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(20 versioni intermedie di 5 utenti non mostrate)
Riga 1:
[[File:Ostracoderm and placoderm.jpg|right|thumb|250px|I primi pesci fossili: [[ostracodermi]] (sopra) e [[Placodermi]]]]
<center>
L'evoluzione dei pesci non è studiata come un singolo evento da quanto la riorganizzazione sistematica dei [[pesci]] ha portato al disconocimento di questi come gruppo monofiletico{{sfn|Lecointre|Le Guyader|2007|ps=}}.
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I primi pesci apparvero durante la cosidetta [[esplosione cambriana]], circa 530 milioni di anni fa: questi ritrovamenti [[fossili]] appartengono ad un gruppo di piccoli pesci corazzati senza mascelle, noti come [[ostracodermi]], oggi estinti. Un [[clade]] imparentato ed esistente, comprendente le [[Lampreda|lamprede]]. Le prime mascelle appaiono invece nei [[Placodermi]]. La diversità di questi vertebrati indica il vantaggio evolutivo di una [[bocca]] provvista di mascelle, anche se ancora non è chiaro chiaro se il vantaggio comprenda un morso con maggior resistenza, una respirazione migliorata o una combinazione di entrambi i fattori.
!width="150" bgcolor= #6495ED |[[Sottofamiglia]]
 
!width="150" bgcolor= #6495ED |[[Genere (tassonomia)|Genere]]
__TOC__
!width="200" bgcolor= #6495ED |[[Specie]]
{{clear}}
!width="100" bgcolor= #6495ED |Immagine
 
==Overview==
{{multiple image
| align = right
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| width = 360
| header = Vertebrate classes
| header_align =
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| footer =
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| image1 = Fish evolution.png
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| caption1 = Spindle diagram for the evolution of fish and other vertebrate classes.{{sfn|Benton|2005}}
Conventional classification has living [[vertebrate]]s as a subphylum grouped into eight classes based on traditional interpretations of gross [[anatomy|anatomical]] and [[Physiology|physiological]] traits. In turn, these classes are grouped into the vertebrates which have four limbs and those which don't, the [[tetrapod]]s and [[fish]]es. The [[extant taxon|extant]] vertebrate classes are:{{sfn|Romer|1970}}
 
: Fish:
::* [[jawless fishes]] (Agnatha)
::* [[cartilaginous fishes]] (Chondrichthyes)
::* [[ray-finned fishes]] (Actinopterygii)
::* [[lobe-finned fishes]] (Sarcopterygii)
: Tetrapods:
::* [[amphibian]]s (Amphibia)
::* [[reptile]]s (Reptilia)
::* [[bird]]s (Aves)
::* [[mammal]]s (Mammalia)
 
In addition to these are two classes of extinct jawed fishes, the armoured [[Placodermi|placoderms]] and the [[Acanthodii|spiny sharks]].
}}
 
{{further|Vertebrate paleontology|List of transitional fossils#Invertebrates to Fish}}
 
Fish may have evolved from an animal similar to a coral-like [[sea squirt]], whose larvae resemble primitive fish in important ways. The first ancestors of fish may have [[Neoteny|kept the larval form into adulthood]] (as some sea squirts do today), although perhaps the reverse is the case. [[Vertebrate]]s, among them the first [[fishes]], originated about 530 million years ago during the [[Cambrian explosion]], which saw the rise in organism diversity.{{sfn|Dawkins|2004|p=357}}
 
[[File:Ancestors.gif|thumb|left|Outdated evolutionary view of continual gradation (click to animate)|alt=Animation showing life at different evolutionary stages]]
[[File:Haikouichthys cropped.jpg|thumb|left|The early vertebrate ''[[Haikouichthys]]'', from about {{ma|518}} in China, may be the "ancestor to all vertebrates" and is one of the earliest known fish.{{r|Shu2003}}]]
 
The first ancestors of fish, or animals that were probably closely related to fish, were ''[[Pikaia]]'', ''[[Haikouichthys]]'' and ''[[Myllokunmingia]]''.{{r|Shu1999}}{{sfn|Dawkins|2004|p=357}} These three [[Genus (biology)|genera]] all appeared around 530 [[mya (unit)|Ma]]. ''Pikaia'' had a primitive [[notochord]], a structure that could have developed into a [[vertebral column]] later. Unlike the other fauna that dominated the Cambrian, these groups had the basic vertebrate body plan: a notochord, rudimentary vertebrae, and a well-defined head and tail.{{r|Waggoner2011}} All of these early vertebrates lacked [[jaw]]s in the common sense and relied on filter feeding close to the seabed.{{sfn|Haines|Chambers|2005}}
 
These were followed by indisputable fossil vertebrates in the form of heavily armoured fishes discovered in rocks from the [[Ordovician]] Period 500–430 [[mya (unit)|Ma]]. The colonisation of new [[Ecological niche|niche]]s resulted in massive body sizes. In this way, fishes with increasing sizes evolved during the early Paleozoic, such as the titanic [[placoderm]] ''[[Dunkleosteus]]'', which could grow 7&nbsp;meters long. The [[Devonian]] Period (395 to 345 Ma) brought in the changes that allowed primitive air-breathing fish to remain on land as long as they wished, thus becoming the first terrestrial vertebrates, the [[amphibian]]s.
 
The first [[Gnathostomata|jawed vertebrates]] appeared in the late [[Ordovician]] and became common in the [[Devonian]], often known as the "Age of Fishes".{{sfn|Encyclopædia Britannica|1954|p=107}} The two groups of [[bony fishes]], the [[actinopterygii]] and [[sarcopterygii]], evolved and became common.{{sfn|Berg|2004|p=599}} The Devonian also saw the demise of virtually all jawless fishes, save for lampreys and hagfish, as well as the [[Placodermi]], a group of armoured fish that dominated much of the late [[Silurian]]. The Devonian also saw the rise of the first [[Labyrinthodontia|labyrinthodonts]], which was a transitional between fishes and [[amphibians]].
 
The [[reptiles]] appeared from labyrinthodonts in the subsequent [[Carboniferous]] period. The [[anapsid]] and [[synapsid]] reptiles were common during the late [[Paleozoic]], while the [[diapsid]]s became dominant during the [[Mesozoic]]. In the sea, the [[bony fishes]] became dominant. The [[dinosaur]]s gave rise to the [[bird]]s in the [[Jurassic]].{{sfn|Cloudsley-Thompson|2005|p=6}} The demise of the dinosaurs at the end of the [[Cretaceous]] promoted expansion of the [[mammals]], which had evolved from the [[therapsid]]s, a group of synapsid reptiles, during the late [[Triassic]] Period.
 
The later radiations, such as those of fish in the Silurian and Devonian periods, involved fewer taxa, mainly with very similar body plans. The first animals to venture onto dry land were [[arthropod]]s. Some fish had lungs and strong, bony fins and could crawl onto the land also.
 
{{clear}}
 
==Pesci senza mascelle==
[[File:Lamprey attached.png|thumb|260px|right|A modern jawless fish, the [[lamprey]], attached to a modern jawed fish]]
[[File:Petromyzon marinus.003 - Aquarium Finisterrae.JPG|thumb|140px|right|{{center|Lamprey mouth}}]]
{{main|Agnatha}}
Jawless fishes belong to the [[superclass (biology)|superclass]] [[Agnatha]] in the [[phylum (biology)|phylum]] [[Chordata]], subphylum [[Vertebrata]]. Agnatha comes from the [[Greek Language|Greek]], and means "no jaws".<ref>SOED</ref> It excludes all vertebrates with jaws, known as [[gnathostome]]s. Although a minor element of modern marine [[Fauna (animals)|fauna]], jawless fish were prominent among the early fish in the early [[Paleozoic]]. Two types of [[Early Cambrian]] animal apparently having fins, [[vertebrate]] musculature, and gills are known from the early Cambrian [[Maotianshan shales]] of [[China]]: ''[[Haikouichthys]]'' and ''[[Myllokunmingia]]''. They have been tentatively assigned to Agnatha by Janvier. A third possible agnathid from the same region is ''[[Haikouella]]''. A possible agnathid that has not been formally described was reported by Simonetti from the Middle Cambrian [[Burgess Shale]] of [[British Columbia]].
 
Many Ordovician, Silurian, and Devonian agnathians were armoured with heavy bony-spiky plates. The first armoured agnathans—the [[Ostracoderm]]s, precursors to the [[bony fish]] and hence to the [[tetrapods]] (including humans)—are known from the middle [[Ordovician]], and by the Late [[Silurian]] the agnathans had reached the high point of their evolution. Most of the ostracoderms, such as [[thelodont]]s, [[osteostracan]]s, and [[galeaspid]]s, were more closely related to the gnathostomes than to the surviving agnathans, known as cyclostomes. Cyclostomes apparently split from other agnathans before the evolution of dentine and bone, which are present in many fossil agnathans, including [[conodonts]].<ref name="Baker 2008">{{cite journal | year = 2008 | title = The evolution and elaboration of vertebrate neural crest cells | journal = Current Opinion in Genetics & Development | volume = 18 | pages = 536–543 | doi = 10.1016/j.gde.2008.11.006 | author = Baker, Clare V.H. | pmid = 19121930 | issue = 6}}</ref> Agnathans declined in the [[Devonian]] and never recovered.
 
The agnathans as a whole are [[paraphyletic]],<ref>{{cite book|last=Purnell|first=M. A.|authorlink=|editor=[[Derek Briggs|Derek E. G. Briggs]] and Peter R. Crowther|year=2001|title=Palaeobiology II|publisher=[[Blackwell Publishing]]|___location=[[Oxford]]|isbn=0-632-05149-3|page=401}}</ref> because most extinct agnathans belong to the [[stem group]] of gnathostomes.<ref name="Zhao & Zhu 2007">{{cite journal | year = 2007 | title = Diversification and faunal shift of Siluro-Devonian vertebrates of China | journal = Geological Journal | volume = 42 | issue = (3-4) | pages = 351–369 | url = http://www3.interscience.wiley.com/journal/114129423/abstract | doi = 10.1002/gj.1072 | author = Zhao Wen-Jin, Zhu Min}}</ref><ref name="Sansom 2009">{{cite journal | author = Sansom, Robert S. | year = 2009 | title = Phylogeny, classification, & character polarity of the Osteostraci (Vertebrata) | journal = Journal of Systematic Palaeontology | volume = 7 | pages = 95–115 | url = http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=3978288 | doi = 10.1017/S1477201908002551}}</ref> Recent molecular data, both from rRNA<ref name="Mallatt, J., and J. Sullivan. 1998.">{{cite journal | author = Mallatt, J., and J. Sullivan. 1998. | year = 1998 | title = 28S and 18S ribosomal DNA sequences support the monophyly of lampreys and hagfishes. | journal = Molecular Biology and Evolution | volume = 15 | pages = 1706–1718 | pmid = 9866205 | issue = 12}}</ref> and from mtDNA<ref name="DeLarbre et al. 2002">{{cite journal | author = DeLarbre Christiane ; Gallut Cyril ; Barriel Veronique ; Janvier Philippe ; Gachelin Gabriel | year = 2002 | title = Complete mitochondrial DNA of the hagfish, Eptatretus burgeri: The comparative analysis of mitochondrial DNA sequences strongly supports the cyclostome monophyly. | journal = Molecular Phylogenetics & Evolution | volume = 22 | issue = 2 | pages = 184–192 | pmid = 11820840 | doi = 10.1006/mpev.2001.1045}}</ref> strongly supports the theory that living agnathans, known as [[cyclostomata|cyclostomes]], are [[monophyletic]].<ref>[[Philippe Janvier|Janvier, P.]] 2010. "MicroRNAs revive old views about jawless vertebrate divergence and evolution." Proceedings of the National Academy of Sciences (USA) 107:19137-19138. [http://www.pnas.org/content/107/45/19137.full.pdf+html] "''Although I was among the early supporters of vertebrate paraphyly, I am impressed by the evidence provided by Heimberg et al. and prepared to admit that cyclostomes are, in fact, monophyletic. The consequence is that they may tell us little, if anything, about the dawn of vertebrate evolution, except that the intuitions of 19th century zoologists were correct in assuming that these odd vertebrates (notably, hagfishes) are strongly degenerate and have lost many characters over time.''"</ref> In [[Phylogenetics|phylogenetic taxonomy]], the relationships between animals are not typically divided into ranks, but illustrated as a nested "family tree" known as a [[cladogram]]. Phylogenetic groups are given definitions based on their relationship to one another, rather than purely on physical traits such as the presence of a backbone. This nesting pattern is often combined with traditional taxonomy, in a practice known as [[evolutionary taxonomy]].
 
[[File:Evolution of jawless fish.png|thumb|260px|right|Evolution of [[Agnatha|jawless fishes]]. The diagram is based on [[Michael Benton]], 2005.<ref name="Benton, M. J. 2005 page 73">Benton, M. J. (2005) Vertebrate Palaeontology, Blackwell, 3rd edition, Fig 3.25 on page 73.</ref>]]
 
Il [[cladogramma]] sottostante dei pesci senza mandibole è basato sugli studi effettuati [[Philippe Janvier]] e altri per il progetto ''Tree of Life Web Project''.<ref name=tol>Janvier, Philippe (1997) [http://tolweb.org/Vertebrata/14829/1997.01.01 Vertebrata. Animals with backbones]. Version 01 January 1997 in [http://tolweb.org/ The Tree of Life Web Project]</ref> <small>(† = indica che il gruppo è [[estinzione|estinto]])</small>
 
{{clade| style=font-size:85%;line-height:70%
|label1=[[Agnatha|'''Pesci senza mascelle''']]
|1={{clade
|1=[[Hyperoartia]] (lamprede)
|2=?†[[Euconodonta]] (Animali simili ad anguille)
|label3=<span style="color:white;">unnamed</span>
|3={{clade
|1=†[[Pteraspidomorphi]] (Pesci senza mascelle)
|2=?†[[Thelodonti]] (Pesci senza mascelle con scaglie)
|label3=<span style="color:white;">unnamed</span>
|3={{clade
|1=?†[[Anaspida]] (antenati senza mascelle delle lamprede)
|label2=<span style="color:white;">unnamed</span>
|2={{clade
|1=†[[Galeaspida]] (Pesci senza mascelle con scudo osseo cranico)
|label2=<span style="color:white;">unnamed</span>
|2={{clade
|1=?†[[Pituriaspida]] (Pesci senza mandibola corazzati con rostro)
|2=†[[Osteostraci]] (Pesci senza mandibola corazzati con scudo osseo cranico)
|3=[[Gnathostomata|'''Vertebrati con mandibola''']] (vedi la prossima sezione)
}}
}}
}}
}}
}}
}}
 
{{clear}}
 
===<sup>†</sup>Conodonti===
[[File:Euconodonta.gif|thumb|140px|right|{{center|<sup>†</sup>[[Conodonts]] (extinct) resembled primitive jawless eels}}]]
 
[[Conodont]]s resembled primitive jawless eels. They appeared 495 Ma and were wiped out 200 Ma.{{r|Renzi1996}} Initially they were known only from tooth-like microfossils called ''conodont elements''. These "teeth" have been variously interpreted as filter-feeding apparatuses or as a "grasping and crushing array".{{r|Gabbott1995}} Conodonts ranged in length from a centimeter to the 40&nbsp;cm ''[[Promissum]]''.{{r|Gabbott1995}} Their large eyes had a lateral position of which makes a predatory role unlikely. The preserved musculature hints that some conodonts (''Promissum'' at least) were efficient cruisers but incapable of bursts of speed.{{r|Gabbott1995}} In 2012 researchers classify the conodonts in the [[Phylum (biology)|phylum]] [[Chordata]] on the basis of their fins with fin rays, [[Chevron (insignia)|chevron]]-shaped muscles and [[notochord]].{{r|Briggs1992}} Some researchers see them as vertebrates similar in appearance to modern [[hagfish]] and [[lamprey]]s,{{r|Milsom2004}} though [[phylogenetic]] analysis suggests that they are more [[derived]] than either of these groups.{{r|Donoghue2000}}
 
===<sup>†</sup>Ostracodermi===
[[File:Astraspis desiderata.gif|thumb|140px|right|{{center|<sup>†</sup>[[Ostracoderms]] (extinct) were armoured jawless fishes}}]]
 
[[Ostracoderms]] ''(shell-skinned)'' are [[Armour (zoology)|armoured]] [[Agnatha|jawless fish]]es of the [[Paleozoic]]. The term does not often appear in classifications today because it is [[paraphyletic]] or [[polyphyletic]], and has no phylogenetic meaning.{{sfn|Benton|1999|p=44}} However, the term is still used informally to group together the armoured jawless fishes.
 
