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DESCRIPTION OF THE CLASS OSTEICHTHYES

DESCRIPTION OF THE CLASS OSTEICHTHYES (HUXLEY 1880)

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CLASS OSTEICHTHYES LINKS

Hierarchical Classification

Osteichthyes (o-sti-IK-thes) is made of two Greek roots that mean “bony fish” [bony -osteinos (οστέινος); and fish -ichthys (ιχθύς)]. This is a reference to the presence of bone in the skeleton rather than cartilage of most members of this group. The formal name was coined by Huxley (1880).
INTRODUCTION TO THE OSTEICHTHYES

Osteichthyes is the most speciose class of the Vertebrata, and comprises nearly 50% of all known vertebrates. Such diversity defies a simple description; however, the osteichthyes tend to have the following characters: gills covered by an operculum, one or more dorsal fins, usually one anal fin, most have a homocercal tail and a body covered with scales, usually imbricate or overlapping. The primitive condition in the Osteichthyes is the occurrence of a lung and a bony skeleton in the paired fins, producing a lobe-like base from which rays emerge. The class is formed of two unequal clades (presented here as subclasses) defined by the structure of their paired fins: Actinopterygii (the ray-finned fishes) and the Sarcopterygii (the lobe-finned fishes). Both Figure 1-A and 1-B illustrate the paraphyletic nature of the Osteichthyes with the tetrpods emerging from the Sarcopterygii.
FIGURE 1A. A cladogram that illustrates the relative position of the Osteichthyes (Actinopterygii + Sarcopterygii) in the gnathostome fishes.
MAJOR CLADES OF THE OSTEICHTHYES

