-Lagerstätten'' preserv ...
near Tepexi de Rodriguez, in Central
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
. 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 ...
(IGM 7441) is a partial skeleton, consisting of a skull, the left forelimb, and the front half of the presacral vertebral column. Another specimen, IGM 7443, preserves most of the right forelimb and the rear half of the body, minus the tail. All specimens were recovered from the Tlayúa Quarry, which, represents the Middle
Member Member may refer to: * Military jury, referred to as "Members" in military jargon * Element (mathematics), an object that belongs to a mathematical set * In object-oriented programming, a member of a class ** Field (computer science), entries in ...
of the
Tlayúa Formation The Tlayúa Formation is an Cretaceous Konservat-Lagerstätte near Tepexi de Rodríguez, Puebla, Central Mexico. It consists of three members (Lower, Middle and Upper), spanning the lower Albian of the Early Cretaceous to the lower Cenomanian of ...
, a
Lagerstätte A Fossil-Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that preserves an exceptionally high amount of palaeontological information. ''Konzentrat-Lagerstätten'' preserv ...
rich in vertebrate fossils. On collection, the specimens were transported to the Geological Institute of the
National Autonomous University of Mexico The National Autonomous University of Mexico (, UNAM) is a public university, public research university in Mexico. It has several campuses in Mexico City, and many others in various locations across Mexico, as well as a presence in nine countri ...
. In 2000, Víctor-Hugo Reynoso formally described the genus. The generic name comes from ''Sphenodon'', the generic name of the modern
tuatara The tuatara (''Sphenodon punctatus'') is a species of reptile endemic to New Zealand. Despite its close resemblance to lizards, it is actually the only extant member of a distinct lineage, the previously highly diverse order Rhynchocephal ...
, and the Greek ἀγκύλος (''ankylos''), meaning "fused" or "bent", referring to the thickening of its ribs and vertebrae.


Classification

The strict consensus tree in Reynoso's paper on ''Ankylosphenodon'' recovers it as part of a small, unnamed clade within
Sphenodontidae Sphenodontidae is a family within the reptile group Rhynchocephalia, comprising taxa most closely related to the living tuatara (''Sphenodon punctatus''). Historically the taxa included within Sphenodontidae have varied greatly between analyses, ...
, one that also includes ''
Eilenodon ''Eilenodon'' is an extinct genus of rhynchocephalian reptile from the Late Jurassic Morrison Formation of western North America,Foster, J. (2007). "Table 2.1: Fossil Vertebrates of the Morrison Formation." ''Jurassic West: The Dinosaurs of the M ...
'' and ''
Toxolophosaurus ''Toxolophosaurus'' was a sphenodont which lived in North America during the Early Cretaceous. The first specimen, consisting of a pair of lower jaws, was found by George Cloud and described by Everett C. Olson in 1960 in the Kootenai Formatio ...
''. Similarities with
sapheosaurs Sapheosaurs are an extinct group of rhynchocephalian reptiles from the Late Jurassic period. "Sapheosaurs" is an informal name for a group of rhynchocephalians closely related to the genus '' Sapheosaurus''. It was first recognized as a group con ...
were noted, though were ascribed to
convergence Convergence may refer to: Arts and media Literature *''Convergence'' (book series), edited by Ruth Nanda Anshen *Convergence (comics), "Convergence" (comics), two separate story lines published by DC Comics: **A four-part crossover storyline that ...
. However, Sebastian Apesteguía, in 2005, suggested that it was in fact a sapheosaur.


Description

''Ankylosphenodon'' had an upper skull length of . The length of the presacral
vertebral column The spinal column, also known as the vertebral column, spine or backbone, is the core part of the axial skeleton in vertebrates. The vertebral column is the defining and eponymous characteristic of the vertebrate. The spinal column is a segmente ...
(the cervical and
dorsal vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebra (anatomy), vertebrae of intermediate size between the ce ...
), measured from the first to nineteenth vertebrae, is estimated at .


Skull and mandible

The skull of ''Ankylosphenodon'' is poorly preserved. Assuming the skull and
mandible 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 ...
were equal in length, the upper temporal fenestra extended for about half the skull's length. The
premaxillae 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 has ...
are separated, and are the only part of the snout that is preserved in the holotype. The mandible is typical among sphenodonts, though is unusually robust. It is slenderer at the front (
anteriorly Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Thi ...
) than it is at the back (
posteriorly Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...
). Little in the way of fine details are preserved, save for the
dentary 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 ...
. The teeth are unique among sphenodonts, in that they extend deep into the dentary, as far as the
Meckelian groove The Meckelian groove (or Meckel's groove, Meckelian fossa, or Meckelian foramen, or Meckelian canal) is an opening in the medial (inner) surface of the mandible (lower jaw) which exposes the Meckelian cartilage.rhynchosaurs 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 ...
. The teeth of ''Ankylosphenodon'' appear to have grown continuously, similar to some mammals, such as
rodents Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
.


Postcranial elements

Among sphenodonts, ''Ankylosphenodon'' is distinct in having pachyostotic (thickened)
vertebrae Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
and ribs, which were massive compared to those of related genera. Its overall morphology is reflective of semi-aquatic reptiles, such as
crocodiles Crocodiles (family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include all extant member ...
and marine iguanas, suggesting that it may have adopted a similar lifestyle and swam similarly to the latter. Like the axial skeletons, the limbs were massive and heavily constructed. The hind limbs were longer than the forelimbs. The fifth pedal digit was smaller and slenderer than the others. ''Ankylosphenodon'''s tail was well-adapted to
lateral Lateral is a geometric term of location which may also refer to: Biology and healthcare * Lateral (anatomy), a term of location meaning "towards the side" * Lateral cricoarytenoid muscle, an intrinsic muscle of the larynx * Lateral release ( ...
motion, though resisted
dorsoventral Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
(up-and-down) motion.


Palaeoenvironment

The palaeoenvironment of the Tlayúa Formation, from which all specimens of ''Ankylosphenodon'' are known, was likely a shallow coastal
lagoon A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into ''coastal lagoons'' (or ''barrier lagoons'') an ...
. It may have formed part of an island, though a connection to the North American mainland cannot be ruled out. A certain degree of influence from freshwater environments is indicated by the presence of fossils from
crocodilians Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchi ...
and freshwater turtles.


References

{{Taxonbar, from=Q21368116 Rhynchocephalia Cretaceous reptiles of North America Prehistoric reptile genera