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''Ymeria'' 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 early stem tetrapod from the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
of
Greenland Greenland ( kl, Kalaallit Nunaat, ; da, Grønland, ) is an island country in North America that is part of the Kingdom of Denmark. It is located between the Arctic and Atlantic oceans, east of the Canadian Arctic Archipelago. Greenland is t ...
. Of the two other genera of stem tetrapods from Greenland, ''
Acanthostega ''Acanthostega'' (meaning "spiny roof") is an extinct genus of stem-tetrapod, among the first vertebrate animals to have recognizable limbs. It appeared in the late Devonian period (Famennian age) about 365 million years ago, and was anatomic ...
'' and ''
Ichthyostega ''Ichthyostega'' (from el, ἰχθῦς , 'fish' and el, στέγη , 'roof') is an extinct genus of limbed tetrapodomorphs from the Late Devonian of Greenland. It was among the earliest four-limbed vertebrates in the fossil record, and was on ...
'', ''Ymeria'' is most closely related to ''Ichthyostega'', though the single known specimen is smaller, the skull about 10 cm in length. A single
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 ...
resembles that of ''Ichthyostega'', an indication ''Ymeria'' may have resembled this genus in the post-cranial skeleton.


Discovery

''Ymeria'' is primarily 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 ...
skull including the lower jaws and
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 ...
, as well as impressions of the shoulder girdle. The holotype comes from the southern slope of Mt. Celsius on
Ymer Island Ymer Island ( da, Ymer Ø) is an island in northeastern Greenland. The island is a part of Northeast Greenland National Park. Ymer Island is named after the Swedish geographical journal ''Ymer'', which published many accounts of Swedish expedit ...
in northeast Greenland. Fossils of Devonian tetrapods like ''Ichthyostega'' have been known from Ymer Island since 1929. The skull of ''Ymeria'' found in 1947 by a team of paleontologists from Sweden and Denmark. It came from a
talus slope Scree is a collection of broken rock fragments at the base of a cliff or other steep rocky mass that has accumulated through periodic rockfall. Landforms associated with these materials are often called talus deposits. Talus deposits typically h ...
or pile of rock fragments at the base of Mount Celsius, encased in a pale red
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ...
. The fossil's origin on the mountain has not been identified. There are four formations preserved at Mt. Celsius, all belonging to the larger
Celsius Bjerg Group The degree Celsius is the unit of temperature on the Celsius scale (originally known as the centigrade scale outside Sweden), one of two temperature scales used in the International System of Units (SI), the other being the Kelvin scale. The d ...
. Since the skull cannot be traced to any one of these formations, its exact age is uncertain.


Classification

Swedish paleontologist
Erik Jarvik Anders Erik Vilhelm Jarvik (30 November 1907 – 11 January 1998) was a Swedish paleontologist who worked extensively on the sarcopterygian (or lobe-finned) fish ''Eusthenopteron''. In a career that spanned some 60 years, Jarvik produced some ...
, a member of the 1947 expedition, could not assign the skull to either ''Ichthyostega'' or ''Acanthostega''. In 1988, English paleontologist
Jennifer Clack Jennifer Alice Clack, (''née'' Agnew; 3 November 1947 – 26 March 2020) was an English palaeontologist and evolutionary biologist. She specialised in the early evolution of tetrapods, specifically studying the "fish to tetrapod" transition: ...
was the first to propose that the material represented a third type of tetrapod from Greenland, based mainly on the teeth. While the skull shape is closest to ''Ichthyostega'', the teeth are smaller, more numerous and less curved, indicating the two exploited different foods. The surface ornamentation on the skull bones are less pronounced, indicating lighter dermal armour. The material was assigned to a new genus and species, ''Ymeria denticulata'', in 2012. The genus is named after Ymer Island, while the specific name refers to the denticulated or bumpy surface of the lower jaw.


Description

The cranium of the holotype only preserves a few fragmentary bones. 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 mammal has b ...
(toothed bones at the tip of the snout) were present and seemingly bore indentations at the front for a median rostral. A small bone is preserved behind the right premaxilla. This bone may be a lateral rostral due to possessing a lateral line, but its shape is more similar to a tectal. Rostrals and tectals are small skull bones scattered around the snout, which are present in
tetrapodomorph The Tetrapodomorpha (also known as Choanata) are a clade of vertebrates consisting of tetrapods (four-limbed vertebrates) and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish. Advance ...
fish but lost in true tetrapods. Both
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 ...
(toothed bones at the side of the snout) are well-preserved, but not particularly specialized. The premaxillae and maxilla bear numerous tapering teeth, with about 11 on each premaxilla and up to 24 on each maxilla. This is slightly more than ''Ichthyostega'' (which has 8–10 on the premaxilla and 16–23 on the maxilla), and ''Ymeria'' further differs by having the largest teeth be slightly further forwards in the snout. Bones 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 ...
(roof of the mouth), such as the
vomers The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxil ...
,
palatines Palatines (german: Pfälzer), also known as the Palatine Dutch, are the people and princes of Palatinates ( Holy Roman principalities) of the Holy Roman Empire. The Palatine diaspora includes the Pennsylvania Dutch and New York Dutch. In 1709 ...
, ectopterygoids, and pterygoids, are poorly preserved, but similar to those of other Devonian stem-tetrapods (in terms of both shape and dentition) when visible. A sliver of bone near the cheek region may represent a branchial element (
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
bone). Preserved fragments of the shoulder girdle resemble those of ''Ichthyostega'', such as smooth
clavicles The clavicle, or collarbone, is a slender, S-shaped long bone approximately 6 inches (15 cm) long that serves as a strut between the shoulder blade and the sternum (breastbone). There are two clavicles, one on the left and one on the right ...
and a pointed rear stalk of the
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 ...
. The lower jaws were thick and well-preserved. Their external bone texturing consisted of indistinct and shallow pits and grooves. This contrasts with ''Ichthyostega'' and ''Acanthostega'', which have much more pronounced systems of ridges and pits. The jaw also has a (mostly) open
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial ...
groove on its outer surface. Like other early tetrapods and their close relatives, there were two main rows of teeth on each lower jaw. The outer (marginal) tooth row was present solely on the dentary bone, which was narrow and had alternating regions of light and absent texturing. It had at most 33 teeth, including a symphysial fang (an enlarged tooth near the chin) which was only slightly larger than the other dentary teeth. The inner tooth row stretched along four plate-like bones: the parasymphysial plate and three coronoid bones. The parasymphysial plate has a large tooth at its front edge, followed by a smaller tooth and a diastema (toothless area), similar to ''Ichthyostega''. About 22 teeth were present in a mostly unbroken row along the coronoids. The largest coronoid teeth were present at the front of the first coronoid, the middle of the second, and (to a lesser extent) about a third the way down the rear coronoid. These large teeth are the same size as the dentary teeth. In contrast, even the largest coronoid teeth of ''Ichthyostega'' are much smaller than the dentary teeth. Another unique feature of ''Ymeria'' is the presence of a large patch of tiny tooth-like denticles on the prearticular bone, which lies directly under the inner main tooth row. This patch of denticles is unknown in ''Ichthyostega'' or any of its relatives. All of ''Ymeria'''s teeth (on both the top and bottom jaws) were sharp but conical, in contrast to the recurved teeth of ''Ichthyostega''.


References

{{Taxonbar, from=Q1948602 Prehistoric tetrapod genera Devonian tetrapods Fossils of Greenland Fossil taxa described in 2012 Taxa named by Jenny Clack