1. ACTINOPTERYGII

2. CLADISTIA

3. CHONDROSTEI

4. NEOPTERYGII

5. TELEOSTEI

6. EUTELEOSTEI

7. NEOTELEOSTEI

8. ACANTHOMORPHA

9. ACANTHOPTERYGII

10. SARCOPTERYGII
FIGURE 1B. A cladogram of the Osteichthyes using the Acanthodii as an outgroup. We have used Benton (2005) and Nelson (2006) as the basis for its structure. Use this cladogram as the structure to understand the descriptive text below. We have indicated numbers for the various clades that are discussed below. Note that clades 4-9 are nested.
  • ACTINOPTERYGII (Clade 1)
    • Benton (2005) describes three successive radiation events for the actinopteryrgian bony fishes:
      • Basal actinopterygian or ‘chondrostean’ radiation: Carboniferous – Triassic
      • Basal neopterygian or ‘holostean’ radiation: Triassic – Jurassic
      • Teleost radiation: Jurassic – present
  • The Chondrostean Radiation (Clades 2 & 3)
    • Remnants of the chondrostean radiation are represented today by two genera of Cladistia (bichirs) and six genera of Chondrostei (sturgeons and paddlefish). The living cladistans are restricted to lakes and rivers of central Africa and characterized by having lungs, lobe-like paired fins, ganoid scales, a square body (in cross-section), and in-line fins punctuated by a series of spines. Thus, they look like snakes with a spiny dorsal fin. Their pectoral fins are strong and can be used to propel themselves over land (Figure 2).
    • The Chondrostei [a name derived from the Greek word for cartilage -chrondros (χόνδρος)] is so-called because their skeleton is poorly ossified. Though once quite diverse, the Chondrostei now are represented by sturgeons and paddlefish, both mainly freshwater taxa of large rivers and lakes in the northern hemisphere. Sturgeons have scales reduced to a few bony plates in a row down their sides. They have a characteristic long rostrum and subterminal mouth (Figure 3). The gigantic beluga sturgeon (Huso), the source of black caviar, and most other sturgeon species experienced dramatic reductions through the 20th Century due to pollution, dams, and over fishing. Paddlefish have a large paddle-like extension of the rostrum and a large mouth/pharynx with which they draw in water and capture plankton (Figure 4).
  • The Holostean Radiation and Neopterygii (Clade 4)
    • The holostean radiation of the Triassic through the Jurassic produced a great diversity of fishes with nearly symmetrical, but still heterocercal, tails. Those alive today are represented by three genera, 2 gars and 1 bowfin. Both the gars and bowfins live in freshwater and both are powerful predators. The gars typically have bodies that are round in cross-section and have elongate jaws filled with needle-like teeth (Figure 5). Thus, they are adapted to quick bursts of speed and snatching prey, often by swiping the narrow jaws sideways. Amia or the Bowfin (Figure 6), the only extant taxon in the Amiaformes, has a more rounded head which is adapted to suck in water along with the prey when it gapes its jaws. Teeth line both the jaws and palate. In addition, Amia has a distinctive long dorsal fin with which it moves by undulation. The Bowfin is the closest living species to the modern fishes, the teleosts.
  • The Teleost Radiation and Basal Teleosts (Clade 5)
    • The teleost radiation began in the Jurassic and continues today. This group of bony fishes is by far the the most diverse of all the fishes, indeed of all the vertebrates. The teleosts are characterized by having true homocercal tails and a moveable premaxilla. In addition, the development of the swim bladder from a lung seems to be a synapomorphy of this whole clade. According to Lundberg (2006) the extant teleosts are represented by members of two basal groups, the Elopomorpha (eels, tarpons, etc.), and Osteoglossomorpha (the bonytongues) together with a monophyletic line of the other teleosts. The Otocephala (Herrings and Carp) are the sisters to the Euteleostei (most of the bony fishes including Salmon, Pike, and Perch), the crown teleost taxa.
    • The radiation of teleosts seems to be associated with a third whole genome duplication, termed Teleost-Specific Genome Duplication (Kuraku and Meyer 2009). [See the description of whole genome duplication (WGD) on the Vertebrata description page.] Such a duplication event was first suggested when Jaillon et al. (2004) published the genome of the freshwater puffer fish (Tetraodon nigroviridis). The distribution of Hox clusters is consistent with a third WGD and the subsequent increase in genetic diversity seems to have driven the remarkable biological diversity in the teleosts that we still observe today.
    • Osteoglossomorpha (bonytongues) have tongue bones that usually possess teeth. Thus, they can tear prey with their jaws and tongues. Most are long, thin, and deep-bodied like Arapaima (Figure 7), a predatory fish of the Amazon Basin and the largest freshwater fish in the Western Hemisphere. Similar in shape, but much smaller are the Old World knifefishes and elephantfishes which orient and find prey by generating a weak electromagnetic field. Most of the bonytongues also exhibit some form of parental care.
    • The Elopomorpha (eels, tarpons and their relatives) are characterized by having a larval form, the leptocephalus, that is long, ribbon-like, and planktonic. Usually, the larvae grow to be longer than the adult that they metamorphose into. The most notable members of this group are the eels, which, as adults, have a continuous in-line fin from the mid dorsal to the mid ventral. The North American and European freshwater eels (sibling species in the genus Anguilla) migrate to the Sargasso Sea where they spawn. The leptocephali return to the respective estuaries, metamorphose into a juvenal form called an elver. Tarpon (Figure 8), a large popular sport fish, has a similar life cycle which is entirely marine.
    • Herrings and Carp (Otocephala) have a series of small bones that connect the ear and the air bladder, which provides a means to detect sound vibrations and transfer them to the inner ear. Thus, very different fish like herrings (sardines, anchovies, and menhadens) are united with carp (including suckers and goldfish) and the catfishes (Figure 9 and Figure 10, respectively). Herrings are very important members of the open water fish community in the oceans where they feed on plankton. They are the base of several important global fisheries like the anchovy catch off the coast of Chile and Peru. Carps, suckers, and catfishes, though present in marine environments, are very common in freshwater. Some of the catfishes in the new world have developed electric field generating organs independently of the old world knifefishes (convergent evolution).