The ostracoderm armour consisted of 3–5&nbsp;mm polygonal plates which shielded the head and gills, and then overlapped further down the body like scales. The eyes were particularly shielded. Earlier [[chordates]] used their [[gill]]s for both respiration and feeding, whereas ostracoderms used their gills for [[Respiration (physiology)|respiration]] only. They had up to eight separate pharyngeal gill pouches along the side of the head, which were permanently open with no protective [[Operculum (fish)|operculum]]. Unlike [[invertebrate]]s that use [[Cilium|cilia]]ted motion to move food, ostracoderms used their muscular pharynx to create a suction that pulled small and slow moving [[prey]] into their mouths.
 
The first fossil fishes that were discovered were ostracoderms. The [[Swiss]] anatomist [[Louis Agassiz]] received some fossils of bony armored fish from [[Scotland]] in the 1830s. He had a hard time classifying them as they did not resemble any living creature. He compared them at first with extant armored fish such as [[catfish]] and [[sturgeon]]s but later realizing that they had no movable jaws, classified them in 1844 into a new group "ostracoderms".<ref>{{cite book |title=Discovering Fossil Fishes |last=Maisey |first=John G. |year=1996 |publisher=Henry Holt & Company|___location=New York |isbn= |page=37 |edition=illustrated}}</ref>
 
Ostracoderms existed in two major groups, the more primitive [[Heterostraci|heterostracans]] and the [[cephalaspidomorphi|cephalaspids]]. Later, about 420 million years ago, the jawed fish evolved from one of the ostracoderms. After the appearance of jawed fish, most ostracoderm species underwent a decline, and the last ostracoderms became extinct at the end of the [[Devonian period]].<ref>[http://www.cosmosmagazine.com/news/4492/vertebrate-jaw-design-locked-down-early Vertebrate jaw design locked down early]</ref>
 
==Pesci con mascella==
[[File:Dunkleosteus skull steveoc.jpg|thumb|360px|right|]]
[[File:DunkleosteusSannoble.JPG|thumb|216px|right|
{{ external media
| align = right
| width = 216px
| video1 = [http://www.youtube.com/watch?v=ga6n2vnM3Nc Evolution Of Jaws 1 of 5] ''Animal Planet''
| video2 = [http://www.youtube.com/watch?feature=endscreen&NR=1&v=vrjAtZZCpdg Evolution Of Jaws 2 of 5] ''Animal Planet''
| video3 = [http://www.youtube.com/watch?v=br6B_m-3NHc&feature=endscreen&NR=1 Evolution Of Jaws 3 of 5] ''Animal Planet''
}}
]]
{{further2|[[Gnathostomata]]|[[Fish jaw]]}}
 
The vertebrate jaw probably originally evolved in the [[Silurian]] period and appeared in the [[Placodermi|Placoderm]] [[fish]] which further diversified in the [[Devonian]]. The two most anterior [[pharyngeal arch]]es are thought to have become the jaw itself and the hyoid arch, respectively. The hyoid system suspends the jaw from the braincase of the skull, permitting great mobility of the jaws. While there is no fossil evidence directly to support this theory, it makes sense in light of the numbers of pharyngeal arches that are visible in extant jawed vertebrates (the [[Gnathostomata|Gnathostomes]]), which have seven arches, and primitive jawless vertebrates (the [[Agnatha]]), which have nine.
 
As in most [[vertebrate]]s, fish jaws are [[bone|bony]] or [[cartilage|cartilaginous]] and oppose vertically, comprising an ''upper jaw'' and a ''lower jaw''. The jaw is derived from the most anterior two [[pharyngeal arch]]es supporting the gills, and usually bears numerous [[teeth]].
 
It is thought that the original selective advantage garnered by the jaw was not related to feeding, but to increased respiration efficiency. The jaws were used in the [[buccal pump]] (observable in modern fish and [[amphibians]]) that pumps water across the gills of fish or air into the lungs in the case of amphibians. Over evolutionary time the more familiar use of jaws (to humans), in feeding, was selected for and became a very important function in vertebrates. Many [[Teleostei|teleost]] fish have substantially modified their jaws for [[suction feeding]] and [[cranial kinesis|jaw protrusion]], resulting in highly complex jaws with dozens of bones involved.
 
Jawed vertebrates and jawed fish evolved from jawless fish, and the [[cladogram]] below for jawed vertebrates is a continuation of the cladogram in the section above. <small>(† = group is extinct)</small>
 
{{clade| style=font-size:85%;line-height:70%
|label1='''[[Gnathostomata|Jawed vertebrates]]'''
|1={{clade
|1=†[[Placodermi]] (armoured fishes)
|label2= <span style="color:white;">unnamed</span>
|2={{clade
|1=[[Chondrichthyes]] (cartilaginous fishes)
|label2=[[Teleostomi]]
|2={{clade
|1=†[[Acanthodii]] ([[spiny shark]]s)
|label2=[[Osteichthyes|Bony fishes]]
|2={{clade
|1=[[Actinopterygii]] (ray-finned fishes) <dominant class of fish today
|label2=[[Sarcopterygii|Lobe-finned fishes]]
|2={{clade
|1=?†[[Onychodontiformes]] (lobe-finned)
|2=[[Actinistia]] (coelacanths)
|label3= <span style="color:white;">unnamed</span>
|3={{clade
|1=†[[Porolepiformes]] (lobe-finned)
|2=[[Dipnoi]] (lungfishes)
}}
|label4= <span style="color:white;">unnamed</span>
|4={{clade
|1=†[[Rhizodontidae|Rhizodontimorpha]] (predatory lobe-finned)
|2={{clade
|1=†[[Tristichopteridae]] ([[tetrapodomorph]]s)
|2=[[Tetrapoda|'''Tetrapods''']] (four legged animals)
}}
}}
}}
}}
}}
}}
}}
}}
 
{{clear}}
 
===<sup>†</sup>Placodermi===
[[File:Evolution of placoderms.png|thumb|260px|right|Evolution of the (now extinct) [[placoderms]]. The diagram is based on [[Michael Benton]], 2005.<ref name="Benton, M. J. 2005 page 73"/>]]
[[File:Coccosteus BW.jpg|thumb|left|<sup>†</sup>[[Placoderm]]s (extinct) were armoured jawed fishes (compare with the ostracoderms above)]]
{{further2|[[Placodermi]]|[[List of placoderms]]}}
 
[[Placoderm]]s, [[Class (biology)|class]] Placodermi ''(plate skinned)'', are [[Extinction (biology)|extinct]] armoured prehistoric fish, which appeared about 430 Ma in the late Silurian. They were mostly wiped out at the end of the Devonian 360 Ma, though a few survived another 5 million years. Their head and thorax were covered with massive and often ornamented armoured plates. The rest of the body was [[Fish scale|scaled]] or naked, depending on the species. The armour shield was articulated, with the head armour hinged to the thoratic armour. This allowed placoderms to lift their heads, unlike ostracoderms. Placoderms were among the first jawed fish; their jaws likely evolved from the first of their [[gill]] arches. The chart on the right shows the rise and demise of the separate placoderm lineages: [[Acanthothoraci]], [[Rhenanida]], [[Antiarchi]], [[Petalichthyidae]], [[Ptyctodontida]] and [[Arthrodira]].
 
===<sup>†</sup>Spiny sharks===
[[File:Acanthodes BW.jpg|thumb|left|<sup>†</sup>[[Spiny shark]]s (extinct) were the earliest known jawed fishes. They resembled sharks but were an independent branch.]]
{{further2|[[Acanthodii]]|[[List of acanthodians]]}}
 
[[Spiny shark]]s, class Acanthodii, are extinct fishes which share features with both bony fishes and cartilaginous fishes. Despite being called "spiny sharks," acanthodians predate sharks. They evolved in the sea at the beginning of the Silurian Period, some 50 million years before the first sharks appeared. Eventually competition from bony fishes proved too much, and the spiny sharks died out in Permian times about 250 Ma. In form they resembled sharks, but their [[Epidermis (skin)|epidermis]] was covered with tiny rhomboid platelets like the scales of [[holostean]]s ([[gar]]s, [[bowfin]]s). They may have been an independent phylogenetic branch of fishes, which had evolved from little-specialized forms close to recent cartilaginous fishes.
 
{{clear}}
 
===Pesci cartilaginei===
[[File:Evolution of cartilaginous fishes.png|thumb|260px|right|Radiation of [[cartilaginous fish]]es, derived from work by [[Michael Benton]], 2005.<ref>Benton, M. J. (2005) Vertebrate Palaeontology, Blackwell, 3rd edition, Fig 7.13 on page 185.</ref>]]
{{further2|[[List of transitional fossils#Chondrichthyes]]|[[List of prehistoric cartilaginous fish]]}}
 
Cartilaginous fishes, class [[Chondrichthyes]], consisting of [[shark]]s, [[Batoidea|rays]] and [[chimaera]]s, appeared by about 395 million years ago, in the middle [[Devonian]]. The class contains the sub classes [[Holocephali]] ([[chimera]]) and [[Elasmobranchii]] ([[shark]]s and [[Batoidea|rays]]). The radiation of elasmobranches in the chart on the right is divided into the taxa: [[Cladoselache]], [[Eugeneodontiformes]], [[Symmoriida]], [[Xenacanthiformes]], [[Ctenacanthiformes]], [[Hybodontiformes]], [[Galeomorphi]], [[Squaliformes]] and [[Batoidea]].
 
{{clear}}
 
===Pesci ossei===
{{further2|[[Osteichthyes]]|[[List of transitional fossils#Bony Fish]]|[[List of prehistoric bony fish]]}}
{{ external media
| align =
| width = 250px
| video1 = [http://www.youtube.com/watch?v=k-5oQlnXSTM Chordate Evolution and Bony Fish] ''YouTube''
}}
 
[[Bony fish]]es, class Osteichthyes, are characterised by bony skeleton rather than [[cartilage]]. They appeared in the late [[Silurian]] or early [[Devonian]], about 416 million years ago. Both bony and cartilaginous fishes may have arisen from either the spiny sharks or the placoderms. A subclass of the Osteichthyes, the ray-finned fishes ([[Actinopterygii]]), have become the [[Dominance (ecology)|dominant group]] of fishes in the post-Paleozoic and modern world, with some 30,000 living species.
 
The bony (and cartilaginous) fish groups that emerged after the Devonian, were characterised by steady improvements in foraging and locomotion.{{sfn|Helfman|others|2009|p=198}}
 
====Lobe-finned fishes====
{{further2|[[Sarcopterygii]]|[[List of sarcopterygians|List of lobe-finned fishes]]}}
[[File:Barramunda.jpg|thumb|left|The [[Queensland lungfish]] is a lobe-finned fish which is a [[living fossil]]. Lungfish evolved the first proto-lungs and proto-limbs. They developed the ability to live outside a water environment in the middle [[Devonian]] (397-385 Ma), and have remained virtually the same for over 100 million years.<ref name = Allen>{{cite book | last = Allen | first = G.R., S.H. Midgley, M. Allen | title = Field Guide to the Freshwater Fishes of Australia. Eds. Jan Knight/Wendy Bulgin | publisher = Perth, W.A.: Western Australia Museum, 2002 | pages = 54–55}}</ref>]]
 
[[Lobe-finned fish]]es, fish belonging to the class Sarcopterygii, are mostly extinct bony fishes, basally characterised by robust and stubby lobe fins containing a robust internal skeleton, [[cosmoid scale]]s and internal nostrils. Their fins are fleshy, [[Lobe (anatomy)|lobed]], paired fins, joined to the body by a single bone.<ref>Clack, J. A. (2002) ''Gaining Ground''. Indiana University</ref> The fins of lobe-finned fish differ from those of all other fish in that each is borne on a fleshy, lobelike, scaly stalk extending from the body. The [[Pectoral fin|pectoral]] and [[pelvic fin]]s are articulated in ways resembling the tetrapod limbs they were the precursors to. The fins evolved into the legs of the first tetrapod land vertebrates, [[amphibian]]s. They also possess two dorsal fins with separate bases, as opposed to the single dorsal fin of [[ray-finned fish]]. The braincase of lobe-finned fishes primitively has a hinge line, but this is lost in tetrapods and lungfish. Many early lobe-finned fishes have a symmetrical tail. All lobe-finned fishes possess teeth covered with true [[tooth enamel|enamel]].
 
Lobe-finned fishes, such as [[coelacanth]]s and [[lungfish]], were the most diverse group of bony fishes in the Devonian. Taxonomists who subscribe to the cladistic approach include the grouping [[Tetrapoda]] within the Sarcopterygii, and the tetrapods in turn include all species of four-limbed vertebrates.{{sfn|Nelson|2006}} The fin-limbs of lobe-finned fishes such as the coelacanths show a strong similarity to the expected ancestral form of tetrapod limbs. The lobe-finned fish apparently followed two different lines of development and are accordingly separated into two subclasses, the [[Rhipidistia]] (including the lungfish, and the [[Tetrapodomorpha]] which include the Tetrapoda) and the [[Actinistia]] (coelacanths). The first lobe-finned fishes, found in the uppermost [[Silurian]] (ca 418 Ma), closely resembled [[spiny shark]]s, which went extinct at the end of the Paleozoic. In the early–middle [[Devonian]] (416 - 385 Ma), while the predatory [[placoderms]] dominated the seas, some lobe-finned fishes came into [[freshwater]] habitats.
 
[[File:Coelacanth-bgiu.png|thumb|right|The [[coelacanth]] is another lobe finned fish which is a living fossil. It is thought to have evolved into roughly its current form about 408 million years ago, during the early Devonian,{{r|Johanson2006}} and has not essentially evolved further from its ancient form.{{sfn|Forey|1998}}]]
 
In the Early Devonian (416-397 Ma), the lobe-finned fishes split into two main lineages — the [[coelacanth]]s and the [[rhipidistian]]s. The former never left the oceans and their heyday was the [[Late Devonian]] and [[Carboniferous]], from 385 to 299 Ma, as they were more common during those periods than in any other period in the Phanerozoic; coelacanths still live today in the [[ocean]]s (genus ''[[Latimeria]]''). The Rhipidistians, whose ancestors probably lived in [[estuaries]], migrated into freshwater habitats. They in turn split into two major groups: the [[lungfish]] and the [[tetrapodomorph]]s. The lungfish's greatest diversity was in the [[Triassic]] period; today there are fewer than a dozen genera left. The lungfish evolved the first proto-lungs and proto-limbs; developing the ability to live outside a water environment in the middle Devonian (397-385 Ma). The first tetrapodomorphs, which included the gigantic rhizodonts, had the same general anatomy as the lungfish, who were their closest kin, but they appear not to have left their water habitat until the late Devonian epoch (385 - 359 Ma), with the appearance of [[tetrapods]] (four-legged vertebrates). Tetrapods are the only tetrapodomorphs which survived after the Devonian. Lobe-finned fishes continued until towards the end of Paleozoic era, suffering heavy losses during the [[Permian-Triassic extinction event]] (251 Ma).
 
