Fedexia
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''Fedexia'' 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 carnivorous
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
within the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Trematopidae Trematopidae is a family of dissorophoid temnospondyls spanning the late Carboniferous to the early Permian. Together with Dissorophidae, the family forms Olsoniformes, a clade comprising the medium-large terrestrial dissorophoids. Trematopids a ...
. It lived 300 million years ago during the late
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
period. It is estimated to have been long, and likely resembled a
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
.Pitt student finds fossil of ancient amphibian
Pittsburgh Post-Gazette, March 15, 2010 ''Fedexia'' is known from a single skull found in Moon Township,
Pennsylvania Pennsylvania, officially the Commonwealth of Pennsylvania, is a U.S. state, state spanning the Mid-Atlantic (United States), Mid-Atlantic, Northeastern United States, Northeastern, Appalachian, and Great Lakes region, Great Lakes regions o ...
. It is named after the shipping service
FedEx FedEx Corporation, originally known as Federal Express Corporation, is an American Multinational corporation, multinational Conglomerate (company), conglomerate holding company specializing in Package delivery, transportation, e-commerce, and ...
, which owned the land where the
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 ...
specimen was first found.


Discovery

The holotype skull (CM 76867) was discovered on land owned by the FedEx Corporation near the
Pittsburgh International Airport Pittsburgh International Airport —originally Greater Pittsburgh Airport and later Greater Pittsburgh International Airport—is a civil-military international airport in Findlay Township and Moon Township, Pennsylvania, United States. Abou ...
in 2004. Adam Striegel, a student at the
University of Pittsburgh The University of Pittsburgh (Pitt) is a Commonwealth System of Higher Education, state-related research university in Pittsburgh, Pennsylvania, United States. The university is composed of seventeen undergraduate and graduate schools and colle ...
, found the skull on a field trip to the area, but he mistook the teeth for a
fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
frond A frond is a large, divided leaf. In both common usage and botanical nomenclature, the leaves of ferns are referred to as fronds and some botanists restrict the term to this group. Other botanists allow the term frond to also apply to the lar ...
. It was later recognized as a skull by class lecturer Dr. Charles Jones, and was taken to the
Carnegie Museum of Natural History The Carnegie Museum of Natural History (abbreviated as CMNH) is a natural history museum in the Oakland (Pittsburgh), Oakland neighborhood of Pittsburgh, Pennsylvania. It was founded by List of people from the Pittsburgh metropolitan area, Pit ...
for further study. The specific name of the type species honors Adam Striegel. The area where the holotype was discovered was part of the Casselman Formation of the
Conemaugh Group The Conemaugh Group is a Group (geology), geologic group in West Virginia, Pennsylvania, Ohio, and Maryland. It preserves fossils dating back to the Carboniferous Period (geology), period. See also * List of fossiliferous stratigraphic units i ...
. In North American Carboniferous stratigraphy, these strata are early Virgilian in age. According to ICS standards, they are
Gzhelian The Gzhelian ( ) is an age in the ICS geologic time scale or a stage in the stratigraphic column. It is the youngest stage of the Pennsylvanian, the youngest subsystem of the Carboniferous. The Gzhelian lasted from to Ma. It follows the Ka ...
in age. The skull itself was found lying near the base of a road cut that exposes the lower part of the Casselman Formation. The Casselman Formation overlies the Ames Limestone, which represents the last marine submergence of the
Appalachian Basin The geology of the Appalachians dates back more than 1.2 billion years to the Mesoproterozoic era when two continental cratons collided to form the supercontinent Rodinia, 500 million years prior to the development of the range during the form ...
. The earliest portions of the Casselman Formation, consisting of
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 ...
s and
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility. Although its permeabil ...
s, are overbank
levee A levee ( or ), dike (American English), dyke (British English; see American and British English spelling differences#Miscellaneous spelling differences, spelling differences), embankment, floodbank, or stop bank is an elevated ridge, natural ...