  • The Euteleostei (Clade 6) and Neoteleostei (Clade 7)
    • The Euteleostei includes 9 superorders according to Nelson (2006; Procanthopterygerii + Neoteleostii). The most basal of the Euteleostei, and sisters to the Neoteleosteii, are the Procanthopterygii (salmons and pike). The salmonids (salmons and trout; Figure 11) are powerful predatory fish that are the basis of important sport and commercial fisheries. Many salmon species are anadromous and spawn in freshwater, but their young move to the marine environment where they mature and return to freshwater after years in the open ocean. Pike are ambush predators with a elongate jaws and a body shape that is reminiscent of the gar in which the dorsal and anal fins serve as extensions of the caudal fins.
    • The most derived of the bony fishes are in the Neoteleostii, a clade defined by having both sets of paired fins (pectoral and pelvic) very close together and joined by a bony connection. Taxa in this group are extraordinarily diverse and vary enormously in form and size. Collectively, the 8 superorders that make up this group are the most speciose in the Osteichthyes.
    • Among the most basal of the Neoteleosteii are the Dragonfishes (Stenopterygii), Jelly-Nose Fishes (Ateleopodomorpha), Lizardfishes (Cyclosquamata), and the Lanternfishes (Scopelomorpha). Fish in these groups retain some of the primitive features of the salmons and pikes. Still, they are quite diverse and derived. The Dragonfishes are deepsea predators with huge mouths that articulate behind the eyes. Often, they are covered with a pattern of photophores, light-generating organs (see Figure 12). Lanternfishes (see Figure 13) are small fish that school in immense numbers. Thus, they serve as a primary food source for many marine predators. Presumably, they evolved a mechanism to avoid sight feeders and still feed near the surface, where most of the food is in the open ocean. During the day, Lanternfish drop to more that 1000m, well below the illuminated zone of the ocean. At night, they rise to within 10m of the surface to feed. Curiously, they are attracted to lamps at night, and thus earned their name.
  • The Acanthomorpha (8)
    • The remaining four superorders of bony fish are united by having true fin spines on the dorsal, anal, and pelvic fins. Thus, they are called the spiny-rayed fish (Acanthomorpha), which includes Oarfish (Lampriomorpha), Beardfish (Polymixiomorpha), the Trout-Perch and Cod (Paracanthopterygii), and the Suborder with more than half of all species in the Osteichthyes, Acanthopterygii.
    • Oarfish (Figure 14) are very distinctive both in form and appearance. Some of the species can be very long, 8-16m long, making them the longest fish on earth. In addition, they have no teeth, no scales, and no anal fin. The dorsal fin is very long, running the length of the animal, and the anterior end of the dorsal fin is elongated into a crest. Usually, the entire dorsal fin is brightly colored, often red. Certainly, some tales of sea monsters began with sailors sighting these elusive and secretive creatures.
    • Cod (Figure 15) are among the most important economic resources in the oceans. In particular, the Atlantic Cod has spawned and maintained whole fisheries. The individual fish is relatively small, only attaining lengths of about 2m and between 3 and 8 kg. They are bottom feeders and will eat about anything that they can catch. Atlantic Cod prefer cold temperatures (3-10C), particularly during spawning season; however, they can withstand sub-zero temperatures. Cod grow slowly and can take up to 7 years to reach sexual maturity, factors that have caused them to decline significantly in the face of overfishing.
  • Acanthopterigyii (9)
    • The most derived superordinal group is the Acanthopterygii. The diversity of its nearly 15,000 species is truly astounding. Three different clades have been recognized: Mugilomorpha, Atherinomorpha, and Percomorpha. Both Mugilomorpha and Atherinomorpha have pelvic fins that are abdominal, and generally posterior to the pectoral fins. Fish in the Percomorpha, however, tend to have the pelvic fins very far forward, usually in front of the pectoral fins.
    • The Mugilomorpha includes only the mullets (Figure 16), planktivorous fish that occur in marine and brackish coastal waters of all tropical and temperate oceans. They have very reduced teeth (absent in some) and feed entirely by straining water for plankton with elongate gill rakers. The mullets appear to be sisters to Atherinomorpha (silversides, dories, top minnows, needlefish, and live bearers). Of the species in the Atherinomorpha, guppies (Figure 17) are the most well known. In general, the live bearers show sexual dimorphism in which the female is much larger than the male. The male has modified the pelvic fins to become a gonopodium, a structure which serves to transmit sperm to the eggs in the female. She retains the eggs (ovovipary) until they hatch. Guppies have been bred for aquarists to have many fantastic color patterns and fin shapes.
    • The Percomorpha are characterized by having the pelvic girdle attached directly to the pectoral girdle, thus the pelvic fins are very far forward on the animal. I have attempted to present the diversity of percomorphs by describing 7 different distinctive taxa that occur within that group. The taxa are: Seahorses, Freshwater Sunfishes, Cichlids, Tunas, Wrasse, Sole, and Mola.
      • Seahorses and pipefish are primarily marine in all tropical and warm temperate waters. They are distinctive in that the body has become elongated and encased in bony rings. In addition, they have lost the tail fin, but produce a tail that has become long, tapered, and often prehensile. Almost all of the other fins are lost as well. They move by undulations of the dorsal fin. The seahorses and pipefish also are distinctive in that once the eggs have been fertilized, the males brood the young, usually in a pouch. Figure 18 is a Sargassum Seahorse. The extensions on the body allow it to remain camouflaged in the Sargassum, some of which can be seen in the background.