====Ray-finned fishes====
[[File:Evolution of ray-finned fish.png|thumb|500px|right]]
 
[[Ray-finned fishes]], class Actinopterygii, differ from lobe-finned fishes in that their fins consist of webs of skin supported by spines ("rays") made of bone or horn. There are other differences in respiratory and circulatory structures. Ray-finned fishes normally have skeletons made from true bone, though this is not true of [[sturgeons]] and [[paddlefish]]es.<ref name="ucmp">[http://www.ucmp.berkeley.edu/vertebrates/actinopterygii/actinintro.html Introduction to the Actinopterygii] ''Museum of Palaeontology'', University of California.</ref>
 
Ray-finned fishes are the dominant vertebrate group, containing half of all known vertebrate species. They inhabit abyssal depths in the sea, coastal inlets and freshwater rivers and lakes, and are a major source of food for humans.<ref name="ucmp" />
 
{{clear}}
 
==Timeline==
{{Phanerozoic eon}}
 
===Pre-Devoniano: origine dei pesci===
{| class="wikitable"
|-
! rowspan=6 style=background:#708050 | [[Cambriano|<span style="color:white;">Cambriano</span>]]
|rowspan="240"|[[Pseudocrenilabrinae]]
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Cambriano]] (542–488 milioni di anni fa): l'inizio del Cambriano è caratterizzato dalla cosiddetta "[[Esplosione cambriana]]", un'improvvisa apparizione di quasi tutti i [[phyla]] animali invertebrati ([[molluschi]], [[meduse]], [[vermi]] e [[artropodi]], come i [[crostacei]]) in grande abbondanza. I primi vertebrati apparvero sotto forma di pesci primitivi, che si sono evoluti poi in modo molto diversificato nel [[Siluriano]] e [[Devoniano]].
|rowspan="226"|'''[[Haplochromis]]'''
|-
| [[Haplochromis acidens]] ||
| [[File:Pikaia BW.jpg|140px]]
|-
| ''[[Pikaia]]''
| [[Haplochromis adolphifrederici]] ||
|-
''Pikaia'', along with ''[[Myllokunmingia]]'' and ''[[Haikouichthys ercaicunensis]]'' immediately below, are all candidates in the fossil record for the titles of "first vertebrate" and "first fish". ''Pikaia'' is a [[Genus (biology)|genus]] that appeared about 530 [[mya (unit)|Ma]] during the [[Cambrian explosion]] of multicellular life. [[Pikaia gracilens]] ''(pictured)'' is a [[transitional fossil]] between invertebrates and vertebrates,{{sfn|Dawkins|2004|p=289|ps=: "Obviously vertebrates must have had ancestors living in the Cambrian, but they were assumed to be invertebrate forerunners of the true vertebrates — protochordates. Pikaia has been heavily promoted as the oldest fossil protochordate."}} and may be the earliest known [[chordate]].{{r|Morris1979}}{{r|Morris2012}} In this sense it may have been the original ancestor of fishes. It was a primitive creature with no evidence of eyes, without a well defined head, and less than 2&nbsp;inches (5 centimetres) long. ''Pikaia'' was a sideways-flattened, leaf-shaped animal which swam by throwing its body into a series of S-shaped, zig-zag curves, similar to movement of snakes. Fish inherited the same swimming movement, but they generally have stiffer backbones. It had a pair of large head tentacles and a series of short appendages, which may be linked to gill slits, on either side of its head. ''Pikaia'' shows the essential prerequisites for [[vertebrates]]. The flattened body is divided into pairs of [[myomere|segmented muscle block]]s, seen as faint vertical lines. The muscles lie on either side of a flexible structure resembling a rod that runs from the tip of the head to the tip of the tail.{{sfn|Palmer|2000|p=66-67}}
| [[Haplochromis aelocephalus]] ||
|-
| [[File:Haikouichthys 3d.png|140px]]
| [[Haplochromis aeneocolor]] ||
| ''[[Haikouichthys]]''
|-
| ''[[Haikouichthys]]'' (il cui nome scientifico significa ''pesce di Haikou'') è un altro genere che appare nei ritrovamenti fossili di circa 530 milioni di anni fa, e segnala la transizione da [[invertebrati]] a [[vetrebrati]]{{sfn|Shu2003}} Gli Haikouichthys appartengono ai [[Craniata|craniati]] (animali con teschio e testa distinta dal corpo). Diversamente dai ''Pikaia'', vi è la presenza di occhi. Inoltre hanno un teschio definito e altre caratteristiche che hanno convinto i paleontologi a classificarli come craniati e come il primo vero pesce. Analisi cladistiche successive hanno suggerito che questi animali fossero [[chordata|cordati]] o [[craniata|craniati]]; {{r|Paleos: The Cambrian: 2}} senza dare risultati indiscutibili e incontrovertibili, pertanto si fanno rientrare in entrambi i gruppi.{{r|Donoghue2005}}{{sfn|Shu2003}}
| [[Haplochromis akika]] ||
|-
| [[File:Myllokunmingia.png|140px]]
| [[Haplochromis albertianus]] ||
| ''[[Myllokunmingia]]''
|-
| ''[[Myllokunmingia]]'' è un genere apparso circa 530 milioni di anni fa. Apparteneva ai [[Cordata|cordati]], aveva dimensioni minute (28 mm di lunghezza e 6 di altezza) ed è tra le più antiche forme di vita [[vertebrati|vertebrate]].
| [[Haplochromis altigenis]] ||
|-
| [[File:Euconodonta.gif|140px]]
| [[Haplochromis ampullarostratus]] ||
| [[Conodonti]]
|-
| [[Conodont]]s ''(cone-teeth)'' resembled primitive eels. They appeared 495 Ma and were wiped out 200 Ma.{{r|Renzi1996}} Initially they were known only from tooth-like microfossils called ''conodont elements''. These "teeth" have been variously interpreted as filter-feeding apparatuses or as a "grasping and crushing array".{{r|Gabbott1995}} Conodonts ranged in length from a centimeter to the 40&nbsp;cm ''[[Promissum]]''.{{r|Gabbott1995}} Their large eyes had a lateral position of which makes a predatory role unlikely. The preserved musculature hints that some conodonts (''Promissum'' at least) were efficient cruisers but incapable of bursts of speed.{{r|Gabbott1995}} In 2012 researchers classify the conodonts in the [[Phylum (biology)|phylum]] [[Chordata]] on the basis of their fins with fin rays, [[Chevron (insignia)|chevron]]-shaped muscles and [[notochord]].{{r|Briggs1992}} Some researchers see them as vertebrates similar in appearance to modern [[hagfish]] and [[lamprey]]s,{{r|Milsom2004}} though [[phylogenetic]] analysis suggests that they are more [[derived]] than either of these groups.{{r|Donoghue2000}}
| [[Haplochromis angustifrons]] ||
|-
| [[File:Thelodonti.gif|140px]]
| [[Haplochromis annectidens]] ||
| [[Ostracodermi]]
|-
| [[Ostracoderms]] ''(shell-skinned)'' are any of several groups of extinct, primitive, jawless fishes that were covered in an armour of bony plates. They appeared in the [[Cambrian]], about 510 million years ago, and became [[extinct]] towards the end of the [[Devonian]], about 377 million years ago. Initially Ostracoderms had poorly formed fins, and paired [[Fish fin|fins]], or limbs, first evolved within this group. They were covered with a bony armour or scales and were often less than 30&nbsp;cm (1&nbsp;ft) long.
| [[Haplochromis antleter]] ||
|-
! rowspan=6 style=background:#20A090 | [[Ordoviciano|<span style="color:white;">Ordoviciano</span>]]
| [[Haplochromis apogonoides]] ||
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Ordoviciano]] (488–443 milioni di anni fa): i pesci, la prima forma di vita vertebrata, continuano ad evolvere: compaiono i primi pesci con mandibole ([[Gnathostomata]]). </br>La vita sulla terraferma non si è ancora differenziata.
|-
|-
| [[Haplochromis arcanus]] ||
| [[File:Larnovaspis stensioei.jpg|140px]]
|-
| [[Heterostraci]]
| [[Haplochromis argenteus]] ||
| [[Heterostraci]] è una classe estinta di agnati vertebrati che viveva principalmente in ambienti marini e di estuario. Sono apparsi all'inizio dell'Ordoviciano e si estinse alla fine del Devoniano. Differivano dagli altri pesci senza mandibola del periodo nella disposizione e nell'[[istologia]] delle loro [[scaglia|scaglie]].
|-
|-
| [[Haplochromis artaxerxes]] ||
| [[File:Astraspis desiderata.gif|center|110px]]
|-
| ''[[Astraspis]]''
| [[Haplochromis astatodon]] ||
| ''[[Astraspis]]'' ''(star shield)'' is an extinct genus of primitive jawless fish related to other Ordovician fishes, such as ''[[Sacabambaspis]]'' and ''[[Arandaspis]]''. Fossils show clear evidence of a sensory structure (lateral line system). The arrangement of these organs in regular lines allows the fish to detect the direction and distance from which a disturbance in the water is coming. ''Arandaspis'' are thought to have had a mobile tail covered with small protective plates and a head region covered with larger plates. A specimen described by Sansom ''et al''. had relatively large, lateral eyes and a series of eight [[gill]] openings on each side.{{r|Sansom1997}}
|-
|-
| [[Haplochromis avium]] ||
| [[File:Arandaspis NT.jpg|140px]]
|-
| [[Pteraspidomorphi]]
| [[Haplochromis azureus]] ||
| [[Pteraspidomorphi]] is an extinct class of early jawless fish. The fossils show extensive shielding of the head. Many had hypocercal tails in order to generate lift to increase ease of movement through the water for their armoured bodies, which were covered in dermal bone. They also had sucking mouth parts and some species may have lived in fresh water.
|-
|-
| [[Haplochromis barbarae]] ||
| [[File:Shielia tiati.svg|center|125px]]
|-
| [[Haplochromis bareliThelodonti]] ||
| [[Thelodonts]] ''(nipple teeth)'' are a class of small, extinct jawless fishes with distinctive scales instead of large plates of armour. There is debate over whether these represent a [[Monophyly|monophyletic grouping]], or disparate stem groups to the major lines of [[Agnatha|jawless]] and [[Gnathostome|jawed fish]].{{r|Turner1982}} Thelodonts are united by their characteristic "thelodont scales". This defining character is not necessarily a result of shared ancestry, as it may have been [[Convergent evolution|evolved independently by different groups]]. Thus the thelodonts are generally thought to represent a polyphyletic group.{{sfn|Sarjeant|Halstead}} If they are monophyletic, there is no firm evidence on what their ancestral state was.{{sfn|Donoghue|2000|p=206}} These scales were easily dispersed after death; their small size and resilience makes them the most common vertebrate fossil of their time.{{sfn|Turner1999|p=42–78}}{{r|Palaeos: Thelodonti}} The fish lived in both freshwater and marine environments, first appearing during the [[Ordovician]], and perishing during the [[Late Devonian extinction|Frasnian–Famennian extinction event]] of the Late [[Devonian]]. They were predominantly deposit-feeding bottom dwellers, although some species may have been pelagic.
|-
|-
| [[Haplochromis bartoni]] ||
| colspan="3" style="line-height:16px; background:#dddddd;"| <small>The Ordovician ended with the [[Ordovician–Silurian extinction event]] (450–440 Ma). Two events occurred that killed off 27% of all families, 57% of all genera and 60% to 70% of all species.{{r|Baez2006}} Together they are ranked by many scientists as the second largest of the five major extinctions in Earth's history in terms of percentage of [[genus|genera]] that went extinct.</small>
|-
|-
| [[Haplochromis bayoni]] ||
! rowspan=7 style=background:#60C090 | [[Siluriano]]
|-
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Silurian]] (443–419 Ma): Many evolutionary milestones occurred during this period, including the appearance of armoured jawless fish, jawed fish, spiny sharks and ray-finned fish.
| [[Haplochromis beadlei]] ||
|-
| [[File:Silurianfishes ntm 1905 smit 1929.gif|140px]]
| [[Haplochromis bicolor]] ||
|-
| While it is traditional to refer to the Devonian as the age of fishes, recent findings have shown the Silurian was also a period of considerable diversification. Jawed fish developed movable [[jaw]]s, adapted from the supports of the front two or three [[gills|gill arches]]
| [[Haplochromis boops]] ||
|-
| [[File:Jamoytius kerwoodi.jpg|center|140px]]
| [[Haplochromis brownae]] ||
| [[Anaspida]]
|-
| [[Anaspida]] ''(without shield)'' is an extinct class of primitive jawless vertebrates that lived during the [[Silurian]] and [[Devonian]] periods.{{sfn|Ahlberg|2001|p=188}} They are classically regarded as the ancestors of [[lamprey]]s.{{sfn|Patterson|1987|p=142}} Anaspids were small, primarily marine [[agnatha]]ns that lacked heavy bony shield and paired fins, but have highly exaggerated [[hypocercal]] tails. They first appeared in the early [[Silurian]], and flourished until the [[Late Devonian extinction]],{{sfn|Hall|Hanken|1993|p=131}} where most species, save for [[lamprey]]s, went extinct. Unusually for an [[agnathan]], anaspids did not possess a bony shield or armour. The head is instead covered in an array of smaller, weakly mineralised scales.{{sfn|Janvier|2003}}
| [[Haplochromis bullatus]] ||
|-
| [[File:Acanthodii.jpg|140px]]
| [[Haplochromis bwathondii]] ||
| [[Spiny shark]]s
|-
| [[Spiny shark]]s, more formally called "Acanthodians" ''(having spines)'', comprise of the class Acanthodii. They first appeared by the late Silurian ~420 Ma, and were among the first fishes to evolve jaws. They share features with both [[cartilaginous fish]] and [[bony fish]], but they are not true sharks. They became extinct before the end of the Permian ~250 Ma. However, scales and teeth attributed to this group, as well as more derived jawed fish, such as cartilaginous and bony fish, date from the [[Ordovician]] ~460 Ma. Acanthodians were generally small shark-like fishes varying from toothless filter-feeders to toothed predators. They were once often classified as an order of the class [[Placodermi]], but recent authorities tend to place the acanthodians nearer to or within the living jawed fishes. They are distinguished in two respects: they were the earliest known jawed vertebrates, and they had stout [[Spine (zoology)|spines]] supporting all their [[fin]]s, fixed in place and non-movable (like a [[shark]]'s [[dorsal fin]]), an important defensive adaptation. Their fossils are extremely rare.
| [[Haplochromis cassius]] ||
|-
| [[File:Silurolepis platydorsalis.jpg|140px]]
| [[Haplochromis cavifrons]] ||
| [[Placoderma]]s
|-
| [[Placodermi]]s, ''(plate-like skin)'', are a group of armoured jawed fishes, of the class Placodermi. The oldest fossils appeared during the late Silurian, and became extinct at the end of the Devonian. Recent studies suggest that the placoderms are possibly a [[paraphyletic]] group of [[Basal (phylogenetics)|basal]] jawed fishes, and the closest relatives of all living jawed vertebrates. Some placoderms were small, flattened bottom-dwellers, such as [[antiarch]]s. However many, particularly the arthrodires, were active midwater predators. ''[[Dunkleosteus]]'', which appeared later in the Devonian below, was the largest and most famous of these. The upper jaw was firmly fused to the skull, but there was a hinge joint between the skull and the bony plating of the trunk region. This allowed the upper part of the head to be thrown back and, in [[arthrodire]]s, allowed them to take larger bites.
| [[Haplochromis chilotes]] ||
|-
| [[File:Guiyu BW.jpg|140px]]
| [[Haplochromis chlorochrous]] ||
| ''[[Guiyu oneiros]]''
|-
| ''[[Guiyu oneiros]]'' è il più antico pesce osseo oggi conosciuto. Presenta una combinazione di caratteri tipici degli [[Actinopterygii]] e dei [[Sarcopterygii]], anche se l'analisi scientifica lo pone più vicino a questi ultimi.
| [[Haplochromis chromogynos]] ||
|-
|
| [[Haplochromis chrysogynaion]] ||
| ''[[Andreolepis]]''
|-
| Il genere ''[[Andreolepis]]'' include il più antico [[Actinopterygii|attinopterigio]] oggi conosciuto, ''[[Andreolepis hedei]]'', che comparve nel tardo [[Siluriano]], circa 420 milioni di anni fa.<ref>{{cite journal | author = Min Z | year = 1997 | title = The oldest sarcopterygian fish | url = http://www.researchgate.net/publication/225089993_The_oldest_sarcopterygian_fish/file/d912f4fcb0d2e1d901.pdf | format = PDF | journal = Lethaia | volume = 30 | issue = | pages = 293–304 }}</ref><ref>{{cite journal | author = Märss T | year = 2001 | title = ''Andreolepis'' (Actinopterygii) in the upper Silurian of northern Eurasia | url = http://books.google.co.nz/books?id=0D3ucZP97e4C&pg=PA174&dq=%22Andreolepis+%28Actinopterygii%29+in+the+upper+Silurian+of+northern+Eurasia%22&hl=en&sa=X&ei=tF8IUeHwHYH-lAWo_IGwBA&ved=0CC8Q6AEwAA#v=onepage&q=%22Andreolepis%20%28Actinopterygii%29%20in%20the%20upper%20Silurian%20of%20northern%20Eurasia%22&f=false | journal = Proceedings of the Estonian Academy of Sciences | volume = 50 | issue = 3| pages = 174–189 }}</ref>
| [[Haplochromis cinctus]] ||
|- }
 
