Acanthodii Temporal range: | |
---|---|
Nerepisacanthus , a Silurian acanthodian | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Eugnathostomata |
Class: | † Acanthodii Owen, 1846 |
Orders | |
Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes). They are currently considered to represent a paraphyletic grade of various fish lineages basal to extant Chondrichthyes, which includes living sharks, rays, 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 was covered with tiny rhomboid platelets like the scales of holosteians (gars, bowfins). [1]
The popular name "spiny sharks" is because they were superficially shark-shaped, with a streamlined body, paired fins, 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-sharks". Acanthodians had a cartilaginous skeleton, 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 rocks. The earliest acanthodians were marine, but during the Devonian, freshwater species became predominant.[ citation needed ]
Acanthodians have been divided into four orders: Acanthodiformes, Climatiiformes, Diplacanthiformes, and Ischnacanthiformes. [2] "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 rakers. They were the last and most specialized off the traditional acanthodians, as they survived up until the Permian period.[ citation needed ]
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 thelodonts. The earliest unequivocal acanthodian fossils date from the beginning of the Silurian 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 and in the coal swamps of Carboniferous. By this time bony fishes 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 times (approximately 250 million years ago).
Many palaeontologists originally considered the acanthodians close to the ancestors of the bony fishes.[ citation needed ] Although their interior skeletons were made of cartilage, a bonelike material had developed in the skins of these fishes, in the form of closely fitting scales (see above)[ clarification needed ]. Some scales were greatly enlarged and formed a bony covering on top of the head and over the lower shoulder girdle [ citation needed ]. Others developed a bony flap over the gill openings analogous to the operculum in later bony fishes[ citation needed ]. However, most of these characteristics are considered homologous characteristics derived from common placoderm ancestors[ clarification needed ], and present also in basal cartilaginous fish [ citation needed ]. Overall, the acanthodians' jaws are presumed to have evolved from the first gill arch of some ancestral jawless fishes that had a gill skeleton made of pieces of jointed cartilage.[ citation needed ]
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 (cartilaginous fish). The well-known acanthodian Acanthodes was placed within Osteichthyes, despite the presence of many chondrichthyan characteristics in its braincase. [3] However, a newly described Silurian placoderm, Entelognathus , which has jaw anatomy shared with bony fish and tetrapods, 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. [4]
Burrow et al. 2016 provides vindication by finding chondrichthyans to be nested among Acanthodii, most closely related to Doliodus and Tamiobatis . [2] 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 and further reinforcing this idea. [5]
Phylogeny after [6]
| ||||||||||||||||||||||||||
The oldest remains attributed acanthodian-grade chondrichthyans are Fanjingshania and Qianodus from the Early Silurian of China, dating to around 439 million years ago. [7] [8] Compared to other contemporary groups of fish, acanthodians were relatively morphologically and ecologically conservative. Acanthodians rose in diversity during the Late Silurian, reaching their apex of diversity during the Lochkovian stage of the Early Devonian, declining during the Pragian but rising again during the following Emsian, which was followed by a decline in diversity during middle-Late Devonian. The diversity of the group was consistently low but stable during the Carboniferous, slightly decreasing going into the Permian. [9] The youngest records of the group are isolated scales and fin spines from Middle-Late Permian strata in the Paraná Basin of Brazil. [10]
Chondrichthyes is a class of jawed fish that contains the cartilaginous fish or chondrichthyans, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or bony fish, which have skeletons primarily composed of bone tissue. Chondrichthyes are aquatic vertebrates with paired fins, paired nares, placoid scales, conus arteriosus in the heart, and a lack of opercula and swim bladders. Within the infraphylum Gnathostomata, cartilaginous fishes are distinct from all other jawed vertebrates.
Gnathostomata are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. Most gnathostomes have retained ancestral traits like true teeth, a stomach, and paired appendages. Other traits are elastin, a horizontal semicircular canal of the inner ear, myelin sheaths of neurons, and an adaptive immune system which has discrete lymphoid organs, and uses V(D)J recombination to create antigen recognition sites, rather than using genetic recombination in the variable lymphocyte receptor gene.
Sarcopterygii — sometimes considered synonymous with Crossopterygii — is a clade of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates are characterised by prominent muscular limb buds (lobes) within their fins, which are supported by articulated appendicular skeletons. This is in contrast to the other clade of bony fish, the Actinopterygii, which have only skin-covered bony spines supporting the fins.
Placoderms are vertebrate animals of the class Placodermi, an extinct group of prehistoric fish known from Paleozoic fossils during the Silurian and the Devonian periods. While their endoskeletons are mainly cartilaginous, their head and thorax were covered by articulated armoured plates, and the rest of the body was scaled or naked depending on the species.
