Delorhynchus Skull
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''Delorhynchus'' 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 lanthanosuchoid
parareptile Parareptilia ("at the side of reptiles") is a subclass or clade of basal sauropsids (reptiles), typically considered the sister taxon to Eureptilia (the group that likely contains all living reptiles and birds). Parareptiles first arose near the ...
known from the late
Early Permian 01 or '01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * '01 (Richard Müller album), 01'' (Richard Müller album), 2001 * 01 (Son of Dave album), ''01'' (Son of Dave album), 2000 * 01 (Urban ...
(
Artinskian In the geologic timescale, the Artinskian is an age or stage of the Permian. It is a subdivision of the Cisuralian Epoch or Series. The Artinskian likely lasted between and million years ago (Ma) according to the most recent revision of the ...
age)
Garber Formation The Garber Formation is a geologic formation in Oklahoma. It preserves fossils dating back to the Permian period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainme ...
of
Comanche County, Oklahoma Comanche County is a county located in the U.S. state of Oklahoma. As of the 2010 census, the population was 124,098, making it the fourth-most populous county in Oklahoma. Its county seat is Lawton. The county was created in 1901 as part of ...
. It contains three
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
: the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
''D. priscus'' is based on a series of
maxillae 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 ...
. The second species to be described, ''D. cifellii'', is known from a larger number of well-preserved skulls and skeletal material. The third species, ''D. multidentatus'', is based on a fragmentary skull with several rows of teeth on its jaw.


