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Acanthodii or acanthodians is an extinct class of
gnathostomes Gnathostomata (; from Greek: (') "jaw" + (') "mouth") are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, living ...
(jawed
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
es), typically considered a
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
group. They are currently considered to represent a
grade Grade most commonly refers to: * Grade (education), a measurement of a student's performance * Grade, the number of the year a student has reached in a given educational stage * Grade (slope), the steepness of a slope Grade or grading may also ref ...
of various fish lineages leading up to the extant
Chondrichthyes Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
, which includes living
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 ...
s,
rays Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (gra ...
, and chimaeras. Acanthodians possess a mosaic of features shared with both osteichthyans (bony fish) and chondrichthyans (cartilaginous fish). In general body shape, they were similar to modern sharks, but their
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...
was covered with tiny rhomboid platelets like the scales of
holostei Holostei is a group of ray-finned bony fish. It is divided into two major clades, the Halecomorphi, represented by a single living species, the bowfin ('' Amia calva''), as well as the Ginglymodi, the sole living representatives being the gars ...
ans (
gar Gars are members of the family Lepisosteidae, which are the only surviving members of the Ginglymodi, an ancient holosteian group of ray-finned fish, which first appeared during the Triassic, over 240 million years ago. Gars comprise seven livin ...
s, bowfins). A lower Silurian
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, ''
Fanjingshania renovata ''Fanjingshania'' is an extinct genus of acanthodian from the lower 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 beginnin ...
'', attributed to
Climatiiformes The Climatiiformes is an order of extinct fish belonging to the class Acanthodii. Like most other "spiny sharks", the Climatiiformes had sharp spines. These animals were often fairly small in size and lived from the Late Silurian to the Early Ca ...
is the oldest chondrichthyan with known anatomical features. The popular name "spiny sharks" is because they were superficially shark-shaped, with a streamlined body, paired
fins A fin is a thin component or appendage attached to a larger body or structure. Fins typically function as foils that produce lift or thrust, or provide the ability to steer or stabilize motion while traveling in water, air, or other fluids. Fin ...
, a strongly upturned tail, and stout, largely immovable bony spines supporting all the fins except the tail—hence, "spiny sharks". However, acanthodians are not true sharks; their close relation to modern cartilaginous fish can lead them to be considered "
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
-sharks". Acanthodians had a
cartilaginous Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck and ...
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
, but their fins had a wide, bony base and were reinforced on their anterior margin with a dentine spine. As a result, fossilized spines and scales are often all that remains of these fishes in ancient
sedimentary rock Sedimentary rocks are types of rock that are formed by the accumulation or deposition of mineral or organic particles at Earth's surface, followed by cementation. Sedimentation is the collective name for processes that cause these particles ...
s. The earliest acanthodians were marine, but during 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 ...
, freshwater species became predominant. Acanthodians have been divided into four orders:
Acanthodiformes Acanthodiformes is an order of acanthodian fishes which lived from the Early Devonian to Early Permian. Subtaxa * Family Acanthodidae ** Genus ''Acanthodes ''Acanthodes'' (from el, ἄκανθώδης , 'provided with spines') is an ex ...
,
Climatiiformes The Climatiiformes is an order of extinct fish belonging to the class Acanthodii. Like most other "spiny sharks", the Climatiiformes had sharp spines. These animals were often fairly small in size and lived from the Late Silurian to the Early Ca ...
,
Diplacanthiformes Diplacanthiformes (also known as Diplacanthida, Diplacanthoidei, or Diplacanthini) is an order of acanthodian fishes which lived during the Devonian Period The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60. ...
, and
Ischnacanthiformes Ischanacanthiformes is a prehistoric order of Acanthodii or spiny sharks found in Canada, Ukraine and United Kingdom. Members of this order were nektonic carnivores, eating animals that swim rather than plankton. They had slender builds, light ar ...
. "Climatiiformes" is a paraphyletic assemblage of early acanthodians such as climatiids, gyracanthids, and diplacanthids; they had robust bony shoulder girdles and many small sharp spines ("intermediate" or "prepelvic" spines) between the pectoral and pelvic fins. The climatiiform subgroup Diplacanthida has subsequently been elevated to its own order, Diplacanthiformes. Ischnacanthiforms were predators with tooth plates fused to their jaws. Acanthodiforms were filter feeders with a single dorsal fin, toothless jaws, and long
gill raker Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of the ...
s. They were the last and most specialized off the traditional acanthodians, as they survived up until the
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 ...
period.


