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Teleostei - AccessScience from McGraw-Hill Education http://www.accessscience.com/content/teleostei/680400

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Article by: Boschung, Herbert Department of Biological Sciences, University of Alabama, Tuscaloosa, Alabama. Gardiner, Brian Linnean Society of London, Burlington House, Piccadilly, London, United Kingdom. Publication year: 2014 DOI: http://dx.doi.org/10.1036/1097-8542.680400 (http://dx.doi.org/10.1036/1097-8542.680400)

Content

Morphology Bibliography Phylogeny Classification Additional Readings Clupeomorpha and

The most recent group of actinopterygians (rayfin ), first appearing in the Upper (Fig. 1). About 26,840 are contained within the Teleostei, accounting for more than half of all living and over 96% of all living fishes. comprise 517 families, of which 69 are extinct, leaving 448 extant families; of these, about 43% have no record. See also: (/content/actinopterygii/009100); (/content/osteichthyes/478500)

Fig. 1 Cladogram showing the relationships of the extant teleosts with the other extant actinopterygians. (J. S. Nelson, , 4th ed., Wiley, New York, 2006)

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Much of the evidence for (evolving from a common ancestral form) and relationships comes from the caudal and concomitant acquisition of a homocercal tail (upper and lower lobes of the caudal fin are symmetrical). This type of tail primitively results from an ontogenetic fusion of centra (bodies of vertebrae) and the possession of paired bracing bones located bilaterally along the dorsal region of the caudal skeleton, derived ontogenetically from the neural arches (uroneurals) of the ural (tail) centra. The presence of uroneurals is a synapomorphy () for all teleosts and is recognizable in the fossil groups, as is another synapomorphy—the articulation of the first two hypurals with the same vertebral centrum (ural centrum 1, which results from an ontogenetic fusion of two centra). Other characteristics of the teleosts include a mobile premaxillary bone, unpaired basibranchial plates, extension of the posterior myodome (the eye muscle canal) into the basioccipital bone, and development of the .

The swim bladder (gas bladder, air bladder), lying between the gut and kidneys, develops as a diverticulum from the dorsal wall of the esophagus. In the groups of teleosts, the swim bladder usually remains in contact with the esophagus via the pneumatic duct, a condition called physostomous. In more advanced groups, this connection may be lost (physoclistic bladder). Accompanying the development of a physoclistic condition are retia (anatomical networks) of blood vessels that form gas glands that pump gases into or out of the bladder. Thus, the principal function of the physoclistic swim bladder is hydrostatic. See also: Swim bladder (/content/swim-bladder/672500)

Ostariophysans have a chain of ossicles (little bones) and associated structures connecting the swim bladder with the , the so-called . In some osteoglossomorphs, clupeomorphs, and euteleosts, there is a direct connection between the diverticulum of the swim bladder and the labyrinth of the ear, with the diverticulum passing through an opening in the exoccipital bone and terminating in two distinct vesicles (one in the prootic, the other in the pterotic). Thus, in both ostariophysans and clupeomorphs, stimuli are transmitted from the swim bladder to the utriculus. However, the nature of the two otophysic (ear-swim bladder) connections is not regarded as homologous. The Weberian ossicles allow the swim bladder to function as a manometer or pressure receptor. In the Clupeomorpha, there is also a connection between the swim bladder and the , and it has been shown that these swim bladder-ear connections in the are important in shoaling (schooling) behavior. Also, the swim bladder can function as a resonator when beaten by the ribs, as in certain toadfishes. See also: Ear () (/content/ear-vertebrate/208600); Ostariophysi (/content/ostariophysi/802680)

Phylogeny

In the past four or five decades, a surge of interest in teleost fishes resulted in several ideas regarding the details of their origins and lines of descent (phylogenies), any one of which may be closer to the truth than the others. Notwithstanding this high level of interest, much work remains ahead for current and future generations of researchers to unravel the evolutionary history of this tremendously large group of fishes.