Discovery

The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
, ''D. priscus,'' was first described and named by Richard C. Fox in 1962. The generic name "''Delorhynchus''" is derived from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''rhynchos''/''ρυγχος'', meaning "beak" (a common suffix for extinct reptile genera names). The specific name of the type species ''D. priscus'' is derived from Greek ''πρίσκος'', meaning "ancient" or "venerable" in reference to the fragmentary nature of the known remains. ''D. priscus'' is known from the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
KU 11117, a fragmentary left
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 ...
bearing 4 teeth, and from the fragmentary referred specimens KU 11118 and KU 11119, a right and a left maxillae respectively, each bearing 4 teeth. All known specimens of ''D. priscus'' are housed at the
University of Kansas Natural History Museum The University of Kansas Natural History Museum is part of the University of Kansas Biodiversity Institute, a KU designated research center dedicated to the study of the life of the planet. The museum's galleries are in Dyche Hall on the unive ...
in
Lawrence, Kansas Lawrence is the county seat of Douglas County, Kansas, Douglas County, Kansas, United States, and the sixth-largest city in the state. It is in the northeastern sector of the state, astride Interstate 70, between the Kansas River, Kansas and Waka ...
. ''D. cifellii'' was first described and named by Robert R. Reisz, Mark J. Macdougall and Sean P. Modesto in 2014. Its specific name honors Dr. Richard L. Cifelli, a paleontologist at the
Sam Noble Oklahoma Museum of Natural History Sam, SAM or variants may refer to: Places * Sam, Benin * Sam, Boulkiemdé, Burkina Faso * Sam, Bourzanga, Burkina Faso * Sam, Kongoussi, Burkina Faso * Sam, Iran * Sam, Teton County, Idaho, United States, a populated place People and fictio ...
(OMNH), for his contributions to the paleontology of Oklahoma and support in studying ''D. cifellii''. Unlike the type species, ''D. cifellii'' is known from a well-preserved partial subadult
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
, an isolated adult
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
, and other disarticulated elements, all housed at the OMNH. The subadult individual preserves both the partial skull and the postcranial remains in articulation, and thus was chosen as the holotype, represented by OMNH 73515. OMNH 73362 represents the complete skull of the large, adult, individual. Other remains referred to ''D. cifellii'' include OMNH 73363, a complete right mandibular ramus of an individual equal in size to OMNH 73362, as well as the right maxilla OMNH 73524. ''D. multidentatus'' was described by Dylan C.T. Rowe, Diane M. Scott, Joseph J. Bevitt, and Robert Reisz in
2021 File:2021 collage V2.png, From top left, clockwise: the James Webb Space Telescope was launched in 2021; Protesters in Yangon, Myanmar following the 2021 Myanmar coup d'état, coup d'état; A civil demonstration against the October–November 2021 ...
. It is based on ROMVP 87042, a fragmentary skull housed in the
Royal Ontario Museum The Royal Ontario Museum (ROM) is a museum of art, world culture and natural history in Toronto, Ontario, Canada. It is one of the largest museums in North America and the largest in Canada. It attracts more than one million visitors every year ...
of
Toronto Toronto ( ; or ) is the capital city of the Canadian province of Ontario. With a recorded population of 2,794,356 in 2021, it is the most populous city in Canada and the fourth most populous city in North America. The city is the ancho ...
. It was named in reference to the multiple rows of teeth on the lower jaw of the skull, a unique feature which distinguished it from other ''Delorhynchus'' species. ''Delorhynchus'' is only found in the fissure fill
claystone Mudrocks are a class of fine-grained siliciclastic sedimentary rocks. The varying types of mudrocks include siltstone, claystone, mudstone, slate, and shale. Most of the particles of which the stone is composed are less than and are too sm ...
and conglomerate of OMNH Locality V51. This locality, also known as
Richards Spur Richards Spur is a Permian fossil locality located at the Dolese Brothers Limestone Quarry north of Lawton, Oklahoma. The locality preserves clay and mudstone fissure fills of a karst system eroded out of Ordovician limestone and dolomite (rock), d ...
, is found at the Dolese Brothers limestone quarry near
Fort Sill Fort Sill is a United States Army post north of Lawton, Oklahoma, about 85 miles (136.8 km) southwest of Oklahoma City. It covers almost . The fort was first built during the Indian Wars. It is designated as a National Historic Landmark ...
in
Comanche County, Oklahoma Comanche County is a county located in the U.S. state of Oklahoma. As of the 2010 census, the population was 124,098, making it the fourth-most populous county in Oklahoma. Its county seat is Lawton. The county was created in 1901 as part of ...
. The fissure fill deposits of Richards Spur are probably equivalent to the
Garber Formation The Garber Formation is a geologic formation in Oklahoma. It preserves fossils dating back to the Permian period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainme ...
of Oklahoma and the Arroyo Formation of
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
. Uranium-Lead dating of speleotherms in the cave deposits suggest that the fossils of Richards Spur were deposited in the
Artinskian In the geologic timescale, the Artinskian is an age or stage of the Permian. It is a subdivision of the Cisuralian Epoch or Series. The Artinskian likely lasted between and million years ago (Ma) according to the most recent revision of the ...
stage of the
Cisuralian The Cisuralian is the first Series (stratigraphy), series/Epoch (geology), epoch of the Permian. The Cisuralian was preceded by the Pennsylvanian (geology), Pennsylvanian and followed by the Guadalupian. The Cisuralian Epoch is named after the w ...
(Early Permian).