| [[Haplochromis cinereus]] ||
===Devoniano: l'era dei pesci===
|-
{{include timeline|Fish}}
| [[Haplochromis cnester]] ||
 
|-
The Devonian Period is broken into the Early, Middle and Late Devonian. By the start of the Early Devonian 419 mya, [[jawed fish]]es had divided into four distinct clades: the [[placoderm]]s and [[spiny shark]]s, both of which are now extinct, and the [[Cartilaginous fish|cartilaginous]] and [[bony fish]]es, both of which are still extant. The modern bony fishes, class [[Osteichthyes]], appeared in the late [[Silurian]] or early Devonian, about 416 million years ago. Both the cartilaginous and bony fishes may have arisen from either the placoderms or the spiny sharks. A subclass of bony fishes, the [[ray-finned fish]]es (Actinopterygii), have become the [[Dominance (ecology)|dominant group]] in the post-Paleozoic and modern world, with some 30,000 living species.
| [[Haplochromis commutabilis]] ||
 
|-
Sea levels in the Devonian were generally high. Marine faunas were dominated by [[bryozoa]], diverse and abundant [[brachiopoda|brachiopods]], the enigmatic [[hederellid|hederelloids]], [[microconchids]] and [[coral]]s. Lily-like [[crinoids]] were abundant, and [[trilobite]]s were still fairly common. Among vertebrates, jawless armoured fish ([[ostracoderms]]) declined in diversity, while the jawed fish (gnathostomes) simultaneously increased in both the sea and [[fresh water]]. Armoured [[placoderms]] were numerous during the lower stages of the Devonian Period and became extinct in the Late Devonian, perhaps because of competition for food against the other fish species. Early cartilaginous ([[Chondrichthyes]]) and bony fishes ([[Osteichthyes]]) also become diverse and played a large role within the Devonian seas. The first abundant genus of shark, ''[[Cladoselache]]'', appeared in the oceans during the Devonian Period. The great diversity of fish around at the time, have led to the Devonian being given the name "The Age of Fish" in popular culture.
| [[Haplochromis coprologus]] ||
every known aquatic environment.
|-
 
| [[Haplochromis crassilabris]] ||
The first [[Actinopterygii|ray-finned]] and [[Sarcopterygii|lobe-finned]] [[bony fish]] appeared in the Devonian, while the [[placoderms]] began dominating almost every known aquatic environment. However, another subclass of Osteichthyes, the [[Sarcopterygii]], including lobe-finned fishes including [[coelacanth]]s and [[lungfish]]) and [[tetrapods]], was the most diverse group of bony fishes in the Devonian. Sarcopterygians are basally characterized by internal nostrils, lobe fins containing a robust internal skeleton, and [[cosmoid scale]]s.
|-
 
| [[Haplochromis crebridens]] ||
During the Middle Devonian 393–383 Ma, the armoured jawless [[ostracoderm]] fishes were declining in diversity; the jawed fish were thriving and increasing in diversity in both the oceans and freshwater. The shallow, warm, oxygen-depleted waters of Devonian inland lakes, surrounded by primitive plants, provided the environment necessary for certain early fish to develop essential characteristics such as well developed lungs, and the ability to crawl out of the water and onto the land for short periods of time. Cartilaginous fishes, class [[Chondrichthyes]], consisting of [[shark]]s, [[Batoidea|rays]] and [[chimaera]]s, appeared by about 395 million years ago, in the middle [[Devonian]]
|-
 
| [[Haplochromis crocopeplus]] ||
During the Late Devonian the first forests were taking shape on land. The first tetrapods appear in the fossil record over a period, the beginning and end of which are marked with extinction events. This lasted until the end of the Devonian 359 mya. The ancestors of all [[tetrapod]]s began adapting to walking on land, their strong pectoral and pelvic fins gradually evolved into legs (see ''[[Tiktaalik]]'').{{r|BBC2010}} In the oceans, primitive [[shark]]s became more numerous than in the [[Silurian]] and the [[late Ordovician]]. The first [[ammonite]] [[mollusk]]s appeared. [[Trilobite]]s, the mollusk-like [[brachiopod]]s and the great [[coral reef]]s, were still common.
|-
 
| [[Haplochromis cronus]] ||
The [[Late Devonian extinction]] severely affected marine life, killing off all placoderms, and all trilobites, save for a few species of the [[order (biology)|order]] [[Proetida]]. This major extinction occurred at the beginning of the last phase of the Devonian period, the Famennian faunal stage, (the Frasnian-Famennian boundary), about {{Geologic Ages Inline|Famennian}} Ma, when all the fossil [[agnathan]] fishes, save for the [[psammosteid]] [[heterostracan]]s, suddenly disappeared. A second strong pulse closed the Devonian period. The Late Devonian extinction was one of five major extinction events in the history of the Earth's biota, more drastic than the familiar extinction event that closed the Cretaceous. The Devonian extinction crisis primarily affected the marine community, and selectively affected shallow warm-water organisms rather than cool-water organisms. The most important group to be affected by this extinction event were the reef-builders of the great Devonian reef-systems.
|-
 
| [[Haplochromis cryptodon]] ||
* [[Hyneria]]
|-
 
| [[Haplochromis cryptogramma]] ||
<div>
|-
:::{| class="wikitable"
| [[Haplochromis cyaneus]] ||
|-
| colspan="5" style="text-align:center; background:#ddf8f8;"| [[Devoniano]] (419–359 mya): The start of Devonian saw the first appearance of [[lobe-finned fish]], precursors to the [[tetrapod]]s (animals with four limbs). Major groups of fish evolved during this period, often referred to as the '''age of fishes'''.{{r|AOF}} See [[:Category:Devonian fish]].
| [[Haplochromis decticostoma]] ||
|-
| colspan="5" |
| [[Haplochromis degeni]] ||
|-
! rowspan=18 style=background:#CD9C50 | <div align="center">'''[[Devoniano|D<br>e<br>v<br>o<br>n<br>i<br>a<br>n<br>o]]'''</div>
| [[Haplochromis dentex]] ||
| rowspan=6 style=background:#FFC060 | <div align="center">[[Devoniano inferiore]]</div>
|-
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Devoniano inferiore]] (419–393 Ma):
| [[Haplochromis dichrourus]] ||
|-
| [[File:Psarolepis BW.jpg|140px]]
| [[Haplochromis diplotaenia]] ||
| ''[[Psarolepis]]''
|-
| ''[[Psarolepis]]'' ''(speckled scale)'' is a genus of extinct [[lobe-finned fish]] which lived around 397 to 418 Ma. Fossils of ''Psarolepis'' have been found mainly in South [[People's Republic of China|China]] and described by [[paleontologist]] [[Xiaobo Yu]] in 1998. It is not known for certain which group ''Psarolepis'' belongs, but paleontologists agree that it probably is a [[basal (biology)|basal]] genus and seems to be close to the common ancestor of lobe-finned and [[ray-finned fish]]es.{{sfn|Benton|2005|p=65}}
| [[Haplochromis dolichorhynchus]] ||
|-
| [[File:Holoptychius nobilissimus.jpg|140px]]
| [[Haplochromis dolorosus]] ||
| ''[[Holoptychius]]''
|-
| ''[[Holoptychius]]'' is an extinct genus from the [[Order (biology)|order]] of [[porolepiform]] lobe-finned fish, extant from 416 to 359 Ma. It was a streamlined predator about {{convert|50|cm|in}} long (though it could grow up to 2.5 m), which fed on other [[bony fish]]. Its rounded scales and body form indicate that it could have swum quickly through the water to catch prey.{{sfn|The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|1999|p=43}}{{r|Holland2010}} Similar to other [[rhipidistia]]ns, it had fang-like [[teeth]] on its [[palate]] in addition to smaller teeth on the jaws. Its asymmetrical tail sported a [[caudal fin]] on its lower end. To compensate for the downward push caused by this fin placement, ''Holoptychius'''s [[pectoral fin]]s were placed high on the body.
| [[Haplochromis eduardianus]] ||
|-
| [[File:Ctenurella gladbachensis.jpg|center|100px]]
| [[Haplochromis eduardii]] ||
| [[Ptyctodontida]]
|-
| The [[ptyctodontids]] (''beak-teeth'') are an extinct [[monotypic]] [[order (biology)|order]] of unarmored [[placoderm]]s, containing only one family. They were extant from the start to the end of the Devonian. With their big heads, big eyes, and long bodies, the ptyctodontids bore a strong resemblance to modern day chimaeras ([[Holocephali]]). Their armor was reduced to a pattern of small plates around the head and neck. Like the extinct and related [[acanthothoracids]], and the living and unrelated holocephalians, most of the ptyctodontids are thought to have lived near the sea bottom and preyed on [[shellfish]].
| [[Haplochromis elegans]] ||
|-
| [[File:Lunaspis.jpg|center|100px]]
| [[Haplochromis empodisma]] ||
| [[Petalichthyida]]
|-
| The [[Petalichthyida]] was an order of small, flattened [[placoderm]]s which were extant from the beginning of the Devonian to the Late Devonian. They were typified by splayed fins and numerous tubercles that decorated all of the plates and scales of their armour. They reached a peak in diversity during the Early Devonian and were found throughout the world. Because they had compressed body forms, it is supposed they were bottom-dwellers that chased after or ambushed smaller fish. Their diet is not clear, as none of the fossil specimens found have preserved mouth parts.
| [[Haplochromis engystoma]] ||
|-
| [[File:Laccognathus embryi - reconstruction.jpg|center|100px]]
| [[Haplochromis erythrocephalus]] ||
| ''[[Laccognathus]]''
|-
| ''[[Laccognathus]]'' ''(pitted jaw)'' was a genus of [[amphibious fish|amphibious]] lobe-finned fish that existed 398–360 Ma.<ref name="ngs">{{cite web|url=http://news.nationalgeographic.com/news/2011/09/110912-ancient-fish-arctic-predator-devonian-fossils-animals-science/|title=Ancient Toothy Fish Found in Arctic—Giant Prowled Rivers|author=[[Christine Dell'Amore|Dell'Amore, C.]]|date=September 12, 2011|publisher=[[National Geographic Society|National Geographic]] Daily News|accessdate=September 13, 2011}}</ref> They were characterized by the three large pits (fossae) on the external surface of the lower jaw which may have had sensory functions.<ref name="avril">{{cite web|url=http://articles.philly.com/2011-09-12/news/30145487_1_tiktaalik-roseae-ted-daeschler/2|title=Fish fossil sheds light on 'Euramerica' phase|author=Tom Avril|date=September 12, 2011|publisher=The Inquirer|accessdate=September 15, 2011}}</ref> ''Laccognathus'' grew to {{convert|1|-|2|m|ft|0}} in length. They had very short dorsoventrally flattened heads, less than one-fifth the length of the body.<ref name="vorobyeva">{{cite journal|author=[[Emiliya Ivanovna Vorobyeva|Vorobyeva, E.I.]]|year=2006|title=A new species of ''Laccognathus'' (Porolepiform Crossopterygii) from the Devonian of Latvia |journal=[[Paleontological Journal|Palaeont. J.]]|volume=40|issue=3|pages=312&ndash;322|publisher=Physorg.com|doi=10.1134/S0031030106030129}}</ref> The skeleton was structured so large areas of skin were stretched over solid plates of bone. This bone was composed of particularly dense fibers – so dense that exchange of oxygen through the skin was unlikely. Rather, the dense [[ossification]]s served to retain water inside the body as ''Laccognathus'' traveled on land between bodies of water.<ref name = "witzmann">{{cite journal|author=[[Florian Witzman|Witzman, F.]]|journal=[[Paleodiversity]]|volume=31|issue=5|pages=981–996|year=2011|publisher=The Society of Vertebrate Paleontology|title=A New Species of ''Laccognathus'' (Sarcopterygii, Porolepiformes) from the Late Devonian of Ellesmere Island, Nunavut, Canada}}</ref>
| [[Haplochromis erythromaculatus]] ||
|-
| rowspan=7 style=background:#FFD090 | <div align="center">[[Devoniano medio]]</div>
| [[Haplochromis estor]] ||
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Devoniano medio]] (397–385 Ma): [[Chondrichthyes|Cartilaginous fishes]], consisting of [[shark]]s, [[Batoidea|rays]] and [[chimaera]]s, appeared about 395 Ma.
|-
|-
| [[Haplochromis eutaenia]] ||
| [[File:Dipterus valenciennesi1.jpg|140px]]
|-
| ''[[Dipterus]]''
| [[Haplochromis exspectatus]] ||
| ''[[Dipterus]]'' ''(two wings)'' is an extinct genus of [[lungfish]] from 376–361 Ma. It was about {{convert|35|cm|in}} long, mostly ate invertebrates, and had lungs, not an air bladder. Like its ancestor ''[[Dipnorhynchus]]'' it had tooth-like plates on its palate instead of real teeth. However, unlike its modern relatives, in which the [[dorsal fin|dorsal]], [[caudal fin|caudal]], and [[anal fin]] are fused into one, its fins were still separated. Otherwise ''Dipterus'' closely resembled modern lungfish.{{sfn|The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|1999|p=45}}
|-
|-
| [[Haplochromis fischeri]] ||
| [[File:Early Shark.jpg|140px]]
|-
| ''[[Cladoselache]]''
| [[Haplochromis flavipinnis]] ||
| ''[[Cladoselache]]'' was the first abundant genus of primitive shark, appearing about 370 Ma.{{sfn|The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|1999|p=26}} It grew to {{convert|6|ft|m}} long, with anatomical features similar to modern [[mackerel shark]]s. It had a streamlined body almost entirely devoid of [[Fish scale|scale]]s, with five to seven [[gill slit]]s and a short, rounded snout that had a terminal mouth opening at the front of the skull.{{sfn|The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|1999|p=26}} It had a very weak jaw joint compared with modern-day sharks, but it compensated for that with very strong jaw-closing muscles. Its teeth were multi-cusped and smooth-edged, making them suitable for grasping, but not tearing or chewing. ''Cladoselache'' therefore probably seized prey by the tail and swallowed it whole.{{sfn|The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|1999|p=26}} It had powerful keels that extended onto the side of the tail stalk and a semi-lunate tail fin, with the superior lobe about the same size as the inferior. This combination helped with its speed and agility which was useful when trying to outswim its probable predator, the heavily armoured {{convert|10|m|ft}} long placoderm fish ''[[Dunkleosteus]]''.{{sfn|The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|1999|p=26}}
|-
|-
| [[Haplochromis flavus]] ||
| [[File:Coccosteus BW.jpg|140px]]
|-
| ''[[Coccosteus]]''
| [[Haplochromis fuscus]] ||
| ''[[Coccosteus]]'' ''(seed bone)'' is an extinct genus of [[arthrodire]] [[placoderm]]. The majority of fossils have been found in freshwater sediments, though they may have been able to enter saltwater. They grew up to {{convert|40|cm|in}} long. Like all other arthrodires, ''Coccosteus'' had a joint between the armour of the body and skull. In addition, it also had an internal joint between its neck [[vertebrae]] and the back of the skull, allowing for the mouth to be opened even wider. Along with the longer jaws, this allowed ''Coccosteus'' to feed on fairly large prey. As with all other arthrodires, ''Coccosteus'' had bony dental plates embedded in its jaws, forming a beak. The beak was kept sharp by having the edges of the dental plates grind away at each other.{{sfn|The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|1999|p=32}}
|-
|-
| [[Haplochromis fusiformis]] ||
| [[File:Bothriolepis canadensis 2.jpg|center|120px]]
|-
| ''[[Bothriolepis]]''
| [[Haplochromis gigas]] ||
|-
''[[Bothriolepis]]'' è stato il genere di [[placodermi]] più diffuso nei ritrovamenti fossili, con oltre 100 specie scoperte negli strati rocciosi del Devoniano medio in ogni continente.
| [[Haplochromis gigliolii]] ||
|-
| [[File:Pituriaspis doylei.jpg|center|80px]]
| [[Haplochromis gilberti]] ||
| [[Pituriaspida]]
|-
| [[Pituriaspida]] ''(hallucinogenic shield)'' is a class containing two bizarre species of armoured jawless fishes with tremendous nose-like [[rostrum (anatomy)|rostrum]]s. They lived in estuaries around 390 Ma. The [[paleontologist]] Gavin Young, named the class after the hallucinogenic drug [[pituri]], since he thought he might be hallucinating upon viewing the bizarre forms.{{sfn|Long|1996|p=}} The better studied species looked like a throwing-dart-like, with an elongate headshield and spear-like rostrum. The other species looked like a guitar pick with a tail, with a smaller and shorter rostrum and a more triangular headshield.
| [[Haplochromis goldschmidti]] ||
|-
| colspan="3" style="line-height:16px; background:#dddddd;"| <small>[[Late Devonian extinction]]: 375–360 Ma. A prolonged series of extinctions eliminated about 19% of all families, 50% of all genera{{r|Baez2006}} and 70% of all species. This extinction event lasted perhaps as long as 20 Ma, and there is evidence for a series of extinction pulses within this period.</small>
| [[Haplochromis gowersii]] ||
|-
| rowspan=5 style=background:#FFE0B0 | <div align="center">[[Devoniano superiore]]</div>
| [[Haplochromis gracilior]] ||
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Devoniano superiore]] (383–359 Ma):
|-
|-
| [[Haplochromis granti]] ||
| [[File:Dunkleosteus intermedius.jpg|140px]]
|-
| ''[[Dunkleosteus]]''
| [[Haplochromis graueri]] ||
| {{ external media
|-
| align =
| [[Haplochromis greenwoodi]] ||
| width = 210px
|-
| video1 = [http://animal.discovery.com/tv-shows/other/videos/animal-armageddon-dunkleosteus ''Dunkleosteus''] – ''Animal Planet''
| [[Haplochromis guiarti]] ||
}}
|-
''[[Dunkleosteus]]'' is a [[genus]] of [[arthrodire]] [[placoderm]]s which was extant from 380 to 360 Ma. It grew up to {{convert|10|m|ft}} long{{r|ScienceNews1999}}{{sfn|The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|1999|p=33}} and weighed up to 3.6 tonnes.{{r|SydneyMorningHerald2006}} It was a [[hypercarnivore|hypercarnivorous]] [[apex predator]]. Apart from its contemporary ''Titanichthys'' (below), no other placoderm rivalled it in size. Instead of teeth, ''Dunkleosteus'' had two pairs of sharp bony plates which formed a beak-like structure. Apart from [[megalodon]], it had the most powerful bite of any fish,{{r|phys.org2006}} generating bite forces in the same league as ''[[Tyrannosaurus|Tyrannosaurus rex]]'' and the modern [[crocodile]].{{r|LiveScience2006}}
| [[Haplochromis harpakteridion]] ||
|-
| [[File:Titanichthys10.jpg|140px]]
| [[Haplochromis heusinkveldi]] ||
| ''[[Titanichthys]]''
|-
| ''[[Titanichthys]]'' is a genus of giant, aberrant marine [[placoderm]] which lived in shallow seas. Many of the species approached ''[[Dunkleosteus]]'' in size and build. Unlike its relative, however, the various species of ''Titanichys'' had small, ineffective-looking mouth-plates that lacked a sharp cutting edge. It is assumed that ''Titanichthys'' was a [[filter feeder]] that used its capacious mouth to swallow or inhale schools of small, [[anchovy]]-like fish, or possibly [[krill]]-like [[zooplankton]], and that the mouth-plates retained the prey while allowing the water to escape as it closed its mouth.
| [[Haplochromis hiatus]] ||
|-
| [[File:Materpiscis02.JPG|140px]]
| [[Haplochromis howesi]] ||
| ''[[Materpiscis]]''
|-
| {{ external media
| [[Haplochromis humilior]] ||
| align = center
|-
| width = 180px
| [[Haplochromis humilis]] ||
| video1 = [http://www.nature.com/nature/videoarchive/themotherfish/ The mother fish] – ''Nature''
|-
}}
| [[Haplochromis igneopinnis]] ||
''[[Materpiscis]]'' ''(mother fish)'' is a [[genus]] of [[ptyctodontid]] [[Placodermi|placoderm]] from about 380 Ma. Known from only one specimen, it is unique in having an unborn [[embryo]] present inside, and with remarkable preservation of a mineralised placental feeding structure ([[umbilical cord]]). This makes ''Materpiscis'' the first known vertebrate to show [[vivipary|viviparity]], or giving birth to live young.{{r|Long2008}} The specimen was named ''Materpiscis attenboroughi'' in honour of [[David Attenborough]].{{r|BBC2008}}
|-
|-
| [[Haplochromis insidiae]] ||
|-
| [[Haplochromis irisRhizodont]] || s
| [[Rhizodont]]s were an order of lobe-finned fish which survived to the end of the Carboniferous, 377–310 Ma. They reached huge sizes. The largest known species, ''[[Rhizodus|Rhizodus hibberti]]'' grew up to 7 metres in length, making it the largest freshwater fish known.
|-
|}
| [[Haplochromis ishmaeli]] ||
</div>
|-
{{clear}}
| [[Haplochromis kamiranzovu]] ||
 