Teleosts probably had their beginning in the middle to late Triassic, about 220 to 200 million years ago. By the end of the , several representatives of Recent teleost taxa had evolved; these included the , , and Clupeomorpha, but the remained the dominant fishes. It was in the succeeding Period that great radiation of the teleost group occurred and at that time they exceeded the Holostei to become by far the largest and most diverse group of vertebrates. The beds at Monte Bolca in Verona, Italy, provide the earliest records (lowermost ) for a large number of higher teleost taxa (Fig. 2). See also: (/content/clupeiformes/142500); (/content/elopiformes/229100); Geologic time (/content/geologic-time-scale/286500); Holostei (/content/holostei /321000); (/content/osteoglossiformes/478600)

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Fig. 2 Phylogeny of the Teleostei.

Teleostei is a of (bow fins). Osteoglossomorpha, Elopomorpha, and Clupeomorpha are now generally regarded as successive (groups) above the level of the fossil, paraphyletic pholidophorids. Clupeomorpha is identified as the sister group of Ostariophysi. See also: Amiiformes (/content/amiiformes/027800)

Clupeomorpha and Ostariophysi

Clupeomorpha and Ostariophysi are collectively named or Ostarioclupeomorpha. Clupeomorpha has one extant (Clupeiformes), with 364 species in 84 genera and 5 families. Ostariophysi accounts for about 68% of the freshwater fishes of the world. Clupeomorpha plus Ostariophysi, comprising 6 orders, are sister to the Euteleostei.

Euteleostei

By far, the largest teleost is Euteleostei, with about 17,419 species arranged in some 346 families. It comprises two major lineages, and (see table).

Table - Two proposed classifications of the Euteleostei

Johnson and Patterson Nelson

Protacanthopterygii Protacanthopterygii

Argentiniformes

Salmoniformes

Salmonoidei Salmoniformes

Osmeroidei

Neognathi Neoteleostei—the remaining teleosts

Esociformes

Neoteleostei

Protacanthopterygii

According to J. S. Nelson, Salmoniformes and Esociformes are sister taxa. Protacanthopterygii comprises four orders and is sister to the Neoteleostei. G. D. Johnson and C. Patterson recognize two orders of protacanthopterygians by combining the

3 of 9 10/7/2015 1:07 PM Teleostei - AccessScience from McGraw-Hill Education http://www.accessscience.com/content/teleostei/680400 osmeroids and salmonids in Salmoniformes, the sister group of the argentinoids. They regard Esociformes as sister to the Neoteleostei.

Neoteleostei

Nelson suggests that support for the monophyly of Neoteleostei is stronger for itself than for its individual member taxa, namely , Ateleopodiformes, , , , Polymixiiformes, , and (see classification below).

The Stomiiformes, a morphologically diverse group of 321 species of deep-water oceanic fishes, seems to be the best candidate as the sister group to all the other neoteleosts. Molecular work of M. Miya and coworkers suggests that Ateleopodiformes is the sister group of Lampriformes and that the two are sister to Myctophiformes. This is unlike the phylogeny shown in Fig. 2. The last four superorders constitute the (spiny-rayed fishes), which left a rich fossil record beginning in the Cretaceous. Scopelomorpha (Myctophiformes) is regarded as the sister group of Acanthomorpha. Lampriomorpha (Lampriformes) is regarded as the primitive sister group to the remaining acanthomorphs, that is, Polymixiomorpha (Polymixiiformes), paracanthopterygians, and acanthopterygians (Fig. 3).

Fig. 3 Cladogram showing the relationships of the Neoteleostei. (J. S. Nelson, Fishes of the World, 4th ed., Wiley, New York, 2006)

Paracanthopterygii is a diverse group of acanthomorphs, which defies a clear-cut definition and lacks firm evidence to support its monophyly. As with the Cyclosquamata (Aulopiformes), it contains various deep-sea groups, including the Lophiiformes (angler fishes) with 297 species and the with about 482 species, many of which are important commercial fishes that constitute over one-quarter of the world's marine fish catch. Other paracanthopterygian groups include the freshwater ( and pirate and cave fishes; 9 species) and the Batrachoidiformes (coastal, benthic toadfishes; 69 species).

