Ymeria
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''Ymeria'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of early
stem tetrapod The Stem Tetrapoda are a cladistically defined group, consisting of all animals more closely related to extant four-legged vertebrates than to their closest extant relatives (the lungfish), but excluding the crown group Tetrapoda. They are thus ...
from the
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
of
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
. Of the two other genera of
stem tetrapod The Stem Tetrapoda are a cladistically defined group, consisting of all animals more closely related to extant four-legged vertebrates than to their closest extant relatives (the lungfish), but excluding the crown group Tetrapoda. They are thus ...
s from Greenland, ''
Acanthostega ''Acanthostega'', from Ancient Greek ἄκανθα (''ákantha''), meaning "spine", and στέγη (''stégē''), meaning "roof", is an extinct genus of stem tetrapoda, stem-tetrapod, among the first vertebrates, vertebrate animals to have recogn ...
'' and ''
Ichthyostega ''Ichthyostega'', from Ancient Greek ἰχθύς (''ikthús''), meaning "fish", and στέγη (''stégē''), meaning "roof", is an Extinction, extinct genus of limbed tetrapodomorpha, tetrapodomorphs from the Devonian, Late Devonian of what is ...
'', ''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 ...
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 (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
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 sep ...
, as well as impressions of the shoulder girdle. The holotype comes from the southern slope of Mt. Celsius on
Ymer Island Ymer Island () 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 expeditions to Spit ...
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. The term ''scree'' is ap ...
or pile of rock fragments at the base of Mount Celsius, encased in a pale red
sandstone Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...
. 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. Since the skull cannot be traced to any one of these formations, its exact age is uncertain.


Classification

Swedish paleontologist Erik Jarvik, a member of the 1947 expedition, could not assign the skull to either ''Ichthyostega'' or ''Acanthostega''. In 1988, English paleontologist Jennifer Clack 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 mammals has ...
(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 Tetrapodomorpha (also known as Choanata) is 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. Advanced for ...
fish but lost in true tetrapods. Both
maxillae In vertebrates, the maxilla (: maxillae ) 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 two maxillar ...
(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 sep ...
(roof of the mouth), such as the vomers,
palatines Palatines () were the citizens and princes of the Palatinates, Holy Roman States that served as capitals for the Holy Roman Emperor. After the fall of the Holy Roman Empire in 1806, the nationality referred more specifically to residents of the ...
, 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 respiration organ, respiratory organ that many aquatic ecosystem, 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 r ...
bone). Preserved fragments of the shoulder girdle resemble those of ''Ichthyostega'', such as smooth
clavicles The clavicle, collarbone, or keybone is a slender, S-shaped long bone approximately long that serves as a strut between the shoulder blade and the sternum (breastbone). There are two clavicles, one on each side of the body. The clavicle is the ...
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 ...
. 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 epithelia ...
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 Stegocephali Prehistoric lobe-finned fish genera Late Devonian sarcopterygians Devonian sarcopterygians of North America Fossils of Greenland Fossil taxa described in 2012 Taxa named by Jenny Clack