      • The Freshwater Sunfishes are among the most common fishes in streams, ponds, and lakes in North America. They are characterized by having a deep, ovoid body that is quite narrow. Usually, sunfish are small, though maximum length is up to 83cm. The long dorsal fin has an anterior spiny and posterior soft portion. The anal fin also has three anterior spines. These fish excavate depressions for nests, which they defend aggressively. Each species has a distinctive color pattern which includes a spot or “ear” on the operculum. The Long-Eared Sunfish (Figure 19) has an especially long extension on the operculum.
      • Cichlids are among the most interesting fishes on earth and are found in almost all tropical, subtropical, and warm temperate freshwater environments. At first glance, they appear similar to sunfish with a somewhat ovoid body that is deep and compressed. Also, they have a long dorsal fin and an anal fin with spines for the anterior rays. From this simple body plan, cichlids have evolved an astounding number of species. Africa alone has more than 900 species of cichlids, most of which have evolved in the last 5 million years (the approximate age of Lake Tanganyika, Lake Malawi, and Lake Victoria, which house most of the African species). Some have evolved to occupy bizarre niches. For example, certain of them become specialized to eat the eyes of other fishes, while other have become specialized to ambush and snatch scales of other fish. Many, including Tilapia and the African Jewel Fish (Figure 20) are more general in their feeding habits. These fish may be substrate brooders or mouth brooders and exhibit fascinating behaviors as well as interesting color patterns, making them among the most popular aquarium fishes.
      • Mackerels and Tunas have evolved a body for speed and efficient motion in the water, some can swim as fast as 60-70 km/h. Those fish that achieve such high speeds have an array of interesting adaptations, including a limited degree of endothermy. They have two dorsal fins that can be retracted into grooves. In addition, they have small finlets behind the dorsal and anal fins, an area called the caudal peduncle, which also has a pair of lateral keels. The Bluefin Tuna (Figure 21) can grow to be nearly 5 m long and travel in schools. The tunas are among the most prized fish and support a multi-billion dollar a year fishery. However, because they are top predators, their muscle can have significant levels of heavy metals and toxic organic compounds through bioaccumulation. Also, the stocks of some species have declined precipitously through overfishing and poaching.
      • Wrasses are the second largest marine family and vary enormously in size and appearance. In general, they are among the few Acanthopterygii that have protractile mouths with teeth angled outward. Most are small, less than 15cm, and bury themselves in the sand at night. Many are cleaner fishes that go over the outsides of larger fish and clean them of parasites. The largest wrasse is the Humphead Maori Wrasse (Figure 22), a common inhabitant of the Great Barrier Reef, and can grow to be more than 2m long.
      • Flatfishes have evolved to lie on the bottom, sometimes partly buried, very much like the rays. However, they evolved from a line of deep-bodied fish that were laterally flattened. Their evolutionary solution was to become asymmetrical. During development, one eye migrates to the up side of the fish (thus, there are dextral and sinistral taxa). Usually, the eyes are raised on small stalks. In this position, the flatfish can remain camouflaged or survey passersby for a meal as it lunges from ambush. Some of the Flounders and Soles can grow to be larger than a meter. The Dover Sole (Figure 23), the common sole of European cuisine, is a dextral species.
      • Molas are entirely marine and restricted to tropical and subtropical portions of the Atlantic, Indian, and Pacific Oceans. Their bodies are almost circular because of the absence of a tail or caudal peduncle (Figure 24). They swim using very long dorsal and anal fins. Molas often “sun” themselves near the surface, thus they are also called the Ocean Sunfish. They feed almost exclusively on jellyfish and can grow to be more than 1 metric ton and about 2m across. A female Mola can produce as many as 300 million eggs at once.
  • SARCOPTERYGII (10)
    • The other major group of bony fishes is the Sarcopterygii or lobe-finned fishes. Rather that having fins of bony rays, they have fins with fleshy lobes from which rays emerge. Of particular interest is the bony architecture of the lobe, which, in the case of the paired fins, has bones that are homologous to those of tetrapod limbs. Nelson (2006) considers the Sarcopterygii to be a separate class, which includes the lungfishes (Figure 25), coelocanths (Figure 26), and all tetrapods. His taxonomy is a strict application of cladistic rules. In fact, a coelocanth or lungfish is more closely related to a rat than it is to a perch. Nevertheless, in this taxonomy, we consider the lobe-finned fishes to be sisters to the tetrapods. The extant lobe-fins are reduced to two species of coelocanth (a deep sea fish in the Indian Ocean with a characteristic diphycercal tail, which has a central lobed tuft), and the lungfishes, which have a few representatives on Australia, South America, and Africa.
    • For some time fossils of Devonian lobe-finned fishes had been found. By the latter part of the Devonian, many of the lobe-fins had been restricted to brackish and freshwater delta areas. Presumably, muscular limb-like paired fins were useful in clambering over debris and moving alone the muddy bottom. There were many candidates that seemed to be promising as intermediates (they had lungs, typical labyrinthine teeth, a jointed skull, and the bony architecture of the paired appendages). However, the earliest tetrapods of the upper Devonian (see Stegocephali) were dorsoventrally flattened, had eyes on the tops of their heads and had toes (with wrists and ankles). Daeschler et al. (2006) reported finding the remains of a Devonian lobefin the spanned the gap in time and structure. They called the animal Tiktaalik (see Figure 27), which had been discovered on a Canadian polar island in 2004. Tiktaalik had a number of tetrapod and sarcopterygian features. Its pectoral girdle was not attached to its skull so that it had a functional neck. Its body was flattened with eyes on the top of its skull. It had the typical upper limb bones, including a functional wrist, but it also had many fish-like fin rays rather than ‘fingers’ and gills, scales, together with a swimming tail. With Tiktaalik, the boundary between tetrapods and fish became much less distinct.