Acanthopterygii is the largest subgroup of the Euteleostei, distributed among 13 orders and 267 families (whose interrelationships are mostly unknown), 2422 genera, and 14,797 species. In Fig. 4, the represent orders , , and . The form the largest of these orders with about 10,033 species in 20 suborders, 160 families, and about 1539 genera (many members of which bear spines on their fins). Of the more advanced members, the (mail-cheeked fishes), with 25 families and about 1271 species, are second to the Perciformes in species richness, whereas the Pleuronectiformes () with 570 species and the ( and pufferfish) with 339 species are a distant third and fourth. Members of a lower include

4 of 9 10/7/2015 1:07 PM Teleostei - AccessScience from McGraw-Hill Education http://www.accessscience.com/content/teleostei/680400 the (, sea horses, pipe fishes, and others; 257 species) and the Atherinomorpha (rainbow fishes, silversides, blue eyes; 285 species).

Fig. 4 Cladogram showing the relationships of the Acanthopterygii. (J. S. Nelson, Fishes of the World, 4th ed., Wiley, New York, 2006)

Classification

Below is a classification of the extant teleost fishes as provided by Nelson, which is adopted for use here.

Division Teleostei

Subdivision Osteoglossomorpha (Jurassic and Cretaceous)

Order (http://www.accessscience.com/content/hiodontiformes/803340) (mooneyes)

Order Osteoglossiformes (http://www.accessscience.com/content/osteoglossiformes/478600) (bonytongues)

Subdivision Elopomorpha ()

Order Elopiformes (http://www.accessscience.com/content/elopiformes/229100) (ladyfishes and )

Order Albuliformes (http://www.accessscience.com/content/albuliformes/803190) (bonefishes)

Order Anguilliformes (http://www.accessscience.com/content/anguilliformes/034600) (eels)

Order Saccopharyngiformes (http://www.accessscience.com/content/saccopharyngiformes/802750) (sackpharynx fishes)

Subdivision Otocephala (=Ostarioclupeomorpha)

Superorder Clupeomorpha (Cretaceous)

Order Clupeiformes (http://www.accessscience.com/content/clupeiformes/142500)

5 of 9 10/7/2015 1:07 PM Teleostei - AccessScience from McGraw-Hill Education http://www.accessscience.com/content/teleostei/680400 Superorder Ostariophysi (Early Cretaceous to )

Order (http://www.accessscience.com/content/gonorynchiformes/295500) (milkfishes)

Order (http://www.accessscience.com/content/cypriniformes/177800) (minnows, , and suckers)

Order (http://www.accessscience.com/content/characiformes/803070) (characins)

Order Siluriformes (http://www.accessscience.com/content/siluriformes/623700) (catfishes)

Order (http://www.accessscience.com/content/gymnotiformes/803360) (American knife-fishes)

Subdivision Euteleostei

Superorder Protacanthopterygii

Order Argentiniformes (http://www.accessscience.com/content/argentiniformes/802830) (marine smelts)

Order Osmeriformes (freshwater smelts)

Order Salmoniformes (http://www.accessscience.com/content/salmoniformes/599300) (trouts)

Order Esociformes (http://www.accessscience.com/content/esociformes/803280) (pikes) [, early Eocene]

Order Neoteleostei

Superorder

Order Stomiiformes (http://www.accessscience.com/content/stomiiformes/803490) (dragonfishes)

Superorder Ateleopodomorpha

Order Ateleopodiformes (http://www.accessscience.com/content/ateleopodiformes/802800) (jellynose fishes)

Superorder Cyclosquamata

Order Aulopiformes (http://www.accessscience.com/content/aulopiformes/802820) (lizard fishes)

Superorder Scopelomorpha

Order Myctophiformes (http://www.accessscience.com/content/myctophiformes/803100) (lanternfishes)

Superorder Lampriomorpha

Order Lampriformes (http://www.accessscience.com/content/lampriformes/369300) (opahs)

Superorder Polymixiomorpha

Order Polymixiiformes (http://www.accessscience.com/content/polymixiiformes/803460)

6 of 9 10/7/2015 1:07 PM Teleostei - AccessScience from McGraw-Hill Education http://www.accessscience.com/content/teleostei/680400 Superorder Paracanthopterygii

