Actinopterygii
Actinopterygii, or the ray-finned fishes, constitute a class or subclass of the bony fishes.
The ray-finned fishes are so called because their fins are webs of skin supported by bony or horny spines, as opposed to the fleshy, lobed fins that characterize the class Sarcopterygii. 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.
Numerically, actinopterygians are the dominant class of vertebrates, comprising nearly 99% of the over 30,000 species of fish. 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, to the massive ocean sunfish, at, and the long-bodied oarfish, at.
Characteristics
Ray-finned fishes occur in many variant forms. The main features of a typical ray-finned fish are shown in the adjacent diagram.Ray-finned fishes have many different types of scales; but all teleosts, the most advanced Actinopterygii, have leptoid scales. The outer part of these scales fan out with bony ridges while the inner part is crossed with fibrous connective tissue. Leptoid scales are thinner and more transparent than other types of scales, and lack the hardened enamel or dentine-like layers found in the scales of many other fish. Unlike ganoid scales, which are found in non-teleost Actinopterygii, new scales are added in concentric layers as the fish grows.
Fin arrangements
Ray-finned fish are very varied in size and shape, and in the number of their ray-fins and the manner in which they arrange them.Reproduction
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. 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 converting to males at some stage, triggered by some internal or external factor. Protandry, where a fish converts from male to female, is much less common than protogyny.Most families use external rather than internal fertilization. Of the oviparous teleosts, most do not provide parental care. 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 of the 422 teleost families; no care is likely the ancestral condition. Viviparity is relatively rare and is found in about 6% of teleost species; male care is far more common than female care. Male territoriality "preadapts" a species for evolving male parental care.
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 and can fertilise eggs that are then spawned by the female. This maintains genetic variability in a species that is otherwise highly inbred.
Fossil record
The earliest known fossil actinopterygian is Andreolepis hedei, dating back 420 million years. Remains have been found in Russia, Sweden, and Estonia.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 that evolved from a related group of fish. Approximate dates are from Near et al., 2012.The polypterids are the sister lineage of all other actinopterygians, the Acipenseriformes are the sister lineage of Neopterygii, and Holostei are the sister lineage of teleosts. The Elopomorpha appear to be the most basal teleosts.
Chondrostei | Chondrostei ' are primarily cartilaginous fish showing some ossification. Earlier definition of Chondrostei is now known to be paraphyletic meaning that this subclass does not contain all the descendants of their common ancestor. There were 52 species divided among two orders, the Acipenseriformes and the Polypteriformes. Reedfish and birchirs are now separated from the Chondrostei into their own sister lineage, the Cladistia. 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. This group had once 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. Additional shared features include spiracles and, in sturgeons, a heterocercal tail. However the fossil record suggests that these fish have more in common with the Teleostei than their external appearance might suggest. | |
Neopterygii | Neopterygii ' 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. |
. The first spine of the dorsal fin of the anglerfish is modified so it functions like a fishing rod with a lure
about 50 million years ago
The listing below follows Phylogenetic Classification of Bony Fishes with notes when this differs from Nelson, ITIS and FishBase and extinct groups from Van der Laan 2016.
- Order †?Asarotiformes Schaeffer 1968
- Order †?Discordichthyiformes Minikh 1998
- Order †?Paphosisciformes Grogan & Lund 2015
- Order †?Scanilepiformes Selezneya 1985
- Order †Cheirolepidiformes Kazantseva-Selezneva 1977
- Order †Paramblypteriformes Heyler 1969
- Order †Rhadinichthyiformes
- Order †Palaeonisciformes Hay 1902
- Order †Tarrasiiformes sensu Lund & Poplin 2002
- Order †Ptycholepiformes Andrews et al. 1967
- Order †Redfieldiiformes Berg 1940
- Order †Haplolepidiformes Westoll 1944
- Order †Aeduelliformes Heyler 1969
- Order †Platysomiformes Aldinger 1937
- Order †Dorypteriformes Cope 1871
- Order †Eurynotiformes Sallan & Coates 2013
- Subclass Cladistii Pander 1860
- * Order †Guildayichthyiformes Lund 2000
- * Order Polypteriformes Bleeker 1859
- Clade Actinopteri Cope 1972 s.s.