Teleostomi is an obsolete taxon of jawed vertebrates that supposedly includes the tetrapods, bony fish, and the wholly extinct acanthodian fish. Key characters of this group include an operculum and a single pair of respiratory openings, features which were lost or modified in some later representatives. The teleostomes include all jawed vertebrates except the chondrichthyans and the extinct class Placodermi.
Acanthodes is an extinct genus of acanthodian fish. Species have been found in Europe, North America, and Asia, spanning the Early Carboniferous to the Early Permian, making it one of the youngest known acanthodian genera.
Psarolepis is a genus of extinct bony fish which lived around 397 to 418 million years ago. Fossils of Psarolepis have been found mainly in South China and described by paleontologist 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 fishes. In 2001, paleontologist John A. Long compared Psarolepis with onychodontiform fishes and refer to their relationships.
Guiyu oneiros is one of the earliest articulated bony fish discovered. Fossils of Guiyu have been found in what is now Qujing, Yunnan, China, in late Silurian marine strata, about 425 million years old.
Most bony fishes have two sets of jaws made mainly of bone. The primary oral jaws open and close the mouth, and a second set of pharyngeal jaws are positioned at the back of the throat. The oral jaws are used to capture and manipulate prey by biting and crushing. The pharyngeal jaws, so-called because they are positioned within the pharynx, are used to further process the food and move it from the mouth to the stomach.
The evolution of fish began about 530 million years ago during the Cambrian explosion. It was during this time that the early chordates developed the skull and the vertebral column, leading to the first craniates and vertebrates. The first fish lineages belong to the Agnatha, or jawless fish. Early examples include Haikouichthys. During the late Cambrian, eel-like jawless fish called the conodonts, and small mostly armoured fish known as ostracoderms, first appeared. Most jawless fish are now extinct; but the extant lampreys may approximate ancient pre-jawed fish. Lampreys belong to the Cyclostomata, which includes the extant hagfish, and this group may have split early on from other agnathans.
Brochoadmones is an extinct genus of acanthodian from the Devonian of what is now Canada. It is the only genus in the suborder Brochoadmonoidei, whose relationship to other acanthodian orders remains currently in flux.
Entelognathus primordialis is an early placoderm from the late Silurian of Qujing, Yunnan, 419 million years ago.
Gogoselachus is an extinct genus of cartilaginous fish known from the late Devonian of Australia. It is one of the earliest well-preserved Devonian chondrichthyans, as much more of the fish than just teeth and scales were preserved. This rare preservation reveals some unique discoveries about the evolution of the cartilage that was inside later cartilaginous fish such as sharks, rays, and chimaeras.
This list of fossil fishes described in 2017 is a list of new taxa of jawless vertebrates, placoderms, acanthodians, fossil cartilaginous fishes, bony fishes and other fishes of every kind that are scheduled to be described during the year 2017, as well as other significant discoveries and events related to paleontology of fishes that are scheduled to occur in the year 2017. The list only includes taxa at the level of genus or species.
Ptomacanthus is an extinct genus of spiny shark, an early relative of living cartilaginous fishes.
Bianchengichthys is a genus of maxillate placoderm fish from the late Silurian Period. Its fossils have been recovered from Yunnan Province, China, and it is represented by only one species: Bianchengichthys micros.
Qianodus is a jawed vertebrate genus that is based on disarticulated teeth from the lower Silurian of China. The type and only species of Qianodus, Q. duplicis, is known from compound dental elements called tooth whorls, each consisting of multiple tooth generations carried by a spiral-shaped base. The tooth whorls of Qianodus represent the oldest unequivocal remains of a toothed vertebrate, predating previously recorded occurrences by about 14 million years. The specimens attributed to the genus come from limestone conglomerate beds of the Rongxi Formation exposed near the village of Leijiatun, Guizhou Province, China. These horizons have been interpreted as tidal deposits1 that form part of the shallow marine sequences of the Rongxi Formation.
Fanjingshania is an extinct genus of "acanthodian" stem-chondrichthyans from the lower Silurian of China around 439 million years old, making it currently the oldest known acanthodian. It comprises a single species, Fanjingshania renovata which is known from over 1,000 isolated elements, including fin spines, branchiostegal plates, sclerotic plates, trunk scales, and tectal and postorbital tesserae. No teeth were ascribed to the taxon, but in the same horizon the tooth taxon Qianodus was found, and these taxa may be synonymous. In a phylogenetic analysis accompanying the description, it was placed in a clade with Climatius, Parexus, and Ptomacanthus.
The evolution of fishes took place over a timeline which spans the Cambrian to the Cenozoic, including during that time in particular the Devonian, which has been dubbed the "age of fishes" for the many changes during that period.