Holostei is a group of

ray-finned 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 ...

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 ...

. It is divided into two major clades, the

Halecomorphi Halecomorphi is a taxon of ray-finned bony fish in the clade Neopterygii. The sole living Halecomorph is the bowfin (''Amia calva''), but the group contains many extinct species in several families (including Amiidae, Caturidae, Liodesmidae, S ...

, represented by a single living species, the bowfin (''

Amia calva The bowfin (''Amia calva'') 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 the sole surviving species of the Halecomorphi ...

''), as well as the

Ginglymodi Ginglymodi is a clade of ray-finned fish containing modern-day gars (Lepisosteidae) and their extinct relatives, including the family Lepidotidae and the orders Semionotiformes and Kyphosichthyiformes, and various other extinct taxa. Ginglymod ...

, the sole living representatives being the

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 (Lepisosteidae), represented by seven living species in two genera (''

Atractosteus ''Atractosteus'' is a genus of gars in the family Lepisosteidae, with three species. The genus first appeared in the Campanian in the Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous ...

'', ''

Lepisosteus ''Lepisosteus'' is a genus of gars in the family Lepisosteidae. Distribution While in the present day, ''Lepisosteus'' is only known across North America, fossil remains show it was much more widespread in the past, with specimens known from t ...

''). The earliest members of the clade appeared during the

Early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which is a un ...

, over 250 million years ago. Holostei was thought to be regarded as

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 ...

. However, a recent study provided evidence that the Holostei are the closest living relates of the

Teleostei Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest class (biology), infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of a ...

, both within the

Neopterygii Neopterygii (from Greek νέος ''neos'' 'new' and πτέρυξ ''pteryx'' 'fin') 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 f ...

. This was found from the morphology of the Holostei, for example presence of a paired

vomer The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxill ...

. Holosteans are closer to teleosts than are the

chondrosteans Chondrostei is a group of non-neopterygian ray-finned fish, while the term originally referred to a paraphyletic group of all non-neopterygian ray-finned fish, it was redefined by Patterson in 1982 to be a clade comprising the Acipenseriformes (w ...

, the other group intermediate between teleosts and cartilaginous fish, which are regarded as (at the nearest) a sister group to the Neopterigii. The

spiracle Spiracle or spiraculum may refer to: * Spiracle (arthropods), opening in the exoskeletons of some arthropods * Spiracle (vertebrates), openings on the surface of some vertebrates * Spiraculum, a genus of land snails in family Cyclophoridae Cycl ...

s of holosteans are reduced to vestigial remnants and the bones are lightly ossified. The thick

ganoid A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as w ...

scales of the gars are more primitive than those of the bowfin.

Characteristics

Holosteans share with other non-teleost

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 ...

a mixture of characteristics of

teleosts Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Teleo ...

and

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. In comparison with the other group of non-teleost ray-finned fish, the

, 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; the skeleton is lightly

ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by Cell (biology), cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes ...

: a thin layer of bone covers a mostly

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 in the bowfins. In gars, the tail is still

heterocercal Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as s ...

but less so than in the chondrosteans. Bowfins have many-rayed dorsal fins and can breathe air like the

bichir Bichirs and the reedfish comprise Polypteridae , a family of archaic ray-finned fishes and the only family in the order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The Diversity of Fishes. West Sussex, UK: Blackwell Pub ...

s. The gars have thick

ganoid scale A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as ...

s typical of

sturgeon Sturgeon is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretace ...

s whereas the bowfin has thin bony scales like the teleosts. The gars are therefore in this regard considered more primitive than the bowfin. The name ''Holostei'' derives from the

Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...

words ''holos'', meaning whole, and ''osteon'', meaning bone: a reference to their bony skeletons.

Systematics of Neopterygii

The

evolutionary relationships Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...

of gars, bowfin and teleosts were a matter of debate. There are two competing

hypotheses A hypothesis (plural hypotheses) is a proposed explanation for a phenomenon. For a hypothesis to be a scientific hypothesis, the scientific method requires that one can test it. Scientists generally base scientific hypotheses on previous obser ...

on the systematics of neopterygians:

Halecostomi hypothesis

The Halecostomi hypothesis proposes

(

bowfin The bowfin (''Amia calva'') 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 the sole surviving species of the Halecomorphi ...

and its fossil relatives) as the

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...

, the major group of living neopterygians, rendering the Holostei

Holostei hypothesis

The Holostei hypothesis is better supported than the Halecostomi hypothesis, rendering the latter paraphyletic. It proposes Halecomorphi as the sister group of

, the group which includes living

s ( Lepisosteiformes) and their fossil relatives. Ginglymodi and Halecomorphi form the

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

Holostei, which is the

to Teleostei. Ginglymodi comprises three

orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

: Lepisosteiformes,

Semionotiformes Semionotiformes is an order of primitive, ray-finned, primarily freshwater fish from the Middle Triassic (Anisian) to the Late Cretaceous (Maastrichtian). The best-known genus is ''Semionotus'' of Europe and North America. Their closest living re ...

and Kyphosichthyiformes. Lepisosteiformes includes 1

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

, 2

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

, and 7

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 ...

that are commonly referred to as gars. Semionotiformes and Kyphosichthyiformes are extinct orders. Halecomorphi contains the orders

Parasemionotiformes 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 speci ...

, Panxianichthyiformes, Ionoscopiformes, and

Amiiformes The Amiiformes order of fish has only one extant species, the bowfin (''Amia calva''). 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, r ...

. In addition to many

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

, Amiiformes includes only 1

extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...

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 Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does include ...

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

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 ...

shows the relationships of holosteans to other living groups of

, the great majority of which are

teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tel ...

s, and to the terrestrial vertebrates (tetrapods) that evolved from a related group of

lobe-finned fish Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includ ...

. Approximate dates are from Near et al. (2012).

Notes

References

External links

Holostei on The University of Liverpool website
{{Taxonbar, from=Q846164 Neopterygii Extant Jurassic first appearances