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''Microleter'' 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 basal
procolophonomorph 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 Ankyr ...
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 which lived in
Oklahoma Oklahoma (; Choctaw language, Choctaw: ; chr, ᎣᎧᎳᎰᎹ, ''Okalahoma'' ) is a U.S. state, state in the South Central United States, South Central region of the United States, bordered by Texas on the south and west, Kansas on the nor ...
during the
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 ...
period. The type and only known
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 ...
is ''Microleter mckinzieorum''. ''Microleter'' is one of several parareptile taxa described from the
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, with the ...
fissure fills, and can be characterized from its high tooth count, lacrimal/ narial contact, short
postfrontal 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, the ...
, and slit-like temporal emargination edged by the
postorbital The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...
,
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 ...
,
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 The quadratojugal is a skull bone present in many vertebrates, including ...
, and
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 ...
. Contrary to ''
Australothyris ''Australothyris'' is an extinct genus of basal procolophonomorph parareptile known from the Middle Permian (middle Capitanian stage) of ''Tapinocephalus'' Assemblage Zone, South Africa. The type and only known species is ''Australothyris smith ...
'', which had a similar
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 as a basal procolophonomorph, ''Microleter'' suggests that early parareptile evolution occurred in
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...
and that multiple lineages developed openings or emarginations in the temporal region.


Discovery

The only known specimen of ''Microleter'' is a well-preserved skull and lower jaw designated as OMNH 71306, 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 ...
specimen. It was found at the Dolese Brothers limestone quarry near
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, with the ...
in Comanche County,
Oklahoma Oklahoma (; Choctaw language, Choctaw: ; chr, ᎣᎧᎳᎰᎹ, ''Okalahoma'' ) is a U.S. state, state in the South Central United States, South Central region of the United States, bordered by Texas on the south and west, Kansas on the nor ...
. Historically referred to as the Fort Sill locality, the quarry has preserved
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 ...
, ~289-286 Ma) fissure fills in an
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
cave system. The fissure fills have yielded many other well preserved
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
fossils, including the most diverse assortment of Permian
parareptiles 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 t ...
in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
. ''Microleter mckinzieorum'' was named by paleontologists Linda A. Tsuji, Johannes Muller, and Robert R. Reisz in
2010 File:2010 Events Collage New.png, From top left, clockwise: The 2010 Chile earthquake was one of the strongest recorded in history; The Eruption of Eyjafjallajökull in Iceland disrupts air travel in Europe; A scene from the opening ceremony of ...
. The genus name combined Greek "''mikros"'' (small) with "-''oleter"'' (murderer), a
suffix In linguistics, a suffix is an affix which is placed after the stem of a word. Common examples are case endings, which indicate the grammatical case of nouns, adjectives, and verb endings, which form the conjugation of verbs. Suffixes can carry ...
common to parareptile genera. The specific name honors the McKinzie family, as the specimen was discovered by Mark McKinzie.


Description

Based on the skull's large
orbits In celestial mechanics, an orbit 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 object or position in space such as a p ...
(eye holes) and weak sutures, the specimen was likely a juvenile. Most of the skull bones were externally textured by radiating pits and furrows, with both sparse large pits and numerous tiny pits as in basal lanthanosuchoids. The only smoothly textured bones of the skull roof were the
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 ...
,
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 The quadratojugal is a skull bone present in many vertebrates, including ...
, and
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 ...
. The maxilla was long and narrow, possessing conical teeth which only differed from each other in a slight shortening trend towards the rear of the maxilla. The maxillary tooth count was 32 or 33, more than any other parareptile apart from ''
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 ...
''. Like several other basal parareptiles, ''Microleter'' had teeth with plicidentine, a type of internally folded
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
which is most common in "
labyrinthodont "Labyrinthodontia" (Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally consid ...
" amphibians. Although the lacrimal was not complete, the internal texture of the overlapping maxilla indicates that it extended to the nares (nostril holes), a trait also observed in millerettids,
pareiasaurs Pareiasaurs (meaning "cheek lizards") are an extinct clade of large, herbivorous parareptiles. Members of the group were armoured with scutes which covered large areas of the body. They first appeared in southern Pangea during the Middle Permian, ...
, and
bolosaurids 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. ...
. The front edge of the orbit was thick due to an internal flange on the prefrontal, akin to that of
procolophonids Procolophonidae is an extinct family of small, lizard-like parareptiles known from the Late Permian to Late Triassic that were distributed across Pangaea, having been reported from Europe, North America, China, South Africa, South America, Antarc ...
and ''
Colobomycter ''Colobomycter'' is an extinct genus of lanthanosuchoid parareptile known from the Early Permian of Oklahoma. Discovery The type species, ''Colobomycter pholeter'', was first described from fossil remains in 1958, at which time it was beli ...
''. The upper edge of the orbit had a slight contribution by the
frontal Front may refer to: Arts, entertainment, and media Films * ''The Front'' (1943 film), a 1943 Soviet drama film * ''The Front'', 1976 film Music *The Front (band), an American rock band signed to Columbia Records and active in the 1980s and ea ...
bone, though more restricted than that of lanthanosuchoids and procolophonoids. 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 ...
and particularly the
postfrontal 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, the ...
are both small and crescent-shaped. The
postorbital The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...
, by contrast, is larger and extends to near the rear edge of the skull roof. Along with the jugal, squamosal, and quadratojugal, the postorbital edges a tall, narrow opening in the rear portion of the skull. This opening, termed a ventral temporal emargination, is likely homologous to the temporal opening present in various other parareptiles. The slit-like opening of ''Microleter'' is proportionally similar to that of '' Nyctiphruretus'', although differs in the participation of the postorbital in its border. However, other parareptiles with postorbital participation (''
Australothyris ''Australothyris'' is an extinct genus of basal procolophonomorph parareptile known from the Middle Permian (middle Capitanian stage) of ''Tapinocephalus'' Assemblage Zone, South Africa. The type and only known species is ''Australothyris smith ...
'' and lanthanosuchids) have their opening fully surrounded by bone, while that of ''Microleter'' is open from below. The rear of the skull is not very long, with a broad parietal, small
supratemporal The supratemporal bone is a paired cranial bone present in many tetrapods and tetrapodomorph fish. It is part of the temporal region (the portion of the skull roof behind the eyes), usually lying medial (inwards) relative to the squamosal and latera ...
, solitary
postparietal Postparietals are cranial bones present in fish and many tetrapods. Although initially a pair of bones, many lineages possess postparietals which were fused into a single bone. The postparietals were dermal bones situated along the midline of the ...
, and tall, boxy squamosal and quadratojugal which are excavated along their rear edge. Most 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 sepa ...
is obscured by overlapping bones. The
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.
is very broad, while the transverse flange of the
pterygoid Pterygoid, from the Greek for 'winglike', may refer to: * Pterygoid bone, a bone of the palate of many vertebrates * Pterygoid processes of the sphenoid bone ** Lateral pterygoid plate ** Medial pterygoid plate * Lateral pterygoid muscle * Medial ...
is oriented forwards. What can be seen of the braincase indicates that ''Microleter'' had tubular paroccipital processes, rather than fan-shaped ones present in other parareptiles. The elongated
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
is ornamented with small pits, but its teeth are obscured and cannot be properly counted. The
surangular The suprangular or surangular is a jaw bone found in most land vertebrates, except mammals. Usually in the back of the jaw, on the upper edge, it is connected to all other jaw bones: dentary, angular, splenial and articular The articular bone i ...
possesses a folded ridge on its outer surface, and encompasses the front half of a hole at the rear of the jaw. The rear half of the hole is edged by the
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 ...
bone. ''Microleter'' is one of the few parareptiles to have preserved part of the
sclerotic ring Sclerotic rings are rings of bone found in the eyes of many animals in several groups of vertebrates, except for mammals and crocodilians. They can be made up of single bones or multiple segments and take their name from the sclera. They are bel ...
, which was formed by tall, concave plates. A few
cervical vertebrae In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In ...
are the only fossilized postcranial elements, but they are poorly preserved.


