Holostei Temporal range: | |
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Spotted gar, Lepisosteus oculatus | |
Bowfin, Amia calva | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Actinopterygii |
Subclass: | Neopterygii |
Infraclass: | Holostei Müller, 1846 |
Clades (with orders) | |
Holostei is a group (infraclass) of ray-finned bony fish. It is divided into two major clades, the Halecomorphi, represented by the single living genus, Amia with two species, the bowfins ( Amia calva and Amia ocellicauda ), as well as the Ginglymodi, the sole living representatives being the gars (Lepisosteidae), represented by seven living species in two genera ( Atractosteus , Lepisosteus ). [3] The earliest members of the clade, which are putative "semionotiforms" such as Acentrophorus and Archaeolepidotus , are known from the Middle to Late Permian and are among the earliest known neopterygians. [4] [5] [1] [2]
Holostei was thought to be regarded as paraphyletic. However, a recent study provided evidence that the Holostei are the closest living relatives of the Teleostei, both within the Neopterygii. This was found from the morphology of the Holostei, for example presence of a paired vomer. [6] Holosteans are closer to teleosts than are the chondrosteans, the other group intermediate between teleosts and cartilaginous fish, which are regarded as (at the nearest [a] ) a sister group to the Neopterygii.
The spiracles of holosteans are reduced to vestigial remnants and the bones are lightly ossified. The thick ganoid scales of the gars are more primitive than those of the bowfin.
Holosteans share with other non-teleost ray-finned fish a mixture of characteristics of teleosts and sharks. In comparison with the other group of non-teleost ray-finned fish, the chondrosteans, the holosteans are closer to the teleosts and further from sharks: the pair of spiracles found in sharks and chondrosteans is reduced in holosteans to a remnant structure: in gars, the spiracles do not even open to the outside; [7] the skeleton is lightly ossified: a thin layer of bone covers a mostly cartilaginous skeleton in the bowfins. In gars, the tail is still heterocercal but less so than in the chondrosteans. Bowfins have many-rayed dorsal fins and can breathe air like the bichirs.
In the holosteans a primary pulmonoid (respiratory) swim bladder is still present, a trait that was independently lost in both chondrostei and teleostei, the only other two lineages of fish with a swim bladder (in some teleosts the swim bladder have since evolved to become secondarily respiratory again). [8]
The gars have thick ganoid scales typical of sturgeons whereas the bowfin has thin bony scales like the teleosts. The gars are therefore in this regard considered more primitive than the bowfin. [9]
The name Holostei derives from the Greek words holos, meaning whole, and osteon, meaning bone: a reference to their bony skeletons.
The evolutionary relationships of gars, bowfin and teleosts were a matter of debate. There are two competing hypotheses on the systematics of neopterygians:
The Halecostomi hypothesis proposes Halecomorphi (bowfin and its fossil relatives) as the sister group of Teleostei, the major group of living neopterygians, rendering the Holostei paraphyletic. [10]
The Holostei hypothesis, where the gars and bowfin form the clade Holostei as the sister group to Teleostei, is better supported than the Halecostomi hypothesis, rendering the latter paraphyletic. [11] [12] [13] [14] It proposes Halecomorphi as the sister group of Ginglymodi, the group which includes living gars (Lepisosteiformes) and their fossil relatives. [15] [16] [3] It is estimated that the last common ancestor of gars and bowfin lived at least 250 million years ago. [17]
Neopterygii |
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Ginglymodi comprises three orders: Lepisosteiformes, Semionotiformes and Kyphosichthyiformes. Lepisosteiformes includes 1 family, 2 genera, and 7 species that are commonly referred to as gars. Semionotiformes and Kyphosichthyiformes are extinct orders.
Halecomorphi contains the orders Parasemionotiformes, Panxianichthyiformes, Ionoscopiformes, and Amiiformes. In addition to many extinct species, Amiiformes includes only 1 extant species that is commonly referred to as the bowfin. Parasemionotiformes, Panxianichthyiformes, and Ionoscopiformes have no living members.
Gars and bowfins are found in North America and in freshwater ecosystems. The differences in each can be spotted very easily from just looking at the fishes. The gars have elongated jaws with fanlike teeth, only 3 branchiostegal rays, and a small dorsal fin. Meanwhile the bowfins have a terminal mouth, 10–13 flattened branchiostegal rays, and a long dorsal fin.
The cladogram shows the relationships of holosteans to other living groups of bony fish (Osteichthyes), the great majority of which are teleosts, [19] and to the terrestrial vertebrates (tetrapods) that evolved from a related group of lobe-finned fish. [20] [21] Approximate dates are from Near et al. (2012). [19]
Euteleostomi/ |
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Osteichthyes |
Actinopterygii, members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of skin supported by radially extended thin bony spines called lepidotrichia, as opposed to the bulkier, fleshy lobed fins of the sister class Sarcopterygii. Resembling folding fans, the actinopterygian fins can easily change shape and wetted area, providing superior thrust-to-weight ratios per movement compared to sarcopterygian and chondrichthyian fins. The fin rays attach directly to the proximal or basal skeletal elements, the radials, which represent the articulation between these fins and the internal skeleton.
