Actinopterygii

Ray-finned fish
Temporal range: Late Silurian–Recent
Rose fish
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Superclass*: Osteichthyes
Class: Actinopterygii
Klein, 1885
Subclasses

Actinopterygii /ˌæktᵻnˌɒptəˈrɪdʒi.aɪ/, or the ray-finned fishes, constitute a class or subclass of the bony fishes.

The ray-finned fishes are so called because they possess lepidotrichia or "fin rays", their fins being webs of skin supported by bony or horny spines ("rays"), as opposed to the fleshy, lobed fins that characterize the class Sarcopterygii which also, however, possess lepidotrichia. These actinopterygian fin rays attach directly to the proximal or basal skeletal elements, the radials, which represent the link or connection between these fins and the internal skeleton (e.g., pelvic and pectoral girdles).

Numerically, actinopterygians are the dominant class of vertebrates, comprising nearly 99% of the over 30,000 species of fish.[1] They are ubiquitous throughout freshwater and marine environments from the deep sea to the highest mountain streams. Extant species can range in size from Paedocypris, at 8 mm (0.3 in), to the massive ocean sunfish, at 2,300 kg (5,070 lb), and the long-bodied oarfish, at 11 m (36 ft).

Characteristics

Anatomy of a typical ray-finned fish
A – dorsal fin: B – fin rays: C – lateral line: D – kidney: E – swim bladder: F – Weberian apparatus: G – inner ear: H – brain: I – nostrils: L – eye: M – gills: N – heart O – stomach: P – gall bladder: Q – spleen: R – internal sex organs (ovaries or testes): S – ventral fins: T – spine: U – anal fin: V – tail (caudal fin). Possible other parts not shown: barbels, adipose fin, external genitalia (gonopodium)

Ray-finned fishes occur in many variant forms. The main features of a typical ray-finned fish are shown in the diagram at the left.

Reproduction

Three-spined stickleback males (red belly) build nests and compete to attract females to lay eggs in them. Males then defend and fan the eggs. Painting by Alexander Francis Lydon, 1879

In nearly all ray-finned fish, the sexes are separate, and in most species the females spawn eggs that are fertilized externally, typically with the male inseminating the eggs after they are laid. Development then proceeds with a free-swimming larval stage.[2] However other patterns of ontogeny exist, with one of the commonest being sequential hermaphroditism. In most cases this involves protogyny, fish starting life as females and transitioning to males at some stage, triggered by some internal or external factor. This may be advantageous as females become less prolific as they age while male fecundity increases with age. Protandry, where a fish transitions from male to female, is much less common than protogyny.[3]

Most families use external rather than internal fertilization.[4] Of the oviparous teleosts, most (79%) do not provide parental care.[5] Viviparity, ovoviviparity, or some form of parental care for eggs, whether by the male, the female, or both parents is seen in a significant fraction (21%) of the 422 teleost families; no care is likely the ancestral condition.[5] Viviparity is relatively rare and is found in about 6% of teleost species; male care is far more common than female care.[5][6] Male territoriality "preadapts" a species for evolving male parental care.[7][8]

There are a few examples of fish that self-fertilise. The mangrove rivulus is an amphibious, simultaneous hermaphrodite, producing both eggs and spawn and having internal fertilisation. This mode of reproduction may be related to the fish's habit of spending long periods out of water in the mangrove forests it inhabits. Males are occasionally produced at temperatures below 19 °C (66 °F) and can fertilise eggs that are then spawned by the female. This maintains genetic variability in a species that is otherwise highly inbred.[9]

Fossil record

The earliest known fossil actinopterygiian is Andreolepis hedei, dating back 420 million years (Late Silurian). Remains have been found in Russia, Sweden, and Estonia.[10]

Classification

Actinopterygians are divided into the subclasses Chondrostei and Neopterygii. The Neopterygii, in turn, are divided into the infraclasses Holostei and Teleostei. During the Mesozoic and Cenozoic the teleosts in particular diversified widely, and as a result, 96% of all known fish species are teleosts. The cladogram shows the major groups of actinopterygians and their relationship to the terrestrial vertebrates (tetrapods) that evolved from a related group of fish.[11][12][13] Approximate dates are from Near et al., 2012.[11]

Osteichthyes
Actinopterygii 400 mya

Chondrostei (sturgeons, paddlefishes, reedfishes, bichirs)


Neopterygii 360 mya

Holostei (bowfins, gars) 275 mya



Teleostei 310 mya




Sarcopterygii

Coelacanths, Lungfish



Tetrapods

Amphibians


Amniota

Mammals



Sauropsids (reptiles, birds)







The polypterids (bichirs and ropefish) are the sister lineage of all other actinopterygians, The Acipenseriformes (sturgeons and paddlefishes) are the sister lineage of Neopterygii, and Holostei (bowfin and gars) are the sister lineage of teleosts. The Elopomorpha (eels and tarpons) appears to be the most basic teleosts.[11]