Order Percopsiformes (http://www.accessscience.com/content/percopsiformes/498000) (trout-perches)

Order Gadiformes (http://www.accessscience.com/content/gadiformes/277100) (cods)

Order (cusk-)

Order Batrachoidiformes (http://www.accessscience.com/content/batrachoidiformes/075100) (toadfishes)

Order Lophiiformes (http://www.accessscience.com/content/lophiiformes/389400) (angler fishes)

Superorder Acanthopterygii

Order Mugiliformes (http://www.accessscience.com/content/mugiliformes/803080) (mullets)

Series Atherinomorpha

Order Atheriniformes (http://www.accessscience.com/content/atheriniformes/058600) (silversides)

Order Beloniformes (http://www.accessscience.com/content/beloniformes/802810) (needlefishes)

Order Cyprinodontiformes (http://www.accessscience.com/content/cyprinodontiformes/802870) ()

Series

Order (http://www.accessscience.com/content/stephanoberyciformes/803500) (pricklefishes)

Order (http://www.accessscience.com/content/beryciformes/079500) (squirrelfishes)

Order (http://www.accessscience.com/content/zeiformes/754300) (dories)

Order Gasterosteiformes (http://www.accessscience.com/content/gasterosteiformes/282000) (sticklebacks)

Order (http://www.accessscience.com/content/synbranchiformes/674200) (swamp eels)

Order Scorpaeniformes (http://www.accessscience.com/content/scorpaeniformes/802720) (mail-cheeked fishes)

Order Perciformes (http://www.accessscience.com/content/perciformes/497900) (perches) [Eocene; ]

Order Pleuronectiformes (http://www.accessscience.com/content/pleuronectiformes/528800) (flatfishes)

Order Tetraodontiformes (http://www.accessscience.com/content/tetraodontiformes/686800) (plectognaths)

Herbert Boschung Brian Gardiner

Bibliography

G. Arratia, Basal teleosts and teleostean phylogeny, Palaeo Ichthyol., 7:1–168, 1997

7 of 9 10/7/2015 1:07 PM Teleostei - AccessScience from McGraw-Hill Education http://www.accessscience.com/content/teleostei/680400 G. Arratia, The monophyly of Teleostei and stem-group teleosts: Consensus and disagreements, pp. 265–334, in G. Arratia and H.-P. Schultze (eds.), Mesozoic Fishes 2— and Fossil Record, Verlag Dr. Friedrich Pfeil, Munich, 1999

G. D. Johnson and C. Patterson, Percomorph phylogeny: A survey of acanthomorphs and a new proposal, Bull. Mar. Sci., 52(1):554–626, 1993

M. Miya et al., Major patterns of higher teleostean phylogenies: A new perspective based on 100 complete mitochondrial DNA sequences, Mol. Phylogenet. Evol., 26(1):121–138, 2003 DOI: 10.1016/S1055-7903(02)00332-9 (http://dx.doi.org/10.1016 /S1055-7903(02)00332-9)

J. S. Nelson, Fishes of the World, 4th ed., Wiley, New York, 2006

J. R. Paxton and W. N. Eschmeyer (eds.), Encyclopaedia of Fishes, 2d ed., Academic Press, San Diego, 1998

M. L. J. Stiassny, L. R. Parenti, and G. D. Johnson (eds.), Interrelationships of Fishes, Academic Press, San Diego, 1996

Additional Readings

B. E. Erisman et al., Phylogenetic perspectives on the of functional hermaphroditism in teleost fishes, Integr. Comp. Biol., 53(4):736–754, 2013 DOI: 10.1093/icb/ict077 (http://dx.doi.org/10.1093/icb/ict077)

G. S. Helfman et al., The Diversity of Fishes: Biology, Evolution, and Ecology, John Wiley & Sons, Chichester, West Sussex, UK, 2009

J. Oriol Sunyer, for mammalian paradigms in the teleost immune system, Nat. Immunol., 14:320–326, 2013 DOI: 10.1038/ni.2549 (http://dx.doi.org/10.1038/ni.2549)

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