The bowfin is a ray-finned fish native to North America. Common names include mudfish, mud pike, dogfish, grindle, grinnel, swamp trout, and choupique. It is regarded as a relict, being one of only two surviving species of the Halecomorphi, a group of fish that first appeared during the Early Triassic, around 250 million years ago. The bowfin is often considered a "living fossil" because they have retained some morphological characteristics of their early ancestors. It is one of two species in the genus Amia, along with Amia ocellicauda, the eyespot bowfin. The closest living relatives of bowfins are gars, with the two groups being united in the clade Holostei.
The Amiiformes order of fish has only two extant species, the bowfins: Amia calva and Amia ocellicauda, the latter recognized as a separate species in 2022. These Amiiformes are found in the freshwater systems of North America, in the United States and parts of southern Canada. They live in freshwater streams, rivers, and swamps. The order first appeared in the Triassic, and the extinct members include both marine and freshwater species, many of which are morphologically disparate from bowfins, such as the caturids.
Euteleostomi is a successful clade that includes more than 90% of the living species of vertebrates. Both its major subgroups are successful today: Actinopterygii includes most extant bony fish species, and Sarcopterygii includes the tetrapods.
Actinopteri is the sister group of Cladistia (bichirs) in the class Actinopterygii.
Neopterygii is a subclass of ray-finned fish (Actinopterygii). Neopterygii includes the Holostei and the Teleostei, of which the latter comprise the vast majority of extant fishes, and over half of all living vertebrate species. While living holosteans include only freshwater taxa, teleosts are diverse in both freshwater and marine environments. Many new species of teleosts are scientifically described each year.
Lepisosteiformes is an order of ray-finned fish and the only living members of the clade Ginglymodi. Its only extant representatives are the gar, and it is defined as all members of Ginglymodi that are more closely related to gar than to the extinct Semionotiformes, the other major grouping of ginglymodians. They are one of two extant orders in the infraclass Holostei alongside the Amiiformes.
Pachycormiformes is an extinct order of marine ray-finned fish known from the Early Jurassic to the end of the Cretaceous. It only includes a single family, Pachycormidae. They were characterized by having serrated pectoral fins, reduced pelvic fins and a bony rostrum. Pachycormiformes are morphologically diverse, containing both tuna and swordfish-like carnivorous forms, as well as edentulous suspension-feeding forms.
Halecostomi is the name of a group of neopterygian fish uniting the halecomorphs and the teleosts, the largest group of extant ray-finned fish.
Asialepidotus is an extinct genus of prehistoric marine ray-finned fish that lived during the Ladinian stage of the Middle Triassic epoch. It contains a single species, A. shingyiensis, from Guizhou, China.
Allolepidotus is an extinct genus of prehistoric marine neopterygian ray-finned fish from the Middle Triassic epoch of what is now Italy, Spain, and Switzerland. It was formerly referred to the halecomorph order Panxianichthyiformes, but is now thought to be a member of the Ionoscopiformes.
Kyphosichthys is an extinct genus of basal actinopterygian bony fish known from the lower Middle Triassic (Anisian) marine deposits in Luoping, eastern Yunnan Province, southwestern China. The species is the first known fossil record of highly deep-bodied ginglymodians.
Scheenstia is an extinct genus of neopterygian ray-finned fish from the Late Jurassic–Early Cretaceous of Europe. Fossils have been found in both marine and freshwater environments.
Cipactlichthys is a genus of extinct marine holostean fish from the Lower Cretaceous of Mexico, distantly related to modern bowfins. The only known species is Cipactlichthys scutatus.
Halecomorphi is a taxon of ray-finned bony fish in the clade Neopterygii. The only extant Halecomorph species are the bowfin and eyespot bowfin, but the group contains many extinct species in several families in the order Amiiformes, as well as the extinct orders Ionoscopiformes, Panxianichthyiformes, and Parasemionotiformes. The fossil record of halecomorphs goes back at least to the Early Triassic epoch.
Amia, commonly called bowfin, is a genus of ray-finned fish related to gars in the infraclass Holostei. They are regarded as taxonomic relicts, being the sole surviving species of the order Amiiformes and clade Halecomorphi, which dates from the Triassic to the Eocene, persisting to the present. There are two living species in Amia, Amia calva and Amia ocellicauda, and a number of extinct species which have been described from the fossil record.
Ginglymodi is a clade of ray-finned fish containing modern-day gars (Lepisosteidae) and their extinct relatives in the order Lepisosteiformes, the extinct orders Semionotiformes and Kyphosichthyiformes, and various other extinct taxa. Ginglymodi is one of the two major subgroups of the infraclass Holostei, the other one being Halecomorphi, which contains the bowfin and eyespot bowfin and their fossil relatives.
Parasemionotiformes is an extinct order of neopterygian ray-finned fish that existed globally during the Triassic period. It comprises the families Parasemionotidae and Promecosominidae. Many of the included genera are monotypic and most species lived during the Early Triassic epoch.
Obaichthyidae is an extinct family of ginglymodian ray-finned fish that lived in what is now Africa, South America, and southern Europe during the Cretaceous period. They were close relatives of the modern gars of the family Lepisosteidae, with the two groups making up the superfamily Lepisosteoidea.
Fishes are a paraphyletic group and for this reason, the class Pisces seen in older reference works is no longer used in formal taxonomy. Traditional classification divides fish into three extant classes, and with extinct forms sometimes classified within those groups, sometimes as their own classes:
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