FIGURE 2. Note the square body, and peg-like nostrils, and lobed fins of Polypterus. This living fossil also has lungs and characters that show it to be near the base of the separation between the Sarcopterygian and Acttinopterygian lines.
Image from: http://web.mit.edu/newsoffice/2008/fish-armor-0727.html

FIGURE 3. A Volga Sturgeon (Acipenser). Note the bony scales, heterocercal tail, and whisker-like barbels around the mouth.
Image from: The Systematics Biodiversity Collection

FIGURE 4. Paddlefish, a large freshwater planktivorous fish that occurs in large rivers east of the Rocky Mountains.
Image from: The Systematics Biodiversity Collection

FIGURE 5. Long-Nosed Gar, Lepidosteus, is a predator with narrow jaws filled with needle-like interlocking teeth. The body is covered with ganoid scales.
Image from: The Systematics Biodiversity Collection

FIGURE 6. Amia, the Bowfin. Note the characteristic long, continuous dorsal fin and the spot at the base of the slightly heterocercal tail. The animal undulates its dorsal fin for most of its locomotion.
Image from: http://www.cnr.vt.edu/efish/families/amiidae.html

FIGURE 7. Arapaima, one of the bone-tongued fishes, is largest freshwater fish in the Western Hemisphere, and is native to the Amazon basin.
Image from: The Systematics Biodiversity Collection

FIGURE 8. Tarpon schooling. Note the characteristic long ray of the dorsal fin on the lower tarpon.
Image from: The Systematics Biodiversity Collection

FIGURE 9. The common carp with its large scales, barbels, and distinctive eversible mouth.
Image from: http://nematode.unl.edu/carp.htm

FIGURE 10. A large Blue Catfish. The body has no scales and some of the fins have a spine-like first ray. Note the barbles around the mouth.
Image from: The Systematics Biodiversity Collection

FIGURE 11. Rainbow Trout. Note the adipose second dorsal fin.
Image from: The Systematics Biodiversity Collection

FIGURE 12. The Dragonfish lives in the deep ocean and is covered with photophores, bioluminescent areas on the skin.
Image from: http://coris.noaa.gov/glossary/photophore_186.jpg

FIGURE 13. Lanternfish congregate in immense numbers around artificial lights at night. They undergo a diurnal vertical migration and occur at more than 1000m during the day and up to 10m at night.
Image from: http://oceanexplorer.noaa.gov/explorations/islands01/log/sep20/media/lanternfish_600.jpg

FIGURE 14. The front end of an Oarfish. Note the red crest on its head and the bright red dorsal fin. This species can grow to be the longest fish (up to 16 meters long).
Image from: http://www.nmfs.noaa.gov/speciesid/fish_page/images/fish82.jpg

FIGURE 15. Atlantic Cod is very important to the economies of many nations, including the US. It is a large fish (almost 2m) and has barbels, three dorsal fins and two anal fins.
Image from: http://bioweb.uwlax.edu/bio203/s2008/bortz_nich/Cod4.jpg

