Neopteroplax
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''Neopteroplax'' 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 eogyrinid
embolomere Embolomeri is an Order (biology), order of Tetrapod, tetrapods or Stem-group, stem-tetrapods, possibly members of Reptiliomorpha. Embolomeres first evolution, evolved in the Early Carboniferous (Mississippian age, Mississippian) Period and were th ...
closely related to
European European, or Europeans, or Europeneans, may refer to: In general * ''European'', an adjective referring to something of, from, or related to Europe ** Ethnic groups in Europe ** Demographics of Europe ** European cuisine, the cuisines of Europe ...
genera such as ''
Eogyrinus ''Pholiderpeton'' (from el, φολῐ́δος , 'horny scale' and el, ἑρπετόν , 'creeping thing') is an extinct genus of embolomere amphibian which lived in the Late Carboniferous period ( Bashkirian) of England. The genus was first ...
'' and '' Pteroplax''. Members of this genus were among the largest embolomeres (and
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
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 ...
s in general) 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 ...
. ''Neopteroplax'' is primarily known from a large (~40 cm) skull found in
Ohio Ohio () is a state in the Midwestern region of the United States. Of the fifty U.S. states, it is the 34th-largest by area, and with a population of nearly 11.8 million, is the seventh-most populous and tenth-most densely populated. The sta ...
, although fragmentary embolomere fossils from Texas and New Mexico have also been tentatively referred to the genus. Despite its similarities to specific European embolomeres, it can be distinguished from them due to a small number of skull and jaw features, most notably a lower surangular at the upper rear portion of the lower jaw.


Discovery

The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
, ''Neopteroplax conemaughensis'', is known from a large skull found in Late
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especial ...
during railroad renovations in
Bloomingdale, Ohio Bloomingdale is a village in central Jefferson County, Ohio, United States. The population was 145 at the 2020 census. It is part of the Weirton–Steubenville metropolitan area. Bloomingdale is the site of a Camaldolese monastic community at H ...
. Although damaged by
excavator Excavators are heavy construction equipment consisting of a boom, dipper (or stick), bucket and cab on a rotating platform known as the "house". The house sits atop an undercarriage with tracks or wheels. They are a natural progression fro ...
s, most of the left side of the skull can be reconstructed based on surviving fragments. The only other ''Neopteroplax'' fossils recovered from this site were a few rib and vertebra fragments. The site was geologically slightly younger than the famous Carboniferous deposits of the nearby Linton diamond coal mine. Other fossils found near the ''Neopteroplax'' skull include fragments of fin spines from '' Edaphosaurus'' and several small amphibians (including a ''
Platyhystrix ''Platyhystrix'' (from el, πλατύς , 'flat' and el, ῠ̔́στρῐξ , 'porcupine') was a temnospondyl amphibian with a distinctive sail along its back, similar to the unrelated synapsids, ''Dimetrodon'' and ''Edaphosaurus''. It lived du ...
''-like form), as well as teeth from early chondrichthyan fish like '' Cladodus'', petalodonts, and cochliodonts. The skull was described, reconstructed and named by Alfred Sherwood Romer in 1963, as part of a study on various new Carboniferous embolomere fossils from Eastern North America. Citing similarities to certain European embolomeres (often referred to the genera '' Pteroplax'' or ''Eogyrinus''), he named the skull ''Neopteroplax conemaughensis'', in reference to both ''Pteroplax'' as well as the geological formation which the skull was found in (the Conemaugh series of Ohio). Jaw fragments and vertebrae similar to those of ''N. conemaughensis'' have also been recovered from the Thrifty Formation of
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
, which lies close to the border between the Carboniferous and
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
periods. These Texan remains were assigned to the second species of ''Neopteroplax'', ''Neopteroplax relictus''. These two species are mainly differentiated by their geographical and temporal isolation. Large embolomerous vertebrae and jaws from the
Wild Cow Formation The Atrasado Formation is a geologic formation in New Mexico. Its fossil assemblage dates the formation to the Kasimovian age of the Pennsylvanian. It was formerly known locally as the Wild Cow Formation or the Guadelupe Box Formation. Descript ...
of
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...
have also been tentatively assigned to ''Neopteroplax'' by
Lucas Lucas or LUCAS may refer to: People * Lucas (surname) * Lucas (given name) Arts and entertainment * Luca Family Singers, also known as "lucas ligner en torsk" * ''Lucas'' (album) (2007), an album by Skeletons and the Kings of All Cities * ''L ...
''et al.'' (1999). Some sources, such as Marjanovic & Laurin (2019), have doubted the validity of these western remains' referral to the genus.