deposits that were formed at the shoreline of the retreating Ames Sea. Overlying these deposits is the Grafton Sandstone, a
cross-bedded In geology, cross-bedding, also known as cross-stratification, is layering within a stratum and at an angle to the main bedding plane. The sedimentary structures which result are roughly horizontal units composed of inclined layers. The original ...
channel-phase sandstone deposited by a meandering river. Above the sandstone lies the Birmingham Shale, which includes fine-grained siltstone and
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of Clay mineral, clay minerals (hydrous aluminium phyllosilicates, e.g., Kaolinite, kaolin, aluminium, Al2Silicon, Si2Oxygen, O5(hydroxide, OH)4) and tiny f ...
overbank deposits. A thick
paleosol In Earth science, geoscience, paleosol (''palaeosol'' in Great Britain and Australia) is an ancient soil that formed in the past. The definition of the term in geology and paleontology is slightly different from its use in soil science. In geo ...
is present at the top of the Birmingham Shale, and it is capped by
limestone Limestone is a type of carbonate rock, carbonate sedimentary rock which is the main source of the material Lime (material), lime. It is composed mostly of the minerals calcite and aragonite, which are different Polymorphism (materials science) ...
that is of freshwater
lacustrine A lake is often a naturally occurring, relatively large and fixed body of water on or near the Earth's surface. It is localized in a basin or interconnected basins surrounded by dry land. Lakes lie completely on land and are separate from t ...
(lake) origin. The holotype skull likely came from the limestone that caps the Birmingham Shale or the underlying
paleosol In Earth science, geoscience, paleosol (''palaeosol'' in Great Britain and Australia) is an ancient soil that formed in the past. The definition of the term in geology and paleontology is slightly different from its use in soil science. In geo ...
. Because it was not found ''in situ'', the horizon in which the skull originated cannot be determined definitively. The gray limestone
matrix Matrix (: matrices or matrixes) or MATRIX may refer to: Science and mathematics * Matrix (mathematics), a rectangular array of numbers, symbols or expressions * Matrix (logic), part of a formula in prenex normal form * Matrix (biology), the m ...
that is found inside the skull has been examined by geochemical X-ray reflectance analyses. However, these analyses were not able to determine the stratigraphic origin of the skull.
Carbonate A carbonate is a salt of carbonic acid, (), characterized by the presence of the carbonate ion, a polyatomic ion with the formula . The word "carbonate" may also refer to a carbonate ester, an organic compound containing the carbonate group ...
nodules Nodule may refer to: *Nodule (geology), a small rock or mineral cluster *Manganese nodule, a metallic concretion found on the seafloor *Nodule (medicine), a small aggregation of cells *Root nodule Root nodules are found on the roots of plants, p ...
at the base of the Clarksburg Member and the limestone capping the Birmingham Shale were the most likely sources of the skull based on these analyses, although chemical dissimilarities with the skull matrix existed in both these strata. According to the original describers of ''Fedexia'', the carbonate nodules of the Clarksburg Member were an unlikely source because of
biogenic A biogenic substance is a product made by or of life forms. While the term originally was specific to metabolite compounds that had toxic effects on other organisms, it has developed to encompass any constituents, secretions, and metabolites of p ...
disturbances and an abundance of
siliciclastic Siliciclastic (or ''siliclastic'') rocks are clastic noncarbonate sedimentary rocks that are composed primarily of silicate minerals, such as quartz or clay minerals. Siliciclastic rock types include mudrock, sandstone Sandstone is a Clastic ...
material. The holotype skull is long and is uncrushed and well preserved. Remarkably, the
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other tetrapods 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 f ...
, or middle ear bone, remains intact. Also preserved are both
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
s and the
atlas An atlas is a collection of maps; it is typically a bundle of world map, maps of Earth or of a continent or region of Earth. Advances in astronomy have also resulted in atlases of the celestial sphere or of other planets. Atlases have traditio ...
-
axis An axis (: axes) may refer to: Mathematics *A specific line (often a directed line) that plays an important role in some contexts. In particular: ** Coordinate axis of a coordinate system *** ''x''-axis, ''y''-axis, ''z''-axis, common names ...
complex, which connects to the base of the skull.