Characteristics

The scales of Acanthodii have distinctive ornamentation peculiar to each order. Because of this, the scales are often used in determining relative age of sedimentary rock. The scales are tiny, with a bulbous base, a neck, and a flat or slightly curved diamond-shaped crown. Despite being called "spiny sharks," acanthodians predate sharks. Scales that have been tentatively identified as belonging to acanthodians, or "shark-like fishes" have been found in various Ordovician strata, though, they are ambiguous, and may actually belong to jawless fishes such as
thelodont Thelodonti (from Greek: "feeble teeth")Maisey, John G., Craig Chesek, and David Miller. Discovering fossil fishes. New York: Holt, 1996. is a class (biology), class of extinct jawless fishes with distinctive Fish scale, scales instead of large pl ...
s. The earliest unequivocal acanthodian fossils date from the beginning of the
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 ...
Period, some 50 million years before the first sharks appeared. Later, the acanthodians colonized fresh waters, and thrived in the rivers and lakes during 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 ...
and in the
coal swamp Coal forests were the vast swathes of wetlands that covered much of the Earth's tropical land areas during the late Carboniferous ( Pennsylvanian) and Permian times.Cleal, C. J. & Thomas, B. A. (2005). "Palaeozoic tropical rainforests and their ...
s of
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 ...
. By this time
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 ...
es were already showing their potential to dominate the waters of the world, and their competition proved too much for the spiny sharks, which died out in
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 ...
times (approximately 250 million years ago). Many palaeontologists originally considered the acanthodians close to the ancestors of the bony fishes. Although their interior
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
s were made of
cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck an ...
, a bonelike material had developed in the skins of these fishes, in the form of closely fitting scales (see above). Some scales were greatly enlarged and formed a bony covering on top of the head and over the lower
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 ...
. Others developed a bony flap over the gill openings analogous to the operculum in later bony fishes. However, most of these characteristics are considered homologous characteristics derived from common placoderm ancestors, and present also in basal cartilaginous fish. Overall, the acanthodians'
jaws Jaws or Jaw may refer to: Anatomy * Jaw, an opposable articulated structure at the entrance of the mouth ** Mandible, the lower jaw Arts, entertainment, and media * Jaws (James Bond), a character in ''The Spy Who Loved Me'' and ''Moonraker'' * ...
are presumed to have evolved from the first
gill arch Branchial arches, or gill arches, are a series of bony "loops" present in fish, which support the gills. As gills are the primitive condition of vertebrates, all vertebrate embryos develop pharyngeal arches, though the eventual fate of these arc ...
of some ancestral jawless fishes that had a gill skeleton made of pieces of jointed cartilage.


Taxonomy and phylogeny

In a study of early jawed vertebrate relationships, Davis ''et al.'' (2012) found acanthodians to be split among the two major clades Osteichthyes (bony fish) and
Chondrichthyes Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
(cartilaginous fish). The well-known acanthodian ''
Acanthodes ''Acanthodes'' (from el, ἄκανθώδης , 'provided with spines') is an extinct genus of spiny shark. Fossils have been found in Europe, North America, Asia, and Australia. ''Acanthodes'' was most common in the Carboniferous and Early Perm ...
'' was placed within Osteichthyes, despite the presence of many chondrichthyan characteristics in its braincase. However, a newly described Silurian placoderm, ''
Entelognathus ''Entelognathus primordialis'' (“primordial complete jaw”) is a maxillate placoderm from the late Silurian (Ludlow epoch) of Qujing, Yunnan, 419 million years ago. A team led by Min Zhu of the Academy of Sciences' Institute of Vertebrate Pa ...
'', which has jaw anatomy shared with
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 ...
and
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, has led to revisions of this phylogeny: acanthodians were then considered to be a paraphyletic assemblage leading to cartilaginous fish, while bony fish evolved from placoderm ancestors. Burrow et al. 2016 provides vindication by finding chondrichthyans to be nested among Acanthodii, most closely related to '' Doliodus'' and '' Tamiobatis''. A 2017 study of ''Doliodus'' morphology points out that it appears to display a mosaic of shark and acanthodian features, making it a
transitional fossil A transitional fossil is any fossilized remains of a life form that exhibits traits common to both an ancestral group and its derived descendant group. This is especially important where the descendant group is sharply differentiated by gross a ...
and further reinforcing this idea.


References


Further reading

* * * *


External links


Acanthodii taxonomy






*

{{Authority control Prehistoric fish classes Teleostomi Permian fish Silurian fish Silurian first appearances Permian extinctions Taxa named by Richard Owen Paraphyletic groups