Holostei

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Holostei
Temporal range: Middle Permian to present [1] [2]
Lepisosteus oculatus.jpg
Spotted gar, Lepisosteus oculatus
Amia calva1.jpg
Bowfin, Amia calva
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Subclass: Neopterygii
Infraclass: Holostei
Müller, 1846
Clades (with orders)

Holostei is a group 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]

Contents

Holostei was thought to be regarded as paraphyletic. However, a recent study provided evidence that the Holostei are the closest living relates 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 [lower-alpha 1] ) a sister group to the Neopterigii.

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.

Characteristics

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.

Systematics of Neopterygii

The evolutionary relationships of gars, bowfin and teleosts were a matter of debate. There are two competing hypotheses on the systematics of neopterygians:

Halecostomi hypothesis

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]

Neopterygii
Halecostomi

Teleostei Common carp (white background).jpg

Halecomorphi Amia calva (white background).jpg

Ginglymodi Alligator gar fish (white background).jpg

Holostei hypothesis

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

Teleostei Common carp (white background).jpg

Holostei

Halecomorphi Amia calva (white background).jpg

Ginglymodi Alligator gar fish (white background).jpg

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.

Phylogeny of bony fishes

Cipactlichthys scutatus holotype fossil, from the Lower Cretaceous Tlayua Formation of Mexico Cipactlichthys scutatus holotype.png
Cipactlichthys scutatus holotype fossil, from the Lower Cretaceous Tlayua Formation of Mexico

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/
Sarcopterygii

Actinistia (Coelacanths) Coelacanth flipped.png

Dipnoi (Lungfish) Chinle fish Arganodus cropped cropped.png

Tetrapods

Amphibians Salamandra salamandra (white background).jpg

Amniota

Mammals Ruskea rotta.png

Sauropsids (reptiles, birds) Zoology of Egypt (1898) (Varanus exanthematicus).png

Actinopterygii  400 mya

part of "Chondrostei" [lower-alpha 2] Polypteridae (bichirs) Cuvier-105-Polyptere.jpg

part of "Chondrostei"

Acipenseriformes (sturgeons, paddlefish) Atlantic sturgeon flipped.jpg

Neopterygii  360 mya

Teleostei 310 mya Common carp (white background).jpg

Holostei (bowfins, gars) 275 mya Amia calva (white background).jpg

Osteichthyes

Notes

  1. Depending who you ask, the Chondrostei may be paraphyletic, or the Polypteridae may be considered not part of them.
  2. Thus the former "Chondrostei" is not a clade, but is broken up. See Actinopteri for a possible reclassification.

Related Research Articles

<span class="mw-page-title-main">Actinopterygii</span> Class of ray-finned bony fishes

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.

<span class="mw-page-title-main">Bowfin</span> Bony fish related to gars in the infraclass Holosteii

The bowfin is a bony 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 "primitive fish" 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.

<span class="mw-page-title-main">Amiiformes</span> Order of fishes

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.

<span class="mw-page-title-main">Euteleostomi</span> Clade including most vertebrates

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.

<span class="mw-page-title-main">Actinopteri</span> Group of fishes

Actinopteri is the sister group of Cladistia (bichirs) in the class Actinopterygii.

<span class="mw-page-title-main">Neopterygii</span> Subclass of fishes

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.

<span class="mw-page-title-main">Lepisosteiformes</span> Order of fishes

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.

<span class="mw-page-title-main">Halecostomi</span> Group of ray-finned fishes

Halecostomi is the name of a group of neopterygian fish uniting the halecomorphs and the teleosts, the largest group of extant ray-finned fish.

<i>Asialepidotus</i> Extinct genus of fishes

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.

<i>Ainia <span style="font-style:normal;">(fish)</span></i> Extinct genus of ray-finned fishes

Ainia is an extinct genus of prehistoric ray-finned fish that lived during the Kimmeridgian stage of the Late Jurassic epoch. It contains a single species, A. armata, known from the famous Solnhofen Limestone of Germany. It is a distant relative of the bowfin, although it is more closely related to genera such as Caturus and Osteorachis.

<i>Allolepidotus</i> Extinct genus of fishes

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.

<span class="mw-page-title-main">Euteleostei</span> Clade of ray-finned fishes

Euteleostei, whose members are known as euteleosts, is a clade of bony fishes within Teleostei that evolved some 240 million years ago, although the oldest known fossil remains are only from the Early Cretaceous. It is divided into Protacanthopterygii and Neoteleostei.

<i>Cipactlichthys</i> Extinct genus of ray-finned fishes

Cipactlichthys is a genus of extinct holostean fish from the Lower Cretaceous of Mexico. The only known species is Cipactlichthys scutatus.

<span class="mw-page-title-main">Halecomorphi</span> Clade of fishes

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.

<span class="mw-page-title-main">Percomorpha</span> Clade of ray-finned fishes

Percomorpha is a large clade of ray-finned fish with more than 17 000 known species that includes the tuna, seahorses, gobies, cichlids, flatfish, wrasse, perches, anglerfish, and pufferfish.

<i>Amia</i> (fish) Genus of ray-finned fishes

Amia, commonly called bowfin, is a genus of bony fish related to gars in the infraclass Holostei. They are regarded as taxonomic relicts, being the sole surviving species of the order Amiiformes, which dates from the Jurassic 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.

<span class="mw-page-title-main">Ginglymodi</span> Clade of ray-finned fishes

Ginglymodi is a clade of ray-finned fish containing modern-day gars (Lepisosteidae) & 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.

<span class="mw-page-title-main">Parasemionotiformes</span> Extinct order of ray-finned fishes

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.

<span class="mw-page-title-main">Obaichthyidae</span> Extinct family of ray-finned fishes

Obaichthyidae is an extinct family of ginglymodian ray-finned fish that lived in what is now Africa and South America during the Cretaceous period. It was erected in 2010 by Lance Grande to include the genera Dentilepisosteus and Obaichthys. In 2012, it was defined as a stem-based taxon containing all taxa more closely related to Obaichthys than to the genera Lepisosteus, Pliodetes or Lepidotes.

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:

References

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