Description

''Fedexia'' possesses several characteristics that identify it as a trematopid. One of the most noticeable is a greatly elongated and subdivided external
naris A nostril (or naris , : nares ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbinates, ...
. The posterior portion of the naris may have held a
salt gland The salt gland is an organ (anatomy), organ for excreting excess salt (chemistry), salts. It is found in the cartilaginous fishes subclass elasmobranchii (sharks, rays, and skates), seabirds, and some reptiles. Salt glands can be found in the r ...
. Another characteristic is the presence of large
palatal 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 ...
tusks. Based on small bony elements present in the related trematopid ''
Anconastes ''Anconastes'' is an extinct genus of dissorophoid temnospondyl within the family Trematopidae. It is known from two specimens from the Late Carboniferous Cutler Formation of north-central New Mexico in the southwestern United States. The genus n ...
'', ''Fedexia'' likely had a granular skin texture formed by bony protuberances. These protuberances would have served as protection from predators and would have lessened water loss through the skin. Unlike other trematopids, ''Fedexia'' has a tall, arching lateral profile to the skull. The margin forms a broad arc from the tip of the snout to the
occipital 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 lies over the occipital lobes of the cere ...
region. Other trematopids have a lower, straighter profile to the skull. Another distinguishing feature is that the postorbital length, from the eyes to the back of the skull, is greater than the preorbital length, from the eyes to the tip of the snout. The
orbits In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an physical body, object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an satellite, artificia ...
, or eye sockets, are vertically elongated, extending dorsally to the
skull table The skull roof or the roofing bones of the skull are a set of bones covering the brain, eyes and nostrils in bony fishes, including land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparati ...
. There is also a small notch on 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 mammals h ...
that separates the premaxillary teeth from the maxillary teeth.


Classification

Below is a
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
showing the
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
relations of ''Fedexia'', from Berman ''et al.'', 2010:


Paleobiology

Like all trematopids, ''Fedexia'' was well adapted to a terrestrial lifestyle. Trematopids are the earliest examples of vertebrate life in North America that lived mostly on land, although they likely returned to the water to mate and lay eggs. ''Fedexia'' is one of the oldest known vertebrates to have been primarily terrestrial rather than aquatic. Even older are the trematopids ''
Actiobates ''Actiobates'' is an extinct genus of trematopid temnospondyl that lived during the Late Carboniferous. It is known from the Garnett Quarry in Kansas. History of study ''Actiobates peabodyi'' was named in 1973 by Theodore Eaton. The genus name ...
'' and ''Anconastes'', which are Missourian (
Kasimovian The Kasimovian is a geochronologic age or chronostratigraphic stage in the ICS geologic timescale. It is the third stage in the Pennsylvanian (late Carboniferous), lasting from to Ma.; 2004: ''A Geologic Time Scale 2004'', Cambridge Unive ...
) and early Virgilian in age, respectively. The shift toward a terrestrial lifestyle may have been a response to the warmer and drier global climate that existed during the late Carboniferous. During this time, glaciation caused the climate to rapidly fluctuate and sea levels to drop. This resulted in a loss of coal swamps that were once common at higher latitudes. ''Fedexia'' was an early example of adaptation to the changing climate. It is possible that terrestrial trematopids existed for a few million years prior to ''Fedexia'', ''Actiobates'', and ''Anconastes''. Earlier trematopids may have inhabited upland regions, and would have later immigrated to lowland and coastal regions as the climate continued to change. Some 20 million years after the appearance of ''Fedexia'', during the
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
period, trematopids and other terrestrial vertebrates experienced rapid diversification.


References


External links


Student Fossil Find Is New Species
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CBS News CBS News is the news division of the American television and radio broadcaster CBS. It is headquartered in New York City. CBS News television programs include ''CBS Evening News'', ''CBS Mornings'', news magazine programs ''CBS News Sunday Morn ...
, November 9, 2004 {{Taxonbar, from=Q2849581 Dissorophidae Trematopidae Carboniferous temnospondyls of North America Carboniferous geology of Pennsylvania Fossil taxa described in 2010 Prehistoric amphibian genera FedEx