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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 of
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
procolophonomorph parareptiles 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 Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * Ty ...
and only known species is ''Microleter mckinzieorum''. ''Microleter'' is one of several parareptile taxa described from the Richards Spur fissure fills, and can be characterized from its high tooth count, lacrimal/ narial contact, short postfrontal, and slit-like temporal emargination edged by the postorbital, jugal, squamosal, and quadratojugal. Contrary to '' Australothyris'', which had a similar phylogenetic 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 specimen. It was found at the Dolese Brothers limestone quarry near Richards Spur 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, ~289-286 Ma) fissure fills in an Ordovician cave system. The fissure fills have yielded many other well preserved tetrapod 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 Linda may refer to: As a name * Linda (given name), a female given name (including a list of people and fictional characters so named) * Linda (singer) (born 1977), stage name of Svetlana Geiman, a Russian singer * Anita Linda (born Alice Lake ...
, Johannes Muller, and
Robert R. Reisz Robert Rafael Reisz is a Canadian paleontologist and specialist in the study of early amniote and tetrapod evolution. Research career Reisz received his B.Sc. (1969), M.Sc. (1971) and Ph.D. (1975) from McGill University as Robert L. Carroll ...
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 Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
honors the McKinzie family, as the specimen was discovered by Mark McKinzie.


Description

Based on the skull's large orbits (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, squamosal, and quadratojugal. 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''. Like several other basal parareptiles, ''Microleter'' had teeth with plicidentine, a type of internally folded dentine 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, 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 and '' Colobomycter''. The upper edge of the orbit had a slight contribution by the frontal bone, though more restricted than that of lanthanosuchoids and procolophonoids. The jugal and particularly the postfrontal are both small and crescent-shaped. The postorbital, 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 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 ...
present in various other parareptiles. The slit-like opening of ''Microleter'' is proportionally similar to that of ''
Nyctiphruretus ''Nyctiphruretus'' (meaning "Guardian of the Night") is an extinct genus of nyctiphruretid parareptile known from the Guadalupian series (middle Permian) of European Russia. Many fossils of the type species, ''N. acudens'', were found well pre ...
'', although differs in the participation of the postorbital in its border. However, other parareptiles with postorbital participation ('' Australothyris'' 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, solitary postparietal, and tall, boxy squamosal and quadratojugal which are excavated along their rear edge. Most of the palate is obscured by overlapping bones. The palatine is very broad, while the transverse flange of the pterygoid 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 is ornamented with small pits, but its teeth are obscured and cannot be properly counted. The surangular 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 bone. ''Microleter'' is one of the few parareptiles to have preserved part of the sclerotic ring, 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 Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
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, are both considered to have been Laurasian in origin, and evidence from varanopids that colonization of Gondwana by Laurasian amniotes was more common than the reverse process. The original description of ''Microleter'' used maximum parsimony and bayesian phylogenetic analyses to position it more derived than millerettids and less derived than lanthanosuchoids. In the parsimony analysis, it was in a polytomy with '' Australothyris'' 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 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, Austral ...
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