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''Psarolepis'' (; ''psārolepis'', from Greek ψαρός 'speckled' and λεπίς 'scale') is a
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 extinct
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
which lived around 397 to 418
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
( Pridoli to
Lochkovian The Lochkovian is one of three faunal stages in the Early Devonian Epoch. It lasted from 419.2 ± 3.2 million years ago to 410.8 ± 2.8 million years ago. It marked the beginning of the Devonian Period, and was followed by the Pragian Stage. It i ...
stages Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * S ...
). Fossils of ''Psarolepis'' have been found mainly in South
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
and described by
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Xiaobo Yu in 1998. It is not known certainly in which group ''Psarolepis'' belongs, but paleontologists agree that it probably is a basal genus and seems to be close to the common ancestor of lobe-finned and
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or hor ...
es.Benton, M. J. (2005): ''Vertebrate Palaeontology'', page 65. In 2001, paleontologist John A. Long compared ''Psarolepis'' with
onychodontiform Onychodontiformes (also known as Onychodontida and Struniiformes) is an order of prehistoric sarcopterygian fish that lived during the Devonian period. The onychodontiforms are generally regarded as early-diverging members of the coelacanth ...
fishes and refer to their relationships.


Description

''Psarolepis'' had a pair of 'parasymphysical tooth whorls', teeth which extend up at the front of the
lower jaw In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
. The head was made of several thick dermal plates and covered with deep pock-marks and large pores. Another trait is a large pectoral spine, just in front of the pectoral fin, extending back from the
shoulder 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 ...
, and a dorsal spine located in front of a median fin behind the head, which gives the fish a
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
-like form. The pock-marked head of ''Psarolepis'' was made of plates containing a layer of
porcelain Porcelain () is a ceramic material made by heating substances, generally including materials such as kaolinite, in a kiln to temperatures between . The strength and translucence of porcelain, relative to other types of pottery, arises mainl ...
-like
cosmine Cosmine is a spongy, bony material that makes up the dentine-like layers in the scales of the lobe-finned fishes of the class Sarcopterygii. Fish scales that include layers of cosmine are known as cosmoid scales. Description As traditionally des ...
. Because the cosmine layer obscures the suture lines of the skull, it is difficult to study the exact bone structure. The snout was strangely humped and the
nostrils A nostril (or naris , plural ''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 turbi ...
were located above the eyes, which were just above the
upper jaw 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 ...
. The most spectacular findings were the fin spines. Two are known: one extending back from the
shoulder 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 ...
and another which is associated with the
dorsal fin A dorsal fin is a fin located on the back of most marine and freshwater vertebrates within various taxa of the animal kingdom. Many species of animals possessing dorsal fins are not particularly closely related to each other, though through conv ...
. These fin spines are found only in primitive jawed fishes and are apparently absent from the most primitive sharks, but present in abundance in more derived forms. ''Psarolepis'' had teeth at the very front of the snout with large fangs on the tooth plate. Outstanding feature are the 'parasymphysical tooth whorls' which place the fish in the order of
Onychodontida Onychodontiformes (also known as Onychodontida and Struniiformes) is an order of prehistoric sarcopterygian fish that lived during the Devonian period. The onychodontiforms are generally regarded as early-diverging members of the coelacanth ...
.Zhu; Yu; Janvier, 1999. 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 ...
and the
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
had large inner teeth and irregular array of tiny outer teeth.


Discovery

Most of the fossils of ''Psarolepis'' were discovered at the same locality of the Yulongsi, Xishancun and
Xitun Formation The Xitun Formation is a palaeontological formation which is named after Xitun village in Qujing, a location in South China. This formation includes many remains of fossilized fish and plants of the Early Devonian period (Late Lochkovian). It wa ...
s, about northwest of the city of Qujing,
Yunnan, China Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the ...
. With this discovery other prehistoric lobe-finned fishes came to light such as ''
Youngolepis ''Youngolepis'' is a genus of prehistoric lobe-finned fish which lived during the Early Devonian period (Lochkovian to Pragian stages, about 407-416 million years ago). Fossils of ''Y. praecursor'' have been found in the Xitun Formation of ...
'' and ''
Diabolepis ''Diabolepis'' (or ''Diabolichthys'') is an extinct genus of very primitive lungfish which lived about 400 million years ago, in the Early Devonian period of South China. ''Diabolepis'' is the most basal known dipnoan. A rather small fish, the ...
''. The remains of ''Psarolepis'' were collected in 1981 and 1984 by paleontologist Min Zhu and colleagues and dated from
Early Devonian The Early Devonian is the first of three Epoch (geology), epochs comprising the Devonian period, corresponding to the Lower Devonian Series (stratigraphy), series. It lasted from and began with the Lochkovian Stage , which was followed by the P ...
and Late
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
periods.Ahlberg, 1999. Other fossils, also dated from Late Silurian, have been found in
Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
but the description of the fish was based on the materials from China because they were better preserved.


History and classification

When ''Psarolepis'' was described for the first time in 1998, it was placed in the group of sarcopterygians (lobe-finned fishes) because the skull and jaws resemble those of primitive
lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
es. In 1999 Zhu et al. were unable to locate ''Psarolepis'' in the
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
because they did not know if it was the most primitive lobed-finned fish or the most primitive bony fish. There are some characteristics that bony fish do not have, such as the median spine located behind the head, which is known in
sharks Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimorp ...
and
acanthodians Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes), typically considered a paraphyletic group. They are currently considered to represent a grade of various fish lineages leading up to the extant Chondrichthyes, which ...
, and the pectoral spine extending back from the shoulder girdle which is found in some
placoderms Placodermi (from Greek πλάξ 'plate' and δέρμα 'skin', literally 'plate-skinned') is a class of armoured prehistoric fish, known from fossils, which lived from the Silurian to the end of the Devonian period. Their head and thorax were ...
and acanthodians. Later, in 2001, Zhu and Schultze gave more basis to the theory that ''Psarolepis'' was probably a basal bony fish. The same year Long re-examined the phylogenetic position of the fish and pointed out several similarities between ''Psarolepis'' and Onychodontiform fishes.Long, 2001. He also noted that the presence of a rotational tooth whorl combined with the other characteristics in the skull, and possibly in the shoulder girdle, show that ''Psarolepis'' is better placed as a sister taxon to ''
Onychodus ''Onychodus'' (, from Greek meaning "claw-tooth") is a genus of prehistoric lobe-finned fish which lived during the Devonian period (Eifelian - Famennian stages, around 374 to 397 million years ago). It is one of the best known of the group of ...
'' as the most basal member of the group of Onychodontiforms. Moreover, Long, referring to new fossils collected from
Gogo Formation The Gogo Formation in the Kimberley region of Western Australia is a Lagerstätte that exhibits exceptional preservation of a Devonian reef community. The formation is named after Gogo Station, a cattle station where outcrops appear and fossils ...
,
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to th ...
, said that ''Psarolepis'' and ''Onychodus'' are both basal bony fish and are more primitive than other lobe-finned groups.


References

* * * * * *


Notes


External links


''Psarolepis'' at Palaeos




{{Taxonbar, from=Q291624 Prehistoric bony fish genera Fish enigmatic taxa Transitional fossils Pridoli life Silurian bony fish Devonian bony fish Early Devonian fish Devonian fish of Asia Silurian fish of Asia Silurian China Fossils of China Fossil taxa described in 1998 Taxa named by Xiaobo Yu