- * Order †Elonichthyiformes Kazantseva-Selezneva 1977
- * Order †Phanerorhynchiformes
- * Order †Saurichthyiformes Berg 1937
- * Subclass Chondrostei
- ** Order †Birgeriiformes Jin 2001
- ** Order †Chondrosteiformes
- ** Order Acipenseriformes Berg 1940
- * Subclass Neopterygii Regan 1923 sensu Xu & Wu 2012
- ** Order †Pholidopleuriformes Berg 1937
- ** Order †Peltopleuriformes Lehman 1966
- ** Order †Perleidiformes Berg 1937
- ** Order †Luganoiiformes Lehman 1958
- ** Order †Pycnodontiformes Berg 1937
- ** Infraclass Holostei Muller 1844
- *** Division Halecomorpha Cope 1872 sensu Grande & Bemis 1998
- **** Order †Parasemionotiformes Lehman 1966
- **** Order †Ionoscopiformes Grande & Bemis 1998
- **** Order Amiiformes Huxley 1861 sensu Grande & Bemis 1998
- *** Division Ginglymodi Cope 1871
- **** Order †Dapediiformes Thies & Waschkewitz 2015
- **** Order †Semionotiformes Arambourg & Bertin 1958
- **** Order Lepisosteiformes Hay 1929
- ** Clade Teleosteomorpha Arratia 2000 sensu Arratia 2013
- *** Order †Prohaleciteiformes Arratia 2017
- *** Division Aspidorhynchei Nelson, Grand & Wilson 2016
- **** Order †Aspidorhynchiformes Bleeker 1859
- **** Order †Pachycormiformes Berg 1937
- *** Infraclass Teleostei Müller 1844 sensu Arratia 2013
- **** Order †?Araripichthyiformes
- **** Order †?Ligulelliiformes Taverne 2011
- **** Order †?Tselfatiiformes Nelson 1994
- **** Order †Pholidophoriformes Berg 1940
- **** Order †Dorsetichthyiformes Nelson, Grand & Wilson 2016
- **** Order †Leptolepidiformes
- **** Order †Crossognathiformes Taverne 1989
- **** Order †Ichthyodectiformes Bardeck & Sprinkle 1969
- **** Teleocephala de Pinna 1996 s.s.
- *****Megacohort Elopocephalai Patterson 1977 sensu Arratia 1999
- ****** Order Elopiformes Gosline 1960
- ****** Order Albuliformes Greenwood et al. 1966 sensu Forey et al. 1996
- ****** Order Notacanthiformes Goodrich 1909
- ****** Order Anguilliformes Jarocki 1822 sensu Goodrich 1909
- ***** Megacohort Osteoglossocephalai sensu Arratia 1999
- ******Supercohort Osteoglossocephala sensu Arratia 1999
- ******* Order †Lycopteriformes Chang & Chou 1977
- ******* Order Hiodontiformes McAllister 1968 sensu Taverne 1979
- ******* Order Osteoglossiformes Regan 1909 sensu Zhang 2004
- ****** Supercohort Clupeocephala Patterson & Rosen 1977 sensu Arratia 2010
- *******Cohort Otomorpha Wiley & Johnson 2010
- ******** Subcohort Clupei Wiley & Johnson 2010
- ********* Order †Ellimmichthyiformes Grande 1982
- ********* Order Clupeiformes Bleeker 1859
- ******** Subcohort Alepocephali
- ********* Order Alepocephaliformes Marshall 1962
- ******** Subcohort Ostariophysi Sagemehl 1885
- ********* Section Anotophysa Sagemehl 1885
- ********** Order †Sorbininardiformes Taverne 1999
- ********** Order Gonorynchiformes Regan 1909
- ********* Section Otophysa Garstang 1931
- ********** Order Cypriniformes Bleeker 1859 sensu Goodrich 1909
- ********** Order Characiformes Goodrich 1909
- ********** Order Gymnotiformes Berg 1940
- ********** Order Siluriformes Cuvier 1817 sensu Hay 1929
- ******* Cohort Euteleosteomorpha
- ********Subcohort Lepidogalaxii
- ********* Lepidogalaxiiformes Betancur-Rodriguez et al. 2013
- ******** Subcohort Protacanthopterygii Greenwood et al. 1966 sensu Johnson & Patterson 1996