|-
===Dai pesci ai tetrapodi===
| [[Haplochromis katavi]] ||
{{further|Tetrapod#Evolution|List of transitional fossils#Fish to Tetrapods}}
|-
 
| [[Haplochromis katonga]] ||
[[File:Fishapod evolution.jpg|thumb|300px|right|A cladogram of the evolution of tetrapods showing some of the best-known transitional fossils. It starts with [[Eusthenopteron]] at the bottom, indisputably still a fish, through [[Panderichthys]], [[Tiktaalik]], [[Acanthostega]] and [[Ichthyostega]] to [[Pederpes]] at the top, indisputably a tetrapod
|-
{| class="wikitable collapsible collapsed" style="text-align: left;"
| [[Haplochromis katunzii]] ||
|-
! width=290px | From fins to limbs
| [[Haplochromis kunjunjui]] ||
|-
|{{multiple image
| [[Haplochromis labiatus]] ||
| align = left
|-
| direction = vertical
| [[Haplochromis labriformis]] ||
| width = 280
|-
| header =
| [[Haplochromis lacrimosus]] ||
| header_align = <!-- left/right/center -->
|-
| header_background =
| [[Haplochromis laparogramma]] ||
| footer =
|-
| footer_align = <!-- left/right/center -->
| [[Haplochromis latifasciatus]] ||
| footer_background =
|-
| background color =
| [[Haplochromis limax]] ||
| image1 = Crossopterygii tetrapod hips.JPG
|-
| alt1 =
| [[Haplochromis lividus]] ||
| caption1 = Illustration showing shows how much the hindlimb attachments in lobe-finned fishes need to change in transitioning from lobe-finned fishes (A) to early tetrapods (B) if the fish was to become a terrestrial animal.
|-
| image2 = Fishapods and tetrapods.JPG
| [[Haplochromis loati]] ||
| alt2 =
|-
| caption2 = Comparison between the fins of lobe-finned fishes and the legs of early tetrapods: 1. ''[[Tiktaalik]]'' 2. ''[[Panderichthys]]'' 3. ''[[Eusthenopteron]]'' 4. ''[[Acanthostega]]'' 5. ''[[Ichthyostega]]'' ( hindleg)
| [[Haplochromis longirostris]] ||
}}
|-
|}
| [[Haplochromis luteus]] ||
]]
|-
 
| [[Haplochromis macconneli]] ||
The first [[tetrapods]] are four-legged, air-breathing, terrestrial animals from which the land vertebrates descended, including humans. They evolved from [[lobe-finned fish]], appearing in coastal water in the middle Devonian, and giving rise to the first [[amphibians]].{{r|Niedźwiedzki2010}}
|-
 
| [[Haplochromis macrocephalus]] ||
The group of lobe-finned fishes that were the ancestors of the tetrapod are grouped together as the [[Rhipidistia]],{{r|Gordon2004}} and the first tetrapods evolved from these fish over the relatively short timespan 385–360 Ma. The early tetrapod groups themselves are grouped as [[Labyrinthodontia]]. They retained aquatic, fry-like [[tadpole]]s, a system still seen in [[Lissamphibia|modern amphibians]]. From the 1950s to the early 1980s it was thought that tetrapods evolved from fish that had already acquired the ability to crawl on land, possibly in order to go from a pool that was drying out to one that was deeper. However, in 1987, nearly complete fossils of ''[[Acanthostega]]'' from about {{ma|363|Ma|Devonian}} showed that this Late Devonian [[Transitional fossil|transitional]] animal had legs and both lungs and gills, but could never have survived on land: its limbs and its wrist and ankle joints were too weak to bear its weight; its ribs were too short to prevent its lungs from being squeezed flat by its weight; its fish-like tail fin would have been damaged by dragging on the ground. The current hypothesis is that ''Acanthostega'', which was about {{convert|1|m|ft}} long, was a wholly aquatic predator that hunted in shallow water<!-- , using its limbs to hold on to vegetation while it lay in ambush -->. Its skeleton differed from that of most fish, in ways that enabled it to raise its head to breathe air while its body remained submerged, including: its jaws show modifications that would have enabled it to gulp air; the bones at the back of its skull are locked together, providing strong attachment points for muscles that raised its head; the head is not joined to the [[shoulder girdle]] and it has a distinct neck.{{r|Clack2005}}
|-
 
| [[Haplochromis macrognathus]] ||
[[File:Ancestors of the land vertebrates cropped.png|thumb|250px|left|Until the 1980s early transitional lobe-finned fishes, such as the ''[[Eusthenopteron]]'' shown here, were depicted as emerging onto land. Paleontologists now widely agree this didn't happen, and they were strictly aquatic.{{r|Laurin2007}}
|-
{{ external media
| [[Haplochromis macrops]] ||
| align = center
|-
| width = 240px
| [[Haplochromis macropsoides]] ||
| video1 = [http://www.youtube.com/watch?v=k-5oQlnXSTM Tetrapod Evolution] ''Animal Planet''<br />[http://www.youtube.com/watch?v=k-5oQlnXSTM 1] [http://www.youtube.com/watch?v=B3iFADplW6U 2] [http://www.youtube.com/watch?v=8HCTFe_XZFQ 3] [http://www.youtube.com/watch?v=JKRTrC1B1PI 4] [http://www.youtube.com/watch?v=khUw_OGRcBs 5]
|-
| video2 = [http://www.youtube.com/watch?v=ga6n2vnM3Nc Evolution fish with fingers Transitional fossils] – ''YouTube''
| [[Haplochromis maculipinna]] ||
}}
|-
]]
| [[Haplochromis mahagiensis]] ||
 
|-
The Devonian proliferation of land plants may help to explain why air-breathing would have been an advantage: leaves falling into streams and rivers would have encouraged the growth of aquatic vegetation; this would have attracted grazing invertebrates and small fish that preyed on them; they would have been attractive prey but the environment was unsuitable for the big marine predatory fish; air-breathing would have been necessary because these waters would have been short of oxygen, since warm water holds less dissolved oxygen than cooler marine water and since the decomposition of vegetation would have used some of the oxygen.{{r|Clack2005}}
| [[Haplochromis maisomei]] ||
 
|-
There are three major hypotheses as to how tetrapods evolved their stubby fins (proto-limbs). The traditional explanation is the "shrinking waterhole hypothesis" or "desert hypothesis" posited by the American paleontologist [[Alfred Romer]]. He believed limbs and lungs may have evolved from the necessity of having to find new bodies of water as old waterholes dried up.{{r|science22011}}
| [[Haplochromis malacophagus]] ||
 
|-
The second hypothesis is the "inter-tidal hypothesis" put forward in 2010 by a team of Polish paleontologists lead by Grzegorz Niedźwiedzki. They argued that sarcopterygians may have first emerged unto land from [[intertidal zone]]s rather than inland bodies of water. Their hypothesis is based on the discovery of the 395 million-year-old Zachełmie tracks in [[Zachełmie, Świętokrzyskie Voivodeship|Zachełmie]], Poland, the oldest ever discovered fossil evidence of tetrapods.{{r|Niedźwiedzki2010}}{{r|newscientist2010}}
| [[Haplochromis mandibularis]] ||
 
|-
The third hypothesis, the "woodland hypothesis", was proposed by the American paleontologist Gregory J. Retallack in 2011. He argues that limbs may have developed in shallow bodies of water in woodlands as a means of navigating in environments filled with roots and vegetation. He based his conclusions on the evidence that transitional tetrapod fossils are consistently found in habitats that were formerly humid and wooded [[floodplain]]s.{{r|Retallack2011}}
| [[Haplochromis martini]] ||
 
|-
Research by [[Jennifer A. Clack]] and her colleagues showed that the very earliest tetrapods, animals similar to ''[[Acanthostega]]'', were wholly aquatic and quite unsuited to life on land. This is in contrast to the earlier view that fish had first invaded the land — either in search of prey (like modern [[mudskipper]]s) or to find water when the pond they lived in dried out — and later evolved legs, lungs, etc.
| [[Haplochromis maxillaris]] ||
 
|-
Two ideas about the [[Homology (biology)|homology]] of arms, hands and digits have existed in the past 130 years. First that digits are unique to tetrapods<ref>Holmgren N. (1933). On the origin of the tetrapod limb. Acta Zoologica 14, 185–295.</ref><ref>{{cite journal | author = Vorobyeva EI | year = 1992 | title = The role of development and function in formation of tetrapod like pectoral fins | url = | journal = Zh. Obshch. Biol. | volume = 53 | issue = | pages = 149–158 }}</ref> and second that antecedents were present in the fins of early [[sarcopterygian]] fish.<ref>{{cite journal | author = Watson DMS | year = 1913 | title = On the primitive tetrapod limb | url = | journal = Anat. Anzeiger | volume = 44 | issue = | pages = 24–27 }}</ref> Until recently it was believed that "genetic and fossil data support the hypothesis that digits are evolutionary novelties".<ref>{{cite journal | author = Shubin N, Tabin C, Carroll S | year = 1997 | title = Fossils, genes and the evolution of animal limbs | url = | journal = Nature | volume = 388 | issue = 6643| pages = 639–48 | doi = 10.1038/41710 | pmid = 9262397 }}</ref><sup>p.&nbsp;640.</sup> However new research that created a three-dimensional reconstruction of [[Panderichthys]], a coastal fish from the [[Devonian]] period 385 million years ago, shows that these animals already had many of the homologous bones present in the forelimbs of limbed vertebrates.<ref name="Boisvert"/> For example, they had [[Radius (bone)|radial]] bones similar to rudimentary fingers but positioned in the arm-like base of their fins.<ref name="Boisvert"/> Thus there was in the evolution of [[tetrapod]]s a shift such that the outermost part of the fins were lost and came to be replaced by early digits. This change is consistent with additional evidence from the study of [[actinopterygian]]s, [[shark]]s and [[lungfish]] that the digits of tetrapods arose from pre-existing distal radials present in more primitive fish.<ref name="Boisvert"/><ref>{{cite web
| [[Haplochromis mbipi]] ||
| url = http://news.nationalgeographic.com/news/2008/09/080924-fish-fingers.html?source=rss
|-
| title = Ancient Fish Had Primitive Fingers, Toes
| [[Haplochromis megalops]] ||
| first = Ker | last = Than
|-
| publisher = National Geographic News
| [[Haplochromis melanopterus]] ||
| date = September 24, 2008
|-
| accessdate =
| [[Haplochromis melanopus]] ||
}}</ref> Controversy still exists since [[Tiktaalik]], a vertebrate often considered to be the [[Transitional fossil|missing link]] between fishes and land-living animals, had stubby leg-like limbs that lacked the finger-like radial bones found in the Panderichthys. The researchers of the paper commented that it "is difficult to say whether this character distribution implies that Tiktaalik is [[autapomorph]]ic, that Panderichthys and tetrapods are convergent, or that Panderichthys is closer to tetrapods than Tiktaalik. At any rate, it demonstrates that the fish–tetrapod transition was accompanied by significant character incongruence in functionally important structures.".<ref name="Boisvert">{{cite journal | author = Boisvert CA, Mark-Kurik E, Ahlberg PE | year = 2008 | title = The pectoral fin of Panderichthys and the origin of digits | url = http://uu.diva-portal.org/smash/get/diva2:221262/FULLTEXT01 | journal = Nature | volume = 456 | issue = 7222| pages = 636–8 | pmid = 18806778 | doi=10.1038/nature07339|bibcode = 2008Natur.456..636B }}</ref><sup>p.&nbsp;638.</sup>
|-
 