Classification

Until recently, the
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
position of ''Delorhynchus'' within
Procolophonomorpha Procolophonomorpha is an order or clade containing most parareptiles. Many papers have applied various definitions to the name, though most of these definitions have since been considered synonymous with modern parareptile clades such as Ankyram ...
was uncertain, due to the fragmentary nature of the remains of ''D. priscus''. However, with the description of ''D. cifellii'' by Reisz ''et al.'' (2014), ''Delorhynchus'' could be added for the first time into a
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
to resolve its position. Reisz ''et al.'' (2014) recovered ''Delorhynchus'' in a
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
position to 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, ...
formed by ''
Acleistorhinus ''Acleistorhinus'' (ah-kles-toe-RYE-nuss) is an extinct genus of parareptile known from the Early Permian (middle Kungurian stage) of Oklahoma. It is notable for being the earliest known anapsid reptile yet discovered. The morphology of the l ...
'' and ''
Lanthanosuchus ''Lanthanosuchus'' is an extinct genus of parareptile from the Late Permian. It was found at Isheevo in Tatarstan The Republic of Tatarstan (russian: Республика Татарстан, Respublika Tatarstan, p=rʲɪsˈpublʲɪkə tət ...
''. Thus ''Delorhynchus'' was either considered to be the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
of
Lanthanosuchoidea Lanthanosuchoidea is an extinct superfamily of ankyramorph parareptiles from the middle Pennsylvanian to the middle Guadalupian epoch ( Moscovian - Wordian In the geologic timescale, the Wordian is an age or stage of the Permian. It is th ...
, or a basal lanthanosuchoid, depending on the definition of Lanthanosuchoidea used. The following
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
is simplified after the phylogenetic analysis of MacDougall and Reisz (2014), an updated version of Reisz ''et al.'' (2014), and shows the placement of ''D. cifellii'' within
Parareptilia Parareptilia ("at the side of reptiles") is a subclass or clade of basal sauropsids (reptiles), typically considered the sister taxon to Eureptilia (the group that likely contains all living reptiles and birds). Parareptiles first arose near the ...
. Relationships within bolded clades are not shown.


Paleoecology

The Richards Spur locality presents a very rich Early Permian
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
paleofauna, including species of
chondrichthyes Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
, lepospondyls,
seymouriamorph Seymouriamorpha were a small but widespread group of limbed vertebrates (tetrapods). They have long been considered reptiliomorphs, and most paleontologists may still accept this point of view, but some analyses suggest that seymouriamorphs are s ...
s, basal
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
s, and basal
eureptile Eureptilia ("true reptiles") is one of the two major subgroups of the clade Sauropsida, the other one being Parareptilia. Eureptilia includes Diapsida (the clade containing all modern reptiles and birds), as well as a number of primitive Permo ...
s. At least six species of
parareptile Parareptilia ("at the side of reptiles") is a subclass or clade of basal sauropsids (reptiles), typically considered the sister taxon to Eureptilia (the group that likely contains all living reptiles and birds). Parareptiles first arose near the ...
s have been described apart from the three ''Delorhynchus'' species. They include the basal parareptile ''
Microleter mckinzieorum ''Microleter'' is an extinct genus of basal procolophonomorph parareptiles which lived in Oklahoma during the Early Permian period. The type and only known species is ''Microleter mckinzieorum''. ''Microleter'' is one of several parareptile ...
'', the
bolosaurid Bolosauridae is an extinct family of ankyramorph parareptiles known from the latest Carboniferous (Gzhelian) or earliest Permian (Asselian) to the early Guadalupian epoch (latest Roadian stage) of North America, China, Germany, Russia and France. ...
''
Bolosaurus grandis ''Bolosaurus'' (from Ancient Greek ''bolos'', "lump" + ''sauros'': lizard]) is an extinct genus of bolosaurid ankyramorph parareptile from the Cisuralian epoch (middle Sakmarian to early Kungurian stages) of North Asia and North America (Red ...
'', the
nyctiphruretid Nyctiphruretidae is an extinct family of hallucicranian parareptiles known from the late Early to the late Middle Permian of European Russia and south-central United States. Nyctiphruretidae was named by Efremov in 1938. The type genus is ''Nyct ...
'' Abyssomedon williamsi'', and three other lanthanosuchoids: '' Colobomycter pholeter'', ''Colobomycter vaughni'', and ''
Feeserpeton oklahomensis ''Feeserpeton'' is an extinct genus of parareptile from the Early Permian of Richard's Spur, Oklahoma. It is known from a single species, ''Feeserpeton oklahomensis'', which was named in 2012 on the basis of a nearly complete skull. ''Feeserpet ...
''.


References

{{Taxonbar, from=Q252747 Procolophonomorphs Permian reptiles of North America Fossil taxa described in 1962 Fossil taxa described in 2014 Extinct animals of the United States Paleontology in Oklahoma Prehistoric reptile genera