FIGURE 16. A Striped Mullet, a common planktivore that travels in large schools in tropical seas.
Image from: http://www.dnr.sc.gov/marine/mrri/acechar/specgal/image/photos/mulletst.jpg

FIGURE 17. Guppies are small freshwater fish native to Central America. They are sexually dimorphic (the male is below with a gonopodium). The females retains the eggs, which are fertilized inside her by the male via the gonopodium. Thus, they are known as live bearers.
Image from: http://nas.er.usgs.gov/XIMAGESERVERX/2005/20051107160911.jpg

FIGURE 18. A Sargasso Seahorse with bizarre extensions of the body that emulate the thalli of Sargassum in which they live. Note the tail modified into a prehensile appendage.
Image from: The Systematics Biodiversity Collection

FIGURE 19. Long-ear Sunfish is native to the Eastern US where it can be found in ponds, lakes, and slow-moving streams among aquatic vegetation.
Image from: The Systematics Biodiversity Collection

FIGURE 20. African Jewel Fish, an African Cichlid that has become an exotic invasive in the Everglades.
Image from: http://fisc.er.usgs.gov/everglades_invaders/hemichromis_1a.jpg

FIGURE 21. A school of Blue-fin Tuna, one of the most economically-important fish in the world.
Image from: http://www.noaanews.noaa.gov

FIGURE 22. The Humphead Maori Wrasse is the largest member of a very large and variable family. This fish was photographed on the Great Barrier Reef.
Image from: The Systematics Biodiversity Collection

FIGURE 23. A Dover Sole, of the flat fishes. Note that metamorphosis has caused both eyes to be on the right side of the animal.
Image from: http://www.nmfs.noaa.gov/fishwatch/species/dover_sole.htm

FIGURE 24. The Mola, also called the Ocean Sun Fish, can grow to more than 1 metric ton.
Image from: http://www.mms.gov/swss/Assets/Photos/400FullSize/400MolaMola.jpg

FIGURE 25. A lungfish gulping air. Note the almost thread-like fins.
Image from: http://mit.biology.au.dk/zoophysiology/education/cources/evolution/evol_of_airbreathing_lungfish2.jpg

FIGURE 26. Latimeria, a lobe-finned fish from the Indian Ocean.
Image from: http://www.uni-heidelberg.de/presse/news/latimeria_vv.jpg

FIGURE 27. Tiktaalik, a model of the animal as it was alive and a model of the skeleton. Note the dorsoventrally flattened skull, heavily lobed fins with a wrist, and eyes on the top of the head (see Acanthostega and the early tetrapods.
Image from: http://www-news.uchicago.edu/releases/06/images/060406.tiktaalik-3.jpg

LITERATURE CITED

Barton, M. 2007. Bond’s Biology of Fishes. 3rd ed. Thompson Brooks/Cole. Belmont, CA.

Benton, M. J. 2005. Vertebrate Paleontology. Third Edition. Blackwell Publishing, Malden, MA.

Daeschler, E. B., N. H. Shubin, and F. A. Jenkins. 2006. A Devonian tetrapod-like fish and the evolution of the tetrapod body plan. Nature. 440: 757-763.

Helfman. G. S., B. B. Collette, and D. E. Facey. 1997. The Diversity of Fishes. Blackwell Science, Inc. Malden, Mass.

Huxley, T. H. 1880. On the application of the laws of evolution to the arrangement of the Vertebrata and more particularly of the Mammalia. Proceedings of the Zoological Society of London. 1880: 649–662.
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Kuraku, S. and A. Meyer. 2009. The evolution and maintence of Hox gene clusters in vertebrates and the teleost-specific genome duplication. International Journal of Developmental Biology. 53: 765-773.

Moyle, P. B. and J. J. Cech. 2003. Fishes: An Introduction to Ichthyology. 5th ed. Benjamin Cummings. San Francisco, CA.

Nelson, J. S. 2006. Fishes of the World. 4th edition. John Wiley and Sons, Inc. New York.

Paxton, J. R. and W. N. Eschmeyer, eds. 1998. Encyclopedia of Fishes. Academic Press. San Diego, CA.

Pough, F. H., C. M. Janis, and J. B. Heiser. 2009. Vertebrate Life. 8th ed. Benjamin Cummings. New York. pp. 688.
By Jack R. Holt and Carlos A. Iudica. Last revised: 01/20/2014
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