| [[Haplochromis melichrous]] ||
From the end of the Devonian to the Mid [[Carboniferous]] a 30 million year gap occurs in the fossil record. This gap, called [[Romer's gap]], is marked by the absence of ancestral tetrapod fossils and fossils of other vertebrates that look well-adapted for life on land.{{r|Ahlberg1994}}
|-
 
| [[Haplochromis mentatus]] ||
{{clear}}
|-
<div>
| [[Haplochromis mento]] ||
:::{| class="wikitable"
|-
|-
| [[Haplochromis michaeli]] ||
! colspan=4 | Transition from lobe-finned fishes to tetrapods
|-
|-
| [[Haplochromis microchrysomelas]] ||
| [[File:Eusthenopteron model.jpg|140px]]<small>~385 milioni di anni fa</small>
|-
| ''[[Eusthenopteron]]''
| [[Haplochromis multiocellatus]] ||
| {{ external media
|-
| align =
| [[Haplochromis mylergates]] ||
| width = 220px
|-
| video1 = [http://animal.discovery.com/tv-shows/other/videos/animal-armageddon-eusthenopteron ''Eusthenopteron''] – ''Animal Planet''
| [[Haplochromis mylodon]] ||
}}
|-
Genus of [[extinction|extinct]] [[lobe-finned fish]]es which has attained an iconic status from its close relationships to [[tetrapod]]s. Early depictions of this animal show it emerging onto land, however paleontologists now widely agree that it was a strictly aquatic animal.{{r|Laurin2007}} The genus ''Eusthenopteron'' is known from several species that lived during the Late Devonian period, about 385 Ma. It was the object of intense study from the 1940s to the 1990s by the [[paleoichthyologist]] [[Erik Jarvik]].{{r|Jarvik2008}}
| [[Haplochromis nanoserranus]] ||
|-
| [[File:Gogonasus BW.jpg|140px]]
| [[Haplochromis nigrescens]] ||
| ''[[Gogonasus]]''
|-
| ''[[Gogonasus]] (snout from Gogo)'' was a lobe-finned fish known from 3-dimensionally preserved 380 million-year-old fossils found in the [[Gogo Formation]]. It was a small fish reaching 30–40&nbsp;cm (1&nbsp;ft) in length.<ref>http://dinosaurs.about.com/od/tetrapodsandamphibians/p/gogonasus.htm</ref> Its skeleton shows several tetrapod-like features. They included the structure of its [[middle ear]], and its fins show the precursors of the forearm bones, the [[radius]] and [[ulna]]. Researchers believe it used its forearm-like fins to dart out of the reef to catch prey. ''Gogonasus'' was first described in 1985 by [[John A. Long]]. For almost 100 years ''[[Eusthenopteron]]'' has been the role model for demonstrating stages in the evolution of lobe-finned fishes to tetrapods. ''Gogonasus'' now replaces ''Eusthenopteron'' in being a better preserved representative without any ambiguity in interpreting its anatomy.
| [[Haplochromis nigricans]] ||
|-
| [[File:Panderichthys BW.jpg|140px]]<br /><small>~385 milioni di anni fa</small>
| [[Haplochromis nigripinnis]] ||
| ''[[Panderichthys]]''
|-
| Adapted to muddy shallows, and capable of some kind of shallow water or terrestrial body flexion locomotion. Had the ability to prop itself up.<ref>Nature: [http://www.nature.com/nature/journal/v438/n7071/edsumm/e051222-13.html The pelvic fin and girdle of ''Panderichthys'' and the origin of tetrapod locomotion]</ref><br /> They had large tetrapod-like heads, and are thought to be the most crownward stem fish-tetrapod with paired fins.
| [[Haplochromis nigroides]] ||
|-
| [[File:Tiktaalik BW.jpg|140px]]<br /><small>~375 milioni di anni fa</small>
| [[Haplochromis niloticus]] ||
| ''[[Tiktaalik]]''
|-
| A fish with limb-like fins that could take it onto land.<ref name=Nature>{{cite journal | journal = [[Nature (journal)|Nature]] | volume = 440 | pages = 757–763 | date = 6 April 2006 | doi = 10.1038/nature04639 | title = A Devonian tetrapod-like fish and the evolution of the tetrapod body plan | url = http://www.nature.com/nature/journal/v440/n7085/abs/nature04639.html | author = Edward B. Daeschler, Neil H. Shubin and Farish A. Jenkins, Jr | pmid = 16598249 | issue = 7085}}</ref> It is an example from several lines of ancient sarcopterygian fish developing adaptations to the oxygen-poor shallow-water habitats of its time, which led to the evolution of tetrapods.<ref name="scientificamerican">[[Jennifer A. Clack]], ''Scientific American'', [http://www.scientificamerican.com/article.cfm?id=getting-a-leg-up-on-land ''Getting a Leg Up on Land''] Nov. 21, 2005.</ref> Paleontologists suggest that it is representative of the [[transitional fossil|transition]] between non-tetrapod vertebrates (fish) such as ''[[Panderichthys]]'', known from fossils 380 million years old, and early tetrapods such as ''[[Acanthostega]]'' and ''[[Ichthyostega]]'', known from fossils about 365 million years old. Its mixture of primitive fish and derived tetrapod characteristics led one of its discoverers, [[Neil Shubin]], to characterize ''Tiktaalik'' as a "[[Tetrapodomorpha|fishapod]]".<ref>John Noble Wilford, ''The New York Times'', [http://www.nytimes.com/2006/04/05/science/05cnd-fossil.html?hp&ex=1144296000&en=fe3427d67e965e46&ei=5094&partner=homepage ''Scientists Call Fish Fossil the Missing Link''], Apr. 5, 2006.</ref><ref name="Shubin 2008">{{cite book|last=Shubin|first=Neil|title=Your Inner Fish|publisher=Pantheon|year=2008|isbn=978-0-375-42447-2}}</ref>
| [[Haplochromis nubilus]] ||
|-
| [[File:Acanthostega BW.jpg|140px]]<br /><small>365 milioni di anni fa</small>
| [[Haplochromis nuchisquamulatus]] ||
| ''[[Acanthostega]]''
|-
| A fish-like early [[Labyrinthodontia|labyrinthodont]] which occupied weed-filled swamps and changed views about the early evolution of tetrapods.{{r|Clack2005}} [[Polydactyly in early tetrapods|It had eight digits]] on each hand (the number of digits on the feet is unclear) linked by webbing, it lacked wrists, and was generally poorly adapted to come onto land.<ref>"[http://www.devoniantimes.org/Order/re-acanthostega.html Acanthostega gunneri]," ''Devonian Times''.</ref> Sunsequent discoveries revealed earlier transitional forms between ''Acanthostega'' and completely fish-like animals.{{r|Daeschler2006}}
| [[Haplochromis nyanze]] ||
 
|-
|-
| [[Haplochromis nyererei]] ||
| [[File:Ichthyostega BW.jpg|140px]]<br /><small>374–359 milioni di anni fa</small>
|-
| ''[[Ichthyostega]]''
| [[Haplochromis obesus]] ||
| {{ external media
|-
| align =
| [[Haplochromis obliquidens]] ||
| width = 225px
|-
| video1 = [http://animal.discovery.com/tv-shows/other/videos/animal-armageddon-ichthyostega.htm ''Ichthyostega''] – ''Animal Planet''
| [[Haplochromis obtusidens]] ||
| video2 = [http://www.youtube.com/watch?v=qY6cHGVG8-c Pierce et al Vertebral Architecture<br />1] [http://www.youtube.com/watch?v=8g0_vCd71bw 2] [http://www.youtube.com/watch?v=0IPmHJ_3nug 3] [http://www.youtube.com/watch?v=lzEzAC5DsUc 4] [http://www.youtube.com/watch?v=TWst4N70Wr4 5] – ''YouTube''}}
|-
Until finds of other early tetrapods and closely related fishes in the late 20th century, ''[[Ichthyostega]]'' stood alone as the transitional fossil between fish and tetrapods, combining a fishlike tail and gills with an amphibian skull and limbs. It possessed lungs and limbs with seven digits that helped it navigate through shallow water in weed-filled swamps.
| [[Haplochromis occultidens]] ||
|-
| [[File:Pederpes22small.jpg|140px]]<br /><small>359–345 milioni di anni fa</small>
| [[Haplochromis oligolepis]] ||
| ''[[Pederpes]]''
|-
| ''[[Pederpes]]'' is the earliest known fully terrestrial tetrapod. It is included here to complete the transition of lobe-finned fishes to tetrapods, even though ''Pederpes'' is no longer a fish.
| [[Haplochromis olivaceus]] ||
|- }
</div>
| [[Haplochromis omnicaeruleus]] ||
{{clear}}
|-
 
| [[Haplochromis oregosoma]] ||
By the late Devonian, land plants had stabilized freshwater habitats, allowing the first [[wetland]] [[ecosystem]]s to develop, with increasingly complex [[Food chain|food web]]s that afforded new opportunities. Freshwater habitats were not the only places to find water filled with organic matter and choked with plants with dense vegetation near the water's edge. Swampy habitats like shallow wetlands, coastal lagoons and large brackish river deltas also existed at this time, and there is much to suggest that this is the kind of environment in which the tetrapods evolved. Early fossil tetrapods have been found in marine sediments, and because fossils of primitive tetrapods in general are found scattered all around the world, they must have spread by following the coastal lines — they could not have lived in freshwater only.
|-
* [http://www.scientificamerican.com/article.cfm?id=fossil-illuminates-evolut Fossil Illuminates Evolution of Limbs from Fins] ''Scientific American'', 2 2 April 2004.
| [[Haplochromis orthostoma]] ||
 
|-
===Post-Devoniano===
| [[Haplochromis pachycephalus]] ||
L'[[Mesozoico|Era Mesozoica]] inizia circa 250 milioni di anni fa sulla scia dell'[[Estinzione di massa del Permiano-Triassico]], la più grande [[estinzione]] della storia della [[Terra]], e finisce circa 65 milioni di anni fa, con l'[[Estinzione di massa del Cretaceo-Paleocene]], un'altra estinzione massiva che uccise i [[dinosauri]] e moltre specie di piante e animali. É spesso definita come l'''età dei rettili'', perché i rettili sono stati i vertebrati dominanti in questa era. Durante il Mesozoico inoltre il maxicontinente [[Pangea]] si divide gradualmente in continenti più piccoli. Il clima alterna periodi molto caldi a periodi più freddi, ma nel complesso la Terra era più calda di quanto non lo sia oggi.
|-
 
| [[Haplochromis pallidus]] ||
{| class="wikitable"
|-
|-
| [[Haplochromis paludinosus]] ||
! rowspan=5 style=background:#888584 | [[Carbonifero]]
|-
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Carboniferous]] (359–299 Ma): [[Shark]]s underwent a major [[evolutionary radiation]] during the [[Carboniferous]].<ref name=goldsharks/> It is believed that this evolutionary radiation occurred because the decline of the [[Placodermi|placoderms]] at the end of the Devonian period caused many [[Niche (ecology)|environmental niches]] to become unoccupied and allowed new organisms to evolve and fill these niches.<ref name=goldsharks/>
| [[Haplochromis pancitrinus]] ||
|-
| width="140px" | [[File:Kansas sea2DB.jpg|140px]]<br /><small>''[[Cretoxyrhina]]'' and ''[[Squalicorax]]'' circle a dead ''[[Claosaurus]]''</small>
| [[Haplochromis pappenheimi]] ||
| align="center" |''[[Cretoxyrhina]]''<br /><br /><br />''[[Squalicorax]]''
|-
| The first 15 million years of the Carboniferous has very few terrestrial fossils. This gap in the fossil record, is called [[Romer's gap]] after the American palaentologist [[Alfred Romer]]. While it has long been debated whether the gap is a result of fossilisation or relates to an actual event, recent work indicates the gap period saw a drop in atmospheric oxygen levels, indicating some sort of ecological collapse.<ref name=Ward>{{cite journal | author = Ward P. ''et al.'' | year = 2006 | title = Confirmation of Romer's Gap is a low oxygen interval constraining the timing of initial arthropod and vertebrate terrestrialization | url = | journal = [[Proceedings of the National Academy of Science]] | volume = 103 | issue = 45| pages = 16818–16822 }}</ref> The gap saw the demise of the [[Devonian]] fish-like [[ichthyostegalia]]n labyrinthodonts, and the rise of the more advanced [[Temnospondyli|temnospondyl]] and [[reptiliomorpha]]n amphibians that so typify the Carboniferous terrestrial vertebrate fauna.
| [[Haplochromis paradoxus]] ||
 
|-
The [[Carboniferous]] seas were inhabited by many fish, mainly [[Elasmobranch]]s (sharks and their relatives). These included some, like ''[[Psammodus]]'', with crushing pavement-like teeth adapted for grinding the shells of brachiopods, crustaceans, and other marine organisms. Other sharks had piercing teeth, such as the [[Symmoriida]]; some, the [[Petalodontiformes|petalodonts]], had peculiar cycloid cutting teeth. Most of the sharks were marine, but the [[Xenacanthida]] invaded fresh waters of the coal swamps. Among the [[Osteichthyes|bony fish]], the [[Palaeonisciformes]] found in coastal waters also appear to have migrated to rivers. [[Sarcopterygii|Sarcopterygia]]n fish were also prominent, and one group, the [[Rhizodont]]s, reached very large size.
| [[Haplochromis paraguiarti]] ||
 