Classification

''Microleter'' is one of the most basal parareptiles, and was originally described as the most basal parareptile from
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...
. Before the description of ''Microleter'', parareptiles were hypothesized to have originated in
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
. However, ''Microleter'' appears in Laurasia soon after the earliest known parareptiles, and spurred discussion supporting the origin of parareptiles in Laurasia. Other factors in support of a Laurasian origin are the early appearance of the Laurasian bolosaurians, the fact that the other two clades of
amniotes Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are distingu ...
,
synapsids 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 Sauropsida, sauropsids, the group that inc ...
and
eureptiles 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-C ...
, are both considered to have been Laurasian in origin, and evidence from
varanopids Varanopidae is an extinct family of amniotes that resembled monitor lizards and may have filled a similar niche, hence the name. Typically, they are considered synapsids that evolved from an '' Archaeothyris''-like synapsid in the Late Carbonife ...
that colonization of Gondwana by Laurasian amniotes was more common than the reverse process. The original description of ''Microleter'' used
maximum parsimony In phylogenetics, maximum parsimony is an optimality criterion under which the phylogenetic tree that minimizes the total number of character-state changes (or miminizes the cost of differentially weighted character-state changes) is preferred. ...
and
bayesian Thomas Bayes (/beɪz/; c. 1701 – 1761) was an English statistician, philosopher, and Presbyterian minister. Bayesian () refers either to a range of concepts and approaches that relate to statistical methods based on Bayes' theorem, or a followe ...
phylogenetic analyses 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 position it more derived than
millerettid Millerettidae is an extinct family of parareptiles from the Middle Permian to the Late Permian period ( Capitanian - Changhsingian stages) of South Africa. The millerettids were small insectivores and probably resembled modern lizards in appearan ...
s and less
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
than lanthanosuchoids. In the parsimony analysis, it was in a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with ''
Australothyris ''Australothyris'' is an extinct genus of basal procolophonomorph parareptile known from the Middle Permian (middle Capitanian stage) of ''Tapinocephalus'' Assemblage Zone, South Africa. The type and only known species is ''Australothyris smith ...
'' and more derived taxa, which was resolved in the bayesian analysis to place ''Australothyris'' more basally. Cladogram from the bayesian analysis of Tsuji, Muller, & Reisz (2010): However, since the original description, various other analyses have each had slightly different conclusions on the position of ''Microleter'', placing it as 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 ''Australothyris'', basal to ''Australothyris'', or as a basal
ankyramorph Ankyramorpha ("anchor forms") is an extinct clade of procolophonomorph parareptiles which lived between the early Cisuralian epoch (middle Sakmarian stage) to the latest Triassic period (latest Rhaetian stage) of Africa, Antarctica, Asia, Aus ...
more derived than lanthanosuchoids,


References

{{Taxonbar, from=Q6839611 Procolophonomorphs Fossil taxa described in 2010 Extinct animals of the United States Permian reptiles of North America Prehistoric reptile genera