- ********* Order Argentiniformes
- ********* Order Galaxiiformes
- ********* Order Salmoniformes Bleeker 1859 sensu Nelson 1994
- ********* Order Esociformes Bleeker 1859
- ******** Subcohort Stomiati
- ********* Order Osmeriformes
- ********* Order Stomiatiformes Regan 1909
- ******** Subcohort Neoteleostei Nelson 1969
- *********Infracohort Ateleopodia
- ********** Order Ateleopodiformes
- ********* Infracohort Eurypterygia Rosen 1973
- **********Section Aulopa
- *********** Order Aulopiformes Rosen 1973
- ********** Section Ctenosquamata Rosen 1973
- ***********Subsection Myctophata
- ************ Order Myctophiformes Regan 1911
- *********** Subsection Acanthomorpha Betancur-Rodriguez et al. 2013
- ************Division Lampridacea Betancur-Rodriguez et al. 2013
- ************* Order Lampriformes Regan 1909
- ************ Division Paracanthomorphacea sensu Grande et al. 2013
- ************* Order Percopsiformes Berg 1937
- ************* Order †Sphenocephaliformes Rosen & Patterson 1969
- ************* Order Zeiformes Regan 1909
- ************* Order Stylephoriformes Miya et al. 2007
- ************* Order Gadiformes Goodrich 1909
- ************ Division Polymixiacea Betancur-Rodriguez et al. 2013
- ************* Order †Pattersonichthyiformes Gaudant 1976
- ************* Order †Ctenothrissiformes Berg 1937
- ************* Order Polymixiiformes Lowe 1838
- ************ Division Euacanthomorphacea Betancur-Rodriguez et al. 2013
- *************Subdivision Berycimorphaceae Betancur-Rodriguez et al. 2013
- ************** Order Beryciformes
- ************* Subdivision Holocentrimorphaceae Betancur-Rodriguez et al. 2013
- ************** Order Holocentriformes
- ************* Subdivision Percomorphaceae Betancur-Rodriguez et al. 2013
- **************Series Ophidiimopharia Betancur-Rodriguez et al. 2013
- *************** Order Ophidiiformes
- ************** Series Batrachoidimopharia Betancur-Rodriguez et al. 2013
- *************** Order Batrachoidiformes
- ************** Series Gobiomopharia Betancur-Rodriguez et al. 2013
- *************** Order Kurtiformes
- *************** Order Gobiiformes
- ************** Series Scombrimopharia Betancur-Rodriguez et al. 2013
- *************** Order Syngnathiformes
- *************** Order Scombriformes
- ************** Series Carangimopharia Betancur-Rodriguez et al. 2013
- *************** Subseries Anabantaria Betancur-Rodriguez et al. 2014
- **************** Order Synbranchiformes
- **************** Order Anabantiformes
- *************** Subseries Carangaria Betancur-Rodriguez et al. 2014
- **************** Carangaria incertae sedis
- **************** Order Istiophoriformes Betancur-Rodriguez 2013
- **************** Order Carangiformes
- **************** Order Pleuronectiformes Bleeker 1859
- *************** Subseries Ovalentaria Smith & Near 2012
- **************** Ovalentaria incertae sedis
- **************** Order Cichliformes Betancur-Rodriguez et al. 2013
- **************** Order Atheriniformes Rosen 1964
- **************** Order Cyprinodontiformes Berg 1940
- **************** Order Beloniformes Berg 1940
- **************** Order Mugiliformes Berg 1940
- **************** Order Blenniiformes Springer 1993
- **************** Order Gobiesociformes Gill 1872
- ************** Series Eupercaria