|-
Most species of Carboniferous marine fish have been described largely from teeth, fin spines and dermal ossicles, with smaller freshwater fish preserved whole. Freshwater fish were abundant, and include the genera ''[[Ctenodus]]'', ''[[Uronemus]]'', ''[[Acanthodes]]'', ''[[Cheirodus]]'', and ''[[Gyracanthus]]''.
| [[Haplochromis paraplagiostoma]] ||
|-
| [[File:Stethacanthus1DB.jpg|140px]]
| [[Haplochromis paropius]] ||
| ''[[Stethacanthidae]]''
|-
| [[File:Steth pair1.jpg|140px|right]]
| [[Haplochromis parorthostoma]] ||
As a result of the evolutionary radiation carboniferous sharks assumed a wide variety of bizarre shapes including sharks belonging to the family [[Stethacanthidae]] which possessed a flat brush-like dorsal fin with a patch of [[denticles]] on its top.<ref name=goldsharks/> ''[[Stethacanthus]]''' unusual fin may have been used in mating rituals.<ref name=goldsharks>{{cite web |url=http://www.elasmo-research.org/education/evolution/golden_age.htm |title=A Golden Age of Sharks |accessdate=2008-06-23 |work=Biology of Sharks and Rays |author=R. Aidan Martin}}</ref> Apart from the fins, Stethacanthidae resembled ''Falcatus'' (below).
|-
|-
| [[Haplochromis parvidens]] ||
| [[File:Falcatus.jpg|140px]]
|-
| ''[[Falcatus]]''
| [[Haplochromis paucidens]] ||
| ''[[Falcatus]]'' è un genere di piccoli cladodonti dai denti appuntiti e squaliformi, che vissero da 335 a 318 milioni di anni fa. Di piccole dimensioni, raggiungevano i 25-30 cm di lunghezza. <ref>Pesci fossili [http://www.sju.edu/research/bear_gulch/pages_fish_species/Falcatus_falcatus.php di Bear Gulch 2005 da Richard Lund e Eileen Grogan] </ref> Sono caratterizzati dalla particolare forma della pinna dorsale, che presenta i raggi incurvati verso la testa.
|-
|-
| [[Haplochromis pellegrini]] ||
| [[File:Orodus sp1DB.jpg|140px]]
|-
| ''[[Orodus]]''
| [[Haplochromis percoides]] ||
| ''[[Orodus]]'' è un altro genere di [[squalo]] del Carbonifero, appartenenti alla famiglia [[Orodontidae]], vissuto da 303 a 295 milioni di anni fa. Raggiungeva una lunghezza di 2 metri.
|-
|-
| [[Haplochromis perrieri]] ||
! rowspan=3 style=background:#ED4C35 | [[Permiano]]
|-
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Permiano]] (298–252 Ma):
| [[Haplochromis petronius]] ||
|-
| [[File:Acanthodes BW.jpg|140px]]
| [[Haplochromis pharyngalis]] ||
| ''[[Acanthodes]]''
|-
| ''[[Acanthodes]]'' are an extinct genus of [[Acanthodii|spiny shark]].<ref>{{cite journal|year=1880|title=Proceedings of the Royal Physical Society of Edinburgh|volume= V.|pages=p115|url=http://books.google.com/?id=69kKAAAAIAAJ&pg=PA115&dq=Acanthodes+extinct|doi=10.1111/|author1=Edinburgh, Royal Physical Society of}}</ref> It had [[gill]]s but no [[teeth]],<ref name="Nicholson">{{cite book|last=Nicholson|first=Henry Alleyne|coauthors=Richard Lydekker|title=A Manual of Palaeontology|year=1889|url=http://books.google.com/?id=MoAXAAAAYAAJ&pg=PA966&dq=Acanthodes+pectoral+fins|page=966}}</ref> and was presumably a [[filter feeder]].<ref name=EoDP/> ''Acanthodes'' had only two skull bones and were covered in cubical scales. Each paired [[pectoral fin|pectoral]] and [[pelvic fin]]s had one spine, as did the single [[anal fin|anal]] and [[dorsal fin]]s, giving it a total of six spines, less than half that of many other spiny sharks.<ref name=EoDP>{{cite book |editor=Palmer, D.|year=1999 |title= The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|publisher= Marshall Editions|___location=London|pages= 30–31|isbn= 1-84028-152-9}}</ref> Acanthodians share qualities of both bony fish (osteichthyes) and cartilaginous fish (chondrichthyes), and it has been suggested that they may have been stem chondrichthyans and stem gnathostomes.<ref>Article on Acanthodes as ancestor of Man, http://www.sci-news.com/paleontology/article00396.html, accessed 15 June 2012</ref><ref>Journal article on Acanthodes, http://www.nature.com/nature/journal/v486/n7402/full/nature11080.html, accessed 15 June 2012</ref>
| [[Haplochromis pharyngomylus]] ||
|-
| colspan="3" style="line-height:16px; background:#dddddd;"| <small>The Permian ended with the most extensive [[extinction event]] recorded in [[paleontology]]: the [[Permian-Triassic extinction event]]. 90% to 95% of marine species became [[Extinction|extinct]], as well as 70% of all land organisms. It is also the only known mass extinction of insects.<ref>http://geology.about.com/od/extinction/a/aa_permotrias.htm</ref><ref>http://www.kgs.ku.edu/Extension/fossils/massExtinct.html</ref> Recovery from the Permian-Triassic extinction event was protracted; land ecosystems took 30M years to recover,<ref name="SahneyBenton2008RecoveryFromProfoundExtinction">{{cite journal | url=http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2596898/?tool | author=Sahney, S. and Benton, M.J. | year=2008 | title=Recovery from the most profound mass extinction of all time | journal=Proceedings of the Royal Society: Biological | doi=10.1098/rspb.2007.1370 | volume = 275 | pages = 759–65|format=PDF | pmid=18198148 | issue=1636 | pmc=2596898}}</ref> and marine ecosystems took even longer.{{r|Baez2006}}</small>
| [[Haplochromis phytophagus]] ||
|-
! rowspan=5 style="background:#228b22" | [[Triassico|<span style="color:white;">Triassico</span>]]
| [[Haplochromis piceatus]] ||
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Triassico]] (252–201 Ma): The fish fauna of the Triassic was remarkably uniform, reflecting the fact that very few families survived the Permian extinction. A considerable radiation of ''ray-finned fishes]]'' occurred during the Triassic, laying the foundation for many modern fishes.<ref>[http://www.seaworld.org/animal-info/info-books/bony-fish/scientific-classification.htm Bony fishes] ''[[SeaWorld]]''. Retrieved 2 February 2013.</ref> ''See [[:Category:Triassic fish]].''
|-
|-
| [[Haplochromis pitmani]] ||
| [[File:Perleidus.jpg|140px]]
|-
| ''[[Perleidus]]''
| [[Haplochromis placodus]] ||
| ''[[Perleidus]]'' was a [[ray-finned fish]] from the Early Triassic. About {{convert|15|cm|in}} in length, it was a freshwater predatory fish with jaws that hung vertically under the braincase, allowing them to open wide. ''Perleidus'' had highly flexible [[dorsal fin|dorsal]] and [[anal fin]]s, with a reduced number of fin rays, which would have made the fish more agile in the water.<ref name=EoDP>{{cite book |editor=Palmer, D.|year=1999 |title= The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|publisher= Marshall Editions|___location=London|page= 36|isbn= 1-84028-152-9}}</ref>
|-
|-
| [[Haplochromis plagiodon]] ||
| [[File:Protosphyraena2DB.jpg|140px]]
|-
| [[Pachycormiformes]]
| [[Haplochromis plagiostoma]] ||
| [[File:Hypsocormus2.jpg|140px|right]]
|-
[[Pachycormiformes]] are an extinct order of ray-finned fish which were extant from the Middle Triassic to the [[Cretaceous–Paleogene extinction event|K-Pg extinction]] (below). They were characterized by serrated pectoral fins, reduced pelvic fins and a bony rostrum. Their relations with other fish are unclear.
| [[Haplochromis plutonius]] ||
|-
| [[File:Pholidophorus NT.jpg|140px]]
| [[Haplochromis prodromus]] ||
| ''[[Pholidophorus]]''
|-
| ''[[Pholidophorus]]'' was an [[extinct]] genus of [[teleost]], around {{convert|40|cm|in}} long, from about 240–140 Ma. Although not closely related to the modern [[herring]], it was somewhat like them. It had a single [[dorsal fin]], a symmetrical tail, and an [[anal fin]] placed towards the rear of the body. It had large eyes and was probably a fast swimming predator, hunting [[plankton]]ic [[crustacean]]s and smaller fish.{{sfn|The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|1999|p=38–39}} A very early [[teleost]], ''Pholidophoris'' had many primitive characteristics such as [[ganoid scale]]s and a [[Vertebral column|spine]] that was partially composed of [[cartilage]], rather than [[bone]].{{sfn|The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|1999|p=38–39}}
| [[Haplochromis prognathus]] ||
|-
| colspan="3" style="line-height:16px; background:#dddddd;"| <small>The Triassic ended with the [[Triassic–Jurassic extinction event]]. About 23% of all families, 48% of all genera (20% of marine families and 55% of marine genera) and 70% to 75% of all species went extinct.<ref name="ucr">{{cite web|url=http://math.ucr.edu/home/baez/extinction |title=extinction |publisher=Math.ucr.edu |date= |accessdate=2008-11-09}}</ref> Non-dinosaurian archosaurs continued to dominate aquatic environments, while [[Diapsid#Taxonomy|non-archosaurian diapsids]] continued to dominate marine environments.<ref name="ucr" /></small>
| [[Haplochromis pseudopellegrini]] ||
|-
! rowspan=3 style="background:#3cb371" | [[Giurassico]]
| [[Haplochromis ptistes]] ||
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Giurassico]] (199–145 milioni di anni fa):
|-
|-
| [[Haplochromis pundamilia]] ||
| [[File:Leedsichthys problematicus.jpg|140px]]
|-
| ''[[Leedsichthys]]''
| [[Haplochromis pyrrhocephalus]] ||
| During the [[Jurassic period]], the primary vertebrates living in the seas were [[fish]] and marine [[reptile]]s. The latter include [[ichthyosaur]]s who were at the peak of their diversity, [[plesiosauria|plesiosaurs]], [[pliosaur]]s, and marine [[crocodilia|crocodiles]] of the families [[Teleosauridae]] and [[Metriorhynchidae]].<ref>Motani, R. (2000), Rulers of the Jurassic Seas, Scientific American vol.283, no. 6</ref> Numerous [[turtle]]s could be found in lakes and rivers.<ref name=nws>{{citation
|-
|journal=Naturwissenschaften: The Science of Nature
| [[Haplochromis pyrrhopteryx]] ||
|title=An enormous Jurassic turtle bone bed from the Turpan Basin of Xinjiang, China
|-
|first=Oliver|last= Wings
| [[Haplochromis retrodens]] ||
|first2=Márton|last2= Rabi
|-
|first3=Jörg W.|last3= Schneider
| [[Haplochromis riponianus]] ||
|first4=Leonie|last4= Schwermann
|-
|first5=Ge |last5=Sun
| [[Haplochromis rubescens]] ||
|first6=Chang-Fu|last5= Zhou
|-
|first7=Walter G.|last7= Joyce
| [[Haplochromis rubripinnis]] ||
|volume=114|year=2012
|-
|url=http://www.springerlink.com/content/l55q277254344462/
|doi=10.1007/s00114-012-0974-5}}</ref><ref name=ls>{{citation|url=http://www.cbsnews.com/8301-205_162-57542904/jurassic-turtle-graveyard-found-in-china/
|first=Megan|last= Gannon
|periodical=Livescience.com|date=October 31, 2012
|title=Jurassic turtle graveyard found in China}}</ref> ''See [[:Category:Jurassic fish]].''
|-
| [[File:Ichthyodectidae1.jpg|140px]]
| [[Ichthyodectidae]]
| [[File:Thrissops cf formosus 01.jpg|thumb|160px|right|This fossil Ichthyodectidae from the Lower Jurassic is one of the best conserved fossil fishes worldwide]]
 
