Karenites
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''Karenites'' is an extinct
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
therocephalia Therocephalia is an extinct suborder of eutheriodont therapsids (mammals and their close relatives) from the Permian and Triassic. The therocephalians ("beast-heads") are named after their large skulls, which, along with the structure of their te ...
n
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
s from the
Late Permian Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
of
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
. The only species is ''Karenites ornamentatus'', named in 1995. Several fossil specimens are known from the town of
Kotelnich Kotelnich (russian: Коте́льнич; chm, Кäкшäр) is a river port town in Kirov Oblast, Russia, located on the right bank of the Vyatka River near its confluence with the Moloma, along the route of the Trans-Siberian Railway, southwe ...
in
Kirov Oblast Kirov Oblast (russian: Ки́ровская о́бласть, ''Kirovskaya oblast'') is a federal subject of Russia (an oblast) in Eastern Europe. Its administrative center is the city of Kirov. Population: 1,341,312 ( 2010 Census). Geography Na ...
.


Description

''Karenites'' is known from a partial
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 ...
skeleton, two partial skulls, and isolated jaw bones. Although incomplete, the skulls preserve small and delicate structures like nasal turbinates on the inside of the skull and the
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foot ...
bone of the ear. The skull of ''Karenites'' is about long, with the snout much longer than the temporal region of the skull behind the eye sockets. Viewed from above, the skull is triangular. The snout is broad, and the skull widens toward the
occiput The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobes of the cereb ...
or posterior margin. Two large holes behind the eye socket called
temporal fenestra An infratemporal fenestra, also called the lateral temporal fenestra or simply temporal fenestra, is an opening in the skull behind the orbit in some animals. It is ventrally bordered by a zygomatic arch. An opening in front of the eye sockets, ...
e occupy most of the posterior skull. Between these fenestra is a narrow
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
. In front of this crest, the
skull roof The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparati ...
bones are weakly pitted with small bumps and ridges for blood vessels. Some specimens include parts 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 ...
, a ring of bone embedded in the eye. On each side of the upper jaw are five incisors, two or three precanines, one canine, and eleven or twelve postcanines. The incisors and precanines are long, thin, and slightly curved, separated from each other by a small gap. The canine is much longer, projecting slightly forward from the tooth socket and curving backward along its length. The postcanine teeth are shorter and broader than the incisors and precanines. Toward the back of the skull the tips of the teeth flatten. The lower jaw is thin and curves upward to the arch of the cheek, except for a large
coronoid process The Coronoid process (from Greek , "like a crown") can refer to: * The coronoid process of the mandible, part of the ramus mandibulae of the mandible * The coronoid process of the ulna The coronoid process of the ulna is a triangular process proj ...
that extends to 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 ...
-
quadrate Quadrate may refer to: * Quadrate bone * Quadrate (heraldry) * Quadrate lobe of liver * Quadrate tubercle The quadrate tubercle is a small tubercle found upon the upper part of the femur. It serves as a point of insertion of the quadratus femori ...
jaw joint at the back of the skull. The lower jaw has three small incisors angled slightly forward, a large canine tooth projecting upward, and thirteen small, blunt postcanine teeth. The farthest postcanine teeth have small secondary cusps behind their tips. These multicusped teeth may have been adaptations for crushing food, although they are not as well developed as the teeth of other therocephalians like ''
Ericiolacerta Left may refer to: Music * ''Left'' (Hope of the States album), 2006 * ''Left'' (Monkey House album), 2016 * "Left", a song by Nickelback from the album ''Curb'', 1996 Direction * Left (direction), the relative direction opposite of right * L ...
''. In 1999, thoracic plates were reported to be present in the holotype of ''Karenites''. Thoracic plates are plates of bone on the underside of the rib cage that are typically found in reptiles, and unusual for mammal relatives like therocephalians. This bone was later reinterpreted as an
interclavicle An interclavicle is a bone which, in most tetrapods, is located between the clavicles. Therian mammals ( marsupials and placentals) are the only tetrapods which never have an interclavicle, although some members of other groups also lack one. In th ...
, part of the
pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of t ...
common to all early therapsids.


Paleobiology

The presence of a sclerotic ring in ''Karenites'' may be an indication that it was aquatic. Pits on the skull have been interpreted as evidence for well-developed whiskers, which may have been used in hunting aquatic prey. Some therocephalians like ''
Perplexisaurus ''Perplexisaurus'' is a genus of therocephalian therapsid from the Middle to Late Permian ''Deltavjatia vjatkensis'' Assemblage Zone, Vanyushonki Member of the Urpalov Formation of Russia. It was described by L. P. Tatarinov in 1997, and the t ...
'' have also been interpreted as aquatic predators, and share many similarities with ''Karenites''. While these aquatic forms had strong sutures between cranial bones, which may have stabilized the skull when consuming large aquatic prey like fishes, ''Karenites'' had weaker, slightly movable skull joints associated with feeding on smaller terrestrial prey like insects. Its multicuspid teeth also suggest it fed on insects. ''Karenites'' has long limb bones that indicate a fully terrestrial rather than aquatic lifestyle. Ridges on the inside of the skull of ''Karenites'' form a series of
sinuses Paranasal sinuses are a group of four paired air-filled spaces that surround the nasal cavity. The maxillary sinuses are located under the eyes; the frontal sinuses are above the eyes; the ethmoidal sinuses are between the eyes and the sphenoid ...
. These sinuses may have been associated with improvements in the sense of smell of therocephalians, but they are not thought to be olfactory structures. Bony projections on the underside of the lower jaw of ''Karenites'' may have supported tissues that transmitted sound to the stapes bone in the ear. Early therapsids like ''Karenites'' lack the well-developed auditory system of mammals, which had evolved from a restructuring of bones in the back of the skull and the lower jaw, and probably had a poor sense of hearing. As an early stage in the development of the mammalian auditory system, ''Karenites'' may have been able to hear some sounds by placing its jaw on the ground to detect vibrations.


References

{{Taxonbar, from=Q3206422 Extinct animals of Russia Therocephalia genera Baurioids