Chondrostei
Atlantic sturgeon
 Chondrostei (cartilage bone) are primarily cartilaginous fish showing some ossification. There are 52 species divided among two orders, the Acipenseriformes (sturgeons and paddlefishes) and the Polypteriformes (reedfishes and bichirs). It is thought that the chondrosteans evolved from bony fish but lost the bony hardening of their cartilaginous skeletons, resulting in a lightening of the frame. Elderly chondrosteans show beginnings of ossification of the skeleton, suggesting that this process is delayed rather than lost in these fish.[14] This group has at times been classified with the sharks: the similarities are obvious, as not only do the chondrosteans mostly lack bone, but the structure of the jaw is more akin to that of sharks than other bony fish, and both lack scales (excluding the Polypteriforms). Additional shared features include spiracles and, in sturgeons, a heterocercal tail (the vertebrae extend into the larger lobe of the caudal fin). However the fossil record suggests that these fish have more in common with the Teleostei than their external appearance might suggest.[14] Chondrostei is paraphyletic meaning that this subclass does not contain all the descendants of their common ancestor; reclassification of the Chondrostei is therefore not out of the question.
Neopterygii
Atlantic salmon
 Neopterygii (new fins) appeared somewhere in the Late Permian, before the time of the dinosaurs. There are only few changes during their evolution from the earlier actinopterygians. They are a very successful group of fishes, because they can move more rapidly than their ancestors. Their scales and skeletons began to lighten during their evolution, and their jaws became more powerful and efficient. While electroreception and the ampullae of Lorenzini is present in all other groups of fish, with the exception of hagfish, Neopterygii has lost this sense, though it later re-evolved within Gymnotiformes and catfishes, who possess nonhomologous teleost ampullae.[15]
Skeleton of the angler fish, Lophius piscatorius. The first spine of the dorsal fin of the anglerfish is modified so it functions like a fishing rod with a lure
Skeleton of another ray-finned fish, the lingcod
Hypsospondylus fossil

The listing below follows Phylogenetic Classification of Bony Fishes [16] with notes when this differs from Nelson,[17] ITIS [18] and FishBase.[19]

Notes

  1. ↑ (Davis, Brian 2010).
  2. ↑ Dorit, R.L.; Walker, W.F.; Barnes, R.D. (1991). Zoology. Saunders College Publishing. p. 819. ISBN 978-0-03-030504-7.
  3. ↑ Avise, J.C.; Mank, J.E. (2009). "Evolutionary perspectives on hermaphroditism in fishes". Sexual Development 3: 152–163. doi:10.1159/000223079.
  4. ↑ Pitcher, T (1993). The Behavior of Teleost Fishes. London: Chapman & Hall.
  5. 1 2 3 Reynolds, John; Nicholas B. Goodwin; Robert P. Freckleton (19 March 2002). "Evolutionary Transitions in Parental Care and Live Bearing in Vertebrates". Philosophical Transactions of the Royal Society B: Biological Sciences 357 (1419). doi:10.1098/rstb.2001.0930. PMC 1692951. PMID 11958696.
  6. ↑ Clutton-Brock, T. H. (1991). The Evolution of Parental Care. Princeton, NJ: Princeton UP.
  7. ↑ Werren, John; Mart R. Gross; Richard Shine (1980). "Paternity and the evolution of male parentage". Journal of Theoretical Biology 82 (4). doi:10.1016/0022-5193(80)90182-4. Retrieved 15 September 2013.
  8. ↑ Baylis, Jeffrey (1981). "The Evolution of Parental Care in Fishes, with reference to Darwin's rule of male sexual selection". Environmental Biology of Fishes 6 (2). doi:10.1007/BF00002788. Retrieved 16 September 2013.
  9. ↑ Wootton, Robert J.; Smith, Carl (2014). Reproductive Biology of Teleost Fishes. Wiley. ISBN 978-1-118-89139-1.
  10. ↑ "Fossilworks: Andreolepis".
  11. 1 2 3 Thomas J. Near; et al. (2012). "Resolution of ray-finned fish phylogeny and timing of diversification". PNAS. pp. 13698–13703. doi:10.1073/pnas.1206625109.
  12. ↑ Betancur-R, Ricardo; et al. (2013). "The Tree of Life and a New Classification of Bony Fishes". PLOS Currents Tree of Life (Edition 1). doi:10.1371/currents.tol.53ba26640df0ccaee75bb165c8c26288.
  13. ↑ Laurin, M.; Reisz, R.R. (1995). "A reevaluation of early amniote phylogeny". Zoological Journal of the Linnean Society 113: 165–223.
  14. 1 2 "Chondrosteans: Sturgeon Relatives". paleos.com.
  15. ↑ Theodore Holmes Bullock; Carl D. Hopkins; Arthur N. Popper (2005). Electroreception. Springer Science+Business Media, Incorporated. p. 229. ISBN 978-0-387-28275-6.
  16. ↑ Betancur-R (2014). "Phylogenetic Classification of Bony Fishes --Version 3".
  17. ↑ Nelson, Joseph, S. (2006). Fishes of the World. John Wiley & Sons, Inc. ISBN 0-471-25031-7.
  18. ↑ "Actinopterygii". Integrated Taxonomic Information System. Retrieved 3 April 2006.
  19. ↑ R. Froese and D. Pauly, editors (February 2006). "FishBase".
  20. ↑ In Nelson, Polypteriformes is placed in its own subclass Cladistia.
  21. ↑ In Nelson and ITIS, Syngnathiformes is placed as the suborder Syngnathoidei of the order Gasterosteiformes.
  22. ↑ In ITIS, Gobiesociformes is placed as the suborder Gobiesocoidei of the order Perciformes.

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