Description

Counting the enlarged cheek regions, the skull was 39.5 centimeters (15.6 inches) in length. The snout was moderately long and the skull as a whole was deep, although crushing has reduced this depth and splayed out the cheeks, making the fossil wider than it would have been when alive. The overall shape and texture was similar to that of most other embolomeres, although there is no trace of lateral line canals. Sutures between skull bones are difficult to interpret with certainty, but when visible, their arrangement and structure is similar to that of other embolomeres (particularly
English English usually refers to: * English language * English people English may also refer to: Peoples, culture, and language * ''English'', an adjective for something of, from, or related to England ** English national ide ...
specimens commonly referred to '' Pteroplax'' or ''
Eogyrinus ''Pholiderpeton'' (from el, φολῐ́δος , 'horny scale' and el, ἑρπετόν , 'creeping thing') is an extinct genus of embolomere amphibian which lived in the Late Carboniferous period ( Bashkirian) of England. The genus was first ...
''). For example, the lacrimal bone is excluded from the front edge of the
orbit 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 socket) due to a downward branch of the
prefrontal bone The prefrontal bone is a bone separating the lacrimal and frontal bones in many tetrapod skulls. It first evolved in the sarcopterygian clade Rhipidistia, which includes lungfish and the Tetrapodomorpha. The prefrontal is found in most modern and ...
, and the broad squamosal bone is only weakly attached to skull bones above it. The orbits were roughly circular, in contrast with a well-preserved embolomere skull from Newsham (initially referred to ''
Anthracosaurus ''Anthracosaurus'' is an extinct genus of embolomere, a possible distant relative of reptiles that lived during the Late Carboniferous (around 310 million years ago) in what is now Scotland and England. It was a large, aquatic eel-like predator a ...
'' and now considered the lectotype of ''Eogyrinus''), which has more triangular orbits due to the shape of the
jugal bone 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 ...
, which forms the lower rim of the orbit. On the other hand, the jugal of ''Neopteroplax'' otherwise resembles that of the Newsham skull due to extending down far enough that it wraps under the rim of the mouth. The
premaxilla 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 ...
(at the tip of the snout) had three teeth, with the first two rather large and the last one much smaller. The following
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 ...
bone had about 40 teeth, which were generally small except for a "canine" region of four or five larger teeth near the front part of the bone. All of the teeth were only preserved as broken stumps at best, so specific information about their shape is unobtainable. The front part of the palate (roof of the mouth) has large, elliptical
choanae The choanae (singular choana), posterior nasal apertures or internal nostrils are two openings found at the back of the nasal passage between the nasal cavity and the throat in tetrapods, including humans and other mammals (as well as crocodilia ...
(internal nostrils) separated by narrow, toothless vomer bones. Most of the palate was formed by the plate-like pterygoid bones, which were covered with tiny denticles. The outer edge of each pterygoid connected to a small, rectangular palatine bone and the longer ectopterygoid, which lies immediately behind it. The palatine and ectopterygoid possessed very large fangs, with two on the palatine and six on the ectopterygoid. Very little of the braincase is well preserved, with the exception of the blade-like parasphenoid which extends forwards along the midline of the skull and divides most of the pterygoids from each other. The lower jaw of ''Neopteroplax'' possessed typical embolomere features such as two large meckelian fenestra (holes visible on the inner surface of the jaw), but it also has a few unique and diagnostic traits. It was approximately 34.2 cm (13.5 in) long, deep towards the rear and tapering and curving upwards towards the front. However, the rear portion of the jaw is not as deep as in the European embolomeres. This is because the surangular bone, which forms the upper portion of the jaw behind the tooth row, was only gently convex along its upper edge, rather than strongly arched with a pronounced crest. The outer surface of the jaw has an pronounced lateral line canal which becomes more indistinct towards the jaw joint. Like the rest of the skull, tooth preservation on the jaw is too poor to conclude much about its dentition. However, it is clear that the first of the coronoids (a series of bones adjacent and inwards of the main tooth row) was toothless, in contrast to ''
Eobaphetes ''Eobaphetes'' is an extinct genus of Embolomeri, embolomere which likely lived in the Pennsylvanian (geology), Pennsylvanian (late Carboniferous) of Kansas. The genus is based on several skull and jaw fragments of a single individual. They were ...
'' and ''Eogyrinus'', which have denticles on the first coronoid.


References

Embolomeres Carboniferous amphibians {{paleo-amphibian-stub