The family [[Ichthyodectidae]] (literally "fish-biters") was a family of marine [[actinopterygii|actinopterygian]] fish. They first appeared 156 Ma during the [[Late Jurassic]] and disappeared during the K-Pg extinction event 65 Ma. They were most diverse throughout the Cretaceous period. Sometimes classified in the primitive [[bony fish]] [[order (biology)|order]] [[Pachycormiformes]], they are today generally regarded as members of the "bulldog fish" order Ichthyodectiformes in the far more advanced [[Osteoglossomorpha]]. Most ichthyodectids ranged between 1 and 5 meters (3–15&nbsp;ft) in length. All known [[taxa]] were [[predator]]s, feeding on smaller fish; in several cases, larger Ichthyodectidae preyed on smaller members of the family. Some species had remarkably large teeth, though others, such as ''[[Gillicus arcuatus]]'', had small ones and sucked in their prey. The largest Xiphactinus was 20 feet long, and appeared in the Late Cretaceous (below).
|-
! rowspan=5 style="background:#32cd32" | [[Cretacico]]
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Cretacico]] (145–65 milioni di anni fa)
|-
| [[File:Acipenser ruthenus Prague Vltava 1.jpg|140px]]
| ''[[Acipenseridae|Storioni]]''
| Gli storioni appaiono nei ritrovamenti fossili nel Cretacico superiore e, da quel momento, la loro morfologia ha subito ben pochi cambiamenti evolutivi, dando loro lo status di [[fossile vivente|fossili viventi]]<ref>B. G. Gardiner (1984) Sturgeons as living fossils. Pp. 148–152 in N. Eldredge and S.M. Stanley, eds. Living fossils. Springer-Verlag, New York.</ref><ref name=krieger>{{cite journal | author = Krieger J., Fuerst P.A. | year = 2002 | title = Evidence for a Slowed Rate of Molecular Evolution in the Order Acipenseriformes | url = | journal = Molecular Biology and Evolution | volume = 19 | issue = | pages = 891–897 }}</ref>. </br>
La loro lentissima evoluzione è dovuta ad alcuni fattori fondamentali: un lungo intervallo generazionale, un'ampia tolleranza alle diverse temperature e salinità delle acque, la mancanza di predatori a causa delle loro grosse dimensioni e l'abbondanza di prede nel loro ambiente.
|-
| [[File:Enchodus petrosus.jpg|140px]]
| ''[[Enchodus]]''
|
|-
| [[File:Xiphactinus audax fossil.jpg|140px]]
| ''[[Xiphactinus]]''
|
Il più grande dei [[teleostei]] conosciuti, questo pesce del Cretacico raggiungeva una lunghezza di 4,5 metri <ref>Rafferty, John P (2010) [http://books.google.co.nz/books?id=kFd-1DApcRYC&pg=PA219&dq=%22Cretaceous+fish%22+OR+%22Cretaceous+fishes%22&hl=en&sa=X&ei=HkrtUNPWHonKlAWshYGQBQ&ved=0CKEBEOgBMBM#v=onepage&q=%22Cretaceous%20fish%22%20OR%20%22Cretaceous%20fishes%22&f=false ''The Mesozoic Era: Age of Dinosaurs''] pag. 219, Rosen Publishing Group. ISBN 9781615301935.</ref>.
|-
| colspan="3" style="line-height:16px; background:#dddddd;"| <small>The end of the Cretaceous was marked by the [[Cretaceous–Paleogene extinction event]] (K-Pg extinction). There are substantial fossil records of [[Gnathostomata|jawed]] [[fish]]es across the K–T boundary, which provides good evidence of extinction patterns of these classes of marine vertebrates. Within [[Chondrichthyes|cartilaginous fish]], approximately 80% of the [[shark]]s, [[Rajiformes|rays]], and [[skate]]s families survived the extinction event,<ref name="MacLeod">{{cite journal|author=MacLeod, N, Rawson, PF, Forey, PL, Banner, FT, Boudagher-Fadel, MK, Bown, PR, Burnett, JA, Chambers, P, Culver, S, Evans, SE, Jeffery, C, Kaminski, MA, Lord, AR, Milner, AC, Milner, AR, Morris, N, Owen, E, Rosen, BR, Smith, AB, Taylor, PD, Urquhart, E & Young, JR|title=The Cretaceous–Tertiary biotic transition|year=1997|journal=Journal of the Geological Society|volume=154|issue=2|pages=265–292|url=http://findarticles.com/p/articles/mi_qa3721/is_199703/ai_n8738406/print|doi=10.1144/gsjgs.154.2.0265}}</ref> and more than 90% of [[teleostei|teleost fish]] (bony fish) families survived.<ref>{{cite book |last=Patterson|first=C |year=1993 |title=Osteichthyes: Teleostei. In: The Fossil Record 2 (Benton, MJ, editor) |publisher=Springer |pages=621–656 |isbn=0-412-39380-8}}</ref> There is evidence of a mass kill of bony fishes at a fossil site immediately above the K–T boundary layer on [[Seymour Island]] near Antarctica, apparently precipitated by the K–Pg extinction event.<ref>{{cite journal |title=Discovery of fish mortality horizon at the K–T boundary on Seymour Island: Re-evaluation of events at the end of the Cretaceous |author=Zinsmeister WJ |date=1 May 1998|url=http://jpaleontol.geoscienceworld.org/cgi/content/abstract/72/3/556?maxtoshow=&HITS=10&hits=10&RESULTFORMAT=1&author1=zinsmeister&andorexacttitle=and&field_name=fulltext&searchid=1&FIRSTINDEX=0&sortspec=relevance&fdate=7/1/1927&tdate=7/31/2007&resourcetype=HWCIT |accessdate=2007-08-27 |journal=Journal of Paleontology |volume=72 |issue=3 |pages=556–571 }}</ref> However, the marine and freshwater environments of fishes mitigated environmental effects of the extinction event.<ref name="Robertson">{{cite journal |title= Survival in the first hours of the Cenozoic |author=Robertson DS, McKenna MC, Toon OB, Hope S, Lillegraven JA |journal=GSA Bulletin |year=2004 |volume=116 |issue=5–6 |pages=760–768 |doi=10.1130/B25402.1 |url=http://www.ugcs.caltech.edu/~presto/cenozoic.pdf |format=PDF|accessdate=2007-08-31}}</ref></small>
|-
! rowspan=3 style="background:#FFFF00" | [[Cenozoico]]
| colspan="3" style="text-align:center; background:#ddf8f8;"| [[Cenozoic Era]] (65 Ma to present): The current era has seen great diversification of bony fishes. Over half of all living vertebrate species (about 32,000 species) are fishes (non-tetrapod craniates), a diverse set of lineages that inhabit all the world's aquatic ecosystems, from snow minnows (Cypriniformes) in Himalayan lakes at elevations over 4,600 metres (15,000 feet) to flatfishes (order Pleuronectiformes) in the Challenger Deep, the deepest ocean trench at about 11,000 metres (36,000 feet). Fishes of myriad varieties are the main predators in most of the world’s water bodies, both freshwater and marine.
|-
| [[File:Amphistium.JPG|140px]]
| ''[[Amphistium]]''
| ''[[Amphistium]]'' is a 50-million-year-old fossil fish which has been identified as an early relative of the flatfish, and as a transitional fossil.<ref name="NaEvo">{{cite news | url=http://news.nationalgeographic.com/news/2008/07/080709-evolution-fish.html | title=Odd Fish Find Contradicts Intelligent-Design Argument | publisher=[[National Geographic Society|National Geographic]] |date= July 9, 2008 | first= | last= | accessdate =2008-07-17}}</ref> In a typical modern flatfish, the head is asymmetric with both eyes on one side of the head. In ''Amphistium'', the transition from the typical symmetric head of a vertebrate is incomplete, with one eye placed near the top of the head.<ref>{{cite journal|author=Matt Friedman|title=The evolutionary origin of flatfish asymmetry|journal=Nature|volume=454|number=7201|date=2008-07-10|pages=209–212|doi= 10.1038/nature07108|pmid=18615083|issue=7201}}</ref>
|-
| [[File:Megalodon NT.jpg|140px]]
| ''[[Megalodon]]''
|
''[[Carcharodon megalodon]]'' è una specie estinta di [[squalo]] che visse tra i 28 e 1,5 milioni di anni fa. Fisicamente simile al Sembrava molto simile al [[grande squalo bianco]], era tuttavia decisamente più grande, con una lunghezza stimata su ritrovamenti fossili che raggiunge i 20,3 metri <ref name="GWB" /> Found in all oceans<ref name="AN">{{Cite journal|last=Pimiento|first=Catalina|coauthors=Dana J. Ehret, Bruce J. MacFadden, and Gordon Hubbell|title=Ancient Nursery Area for the Extinct Giant Shark Megalodon from the Miocene of Panama|journal=PLoS ONE|volume=5|issue=5|pages=e10552|publisher=PLoS.org|___location=Panama|date=May 10, 2010|pmid=20479893|url=http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0010552|pmc=2866656|doi=10.1371/journal.pone.0010552|accessdate=12 May 2010|editor1-last=Stepanova|editor1-first=Anna|bibcode = 2010PLoSO...510552P }}, anche se si ipotizza che superasse queste dimensioni </ref> it was one of the largest and most powerful predators in vertebrate history,<ref name="GWB">{{cite journal|last=Wroe|first=S.|coauthors= Huber, D. R. ; Lowry, M. ; McHenry, C. ; Moreno, K. ; Clausen, P. ; Ferrara, T. L. ; Cunningham, E. ; Dean, M. N. ; Summers, A. P.|title=Three-dimensional computer analysis of white shark jaw mechanics: how hard can a great white bite?|url=http://www.bio-nica.info/Biblioteca/Wroe2008GreatWhiteSharkBiteForce.pdf|journal=Journal of Zoology|volume=276|issue=4|pages=336–342|year= 2008|doi=10.1111/j.1469-7998.2008.00494.x}}</ref>, Il ''Megalodon'' viveva in tutti gli oceani ed era uno dei più grandi e potenti predatori nella storia dei vertebrati che ha avuto un profondo impatto sulla vita marina<ref name="LV">{{Cite journal|doi=10.1038/nature09067|last=Lambert|first=Olivier|coauthors=Giovanni Bianucci, Klaas Post, Christian de Muizon, Rodolfo Salas-Gismondi, Mario Urbina and Jelle Reumer|title=The giant bite of a new raptorial sperm whale from the Miocene epoch of Peru|journal=Nature|volume=466|issue=7302|pages=105–108|___location=Peru|date=1 July 2010|url=http://www.nature.com/nature/journal/v466/n7302/full/nature09067.html|pmid=20596020|bibcode = 2010Natur.466..105L }}</ref>
|}
 
</center>
==Pesci preistorici==
{{Vedi anche|Elenco dei pesci preistorici}}
I pesci preistorici sono pesci estinti conosciuti soltanto tramite ritrovamenti [[fossili]]. Considerati tra i [[vertebrati]] più antichi oggi conosciuti, comprendono il primo pesce estinto, vissuto tra il [[Cambriano]] e il [[Terziario]]. La branca della [[paleontologia]] che studia i pesci preistorici si chiama paleoittiologia. Alcune specie viventi di pesci sono classificati come pesci preistorici, come il [[celacanto]] che è anche un [[fossile vivente]], a causa della loro rarità attuale e somiglianza con forme preistoriche estinte.
 
{{clear}}
 
==Fossili viventi==
[[File:Pacific hagfish Myxine.jpg|thumb|right|The jawless [[hagfish]] is a living fossil which has not essentially changed for 300 million years.<ref>[http://www.ucmp.berkeley.edu/vertebrates/basalfish/myxini.html Myxini] - University of California Museum of Paleontology</ref>]]
 
{{div col|3}}
'''Bony fishes'''
* [[Arowana]] and [[Arapaima]]
* [[Bowfin]]
* [[Coelacanth]]
* [[Gar]]
* [[Queensland lungfish]]
* [[Sturgeons]] and [[paddlefish]]
* [[Bichir]]
* [[Polypterus retropinnis]]
 
'''Sharks'''
* [[Blind shark]]
* [[Bullhead shark]]
* [[Elephant shark]]
* [[Frilled shark]]
* [[Goblin shark]]
* [[Gulper shark]]
 
'''Jawless fishes'''
* [[Hagfish]]
* [[Northern brook lamprey]]
 
'''Eels'''
* [[Protoanguilla palau]]
{{div col end}}
 
The [[coelacanth]] was thought to have gone extinct {{Ma|65}}, until a living specimen belonging to the [[order (biology)|order]] was discovered in 1938
 
{{clear}}
 
==Siti di fossili==
[[File:Miguasha affleurement 2.jpg|thumb|right|[[Miguasha National Park]]: outcrop of Devonian beds rich in fossil fish]]
{{see also|List of fossil sites}}
Some fossil sites which have produced notable fish fossils.
 
{{div col|3}}
* [[Abbey Wood SSSI]]
* [[Bracklesham Beds]]
* [[Bear Gulch Limestone]]
* [[Burgess Shale]]
* [[Canowindra]]
* [[Crato Formation]]
* [[Dura Den]]
* [[Feltville Formation]]
* [[Fossil Butte National Monument]]
* [[Fur Formation]]
* [[Gogo Formation]]
* [[Green's Creek]]
* [[Green River Formation]]
* [[Kakwa Provincial Park and Protected Area|Kakwa Provincial Park]]
* [[Land Grove Quarry, Mitcheldean|Land Grove Quarry]]
* [[Maotianshan Shales]]
* [[Matanuska Formation]]
* [[McAbee Fossil Beds]]
* [[Miguasha National Park]]
* [[MoClay]]
* [[Monte Bolca]]
* [[Mount Ritchie]]
* [[Orcadian Basin]]
* [[Portishead Pier to Black Nore SSSI]]
* [[Santana Formation]]
* [[Southerham Grey Pit]]
* [[Thanet Beds]]
* [[Towaco Formation]]
* [[Weydale]]
* [[Zhoukoudian]]
{{div col end}}
 
{{clear}}
 
==Collezioni di fossili==
Some notable fossil fish collections.
 
{{div col|1}}
* [http://www.nhm.ac.uk/research-curation/collections/our-collections/fossil-vertebrate-collections/fishes/index.html Fossil fish collection] ''[[Natural History Museum]]'', Britain.
* [http://www.synthesys.info/de_taf_mfn.htm Collection and expertise] ''[[Museum für Naturkunde]]'', Germany.
* [http://fieldmuseum.org/explore/department/geology/fossil-fishes Fossil fishes] ''[[The Field Museum]]'', United States.
{{div col end}}
 
==Paleoittiologi==
La paleoittiologia è la scienza che studia i pesci preistorici. Nella lista sottostante sono elencati i paleoittiologi che con i loro studi hanno contribuito a svelare i segreti di queste antiche forme di vita.
{{div col|3}}
* [[Louis Agassiz]]
* [[Mary Anning]]
* [[Michael Benton]]
* [[Derek Briggs]]
* [[Hans C. Bjerring]]
* [[John Samuel Budgett]]
* [[Frederick Chapman]]
* [[Jenny Clack]]
* [[Ted Daeschler]]
* [[Bashford Dean]]
* [[Robert Dick]]
* [[Sir Philip Grey Egerton, 10th Baronet|Philip Grey Egerton]]
* [[Edwin Sherbon Hills]]
* [[Jeffrey A. Hutchings]]
* [[Thomas Henry Huxley]]
* [[Johan Aschehoug Kiær]]
* [[Philippe Janvier]]
* [[Erik Jarvik]]
* [[George V. Lauder]]
* [[John A. Long]]
* [[Hugh Miller]]
* [[Charles Moore (geologist)|Charles Moore]]
* [[Paul E. Olsen]]
* [[Heinz Christian Pander]]
* [[Elizabeth Philpot]]
* [[Jean Piveteau]]
* [[Colin Patterson (biologist)|Colin Patterson]]
* [[Alfred Romer]]
* [[Ira Rubinoff]]
* [[Neil Shubin]]
* [[Franz Steindachner]]
* [[Erik Stensiö]]
* [[Ramsay Heatley Traquair]]
* [[Thomas Stanley Westoll]]
* [[Tiberius Cornelis Winkler]]
* [[Arthur Smith Woodward]]
{{div col end}}
 
== Voci correlate ==
* [[Comparative anatomy]]
* [[List of examples of convergent evolution#Fish|Convergent evolution in fish]]
* [[Ichthyolith]]
* [[Old Red Sandstone]]
* [[Prehistoric life]]
* [[Vertebrate paleontology]]
 
{{clear}}
 
== References ==
 
===Citations===
<!------------------------------------------------------------
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the {{r}} and {{sfn}} templates to keep the body
text clean. Please follow existing examples within the text
and refer to the following documentation pages if needed:
 
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Template {{r}}:
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Template {{sfn}}:
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discussion of different citation methods and how to generate
footnotes using the <ref> tags.
------------------------------------------------------------->
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<ref name="Shu2003">{{cite journal | author=Shu, D-G., Conway Morris, S., Han, J., ''et al.'' | title=Head and backbone of the Early Cambrian vertebrate Haikouichthys| journal=Nature | volume=421 | pages=526–529 | date=January 2003 | doi=10.1038/nature01264 | url=http://www.nature.com/nature/journal/v421/n6922/abs/nature01264.html | accessdate=September 21, 2008 | pmid=12556891 | issue=6922 | bibcode=2003Natur.421..526S}}</ref>
 
<ref name="SL2009">[http://www.arn.org/blogs/index.php/literature/2009/03/27/critical_transitions_in_fish_evolution_l Post details: Critical transitions in fish evolution lack fossil documentation] ''Science Literature'', 27 March 2009.</ref>
 
<ref name="SydneyMorningHerald2006">[http://www.smh.com.au/news/science/jaws-of-steel-on-this-fish-tank/2006/11/29/1164777657728.html Monster fish crushed opposition with strongest bite ever]. The Sydney Morning Herald. November 30, 2006.</ref>
 
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{{refend}}
 
==Further reading==
{{ external media
| align = right
| width = 216px
| video1 = [https://www.youtube.com/watch?v=wJunXtPFK-0 Feeding Mechanism of Conodonts] – ''YouTube''
| video2 = [https://www.youtube.com/watch?v=ypYesuV3PoI Chordate evolution] – ''YouTube''
}}
* Benton MJ (1998) [http://palaeo.gly.bris.ac.uk/Essays/vertfr/default.html "The quality of the fossil record of the vertebrates"] Pages 269–303 in Donovan, SK and Paul CRC (eds), ''The adequacy of the fossil record''. Wiley. ISBN 9780471969884.
* {{cite journal | author = Cloutier R | year = 2010 | title = The fossil record of fish ontogenies: Insights into developmental patterns and processes | url = http://www.fishlarvae.com/common/sitemedia/Cloutier%202010%20Sem%20Cell%20Dev%20Biol%20The%20fossil%20record%20of%20fish%20ontogenies.pdf | format = PDF | journal = Seminars in Cell & Developmental Biology | volume = 21 | issue = | pages = 400–413 }}
* Janvier, Philippe (1998) ''Early Vertebrates'', Oxford, New York: Oxford University Press. ISBN 0-19-854047-7
* Long, John A. (1996) [http://books.google.co.nz/books?id=dEP_kQAACAAJ&dq=%22The+rise+of+fishes%22&hl=en&sa=X&ei=8WgLUdOMGo2IkgWZzYGwBw&ved=0CC8Q6AEwAA ''The Rise of Fishes: 500 Million Years of Evolution''] Johns Hopkins University Press. ISBN 0-8018-5438-5
* McKenzie DJ, Farrell AP and Brauner CJ (2011) [http://books.google.co.nz/books?id=gfBc_omOIeAC&printsec=frontcover&dq=%22Fish+physiology%22&hl=en&sa=X&ei=454PUeGDLsGimQXCpYCIBA&ved=0CF4Q6AEwCQ#v=onepage&q=%22Fish%20physiology%22&f=false ''Fish Physiology: Primitive Fishes''] Academic Press. ISBN 9780080549521.
* Maisey JG (1996) [http://books.google.co.nz/books?id=gAiAPwAACAAJ&dq=editions:_y8O0MW7spIC&hl=en&sa=X&ei=UmgLUZGFNMemkQXjzoGoDA&ved=0CC8Q6AEwAA''Discovering fossil fishes''] Holt. ISBN 9780805043662.
* [[Neil Shubin|Shubin, Neil]] (2009) [http://books.google.co.nz/books?id=c008kdNwR1cC&dq=%22Your+Inner+Fish%22&hl=en&sa=X&ei=SePbUKKxHIavkgXllIHAAQ&ved=0CDQQ6AEwAA ''Your inner fish: A journey into the 3.5-billion-year history of the human body''] Vintage Books. ISBN 9780307277459.
* [http://www.ucmp.berkeley.edu/vertebrates/vertintro.html Introduction to the Vertebrates] ''Museum of Palaeontology'', University of California.
 
==External links==
* [http://hoopermuseum.earthsci.carleton.ca/12.html Fossil Fish]
* [http://www.ukwetlandhabitats.co.uk/fishbiologyandbehavior.html Origins of Fish]
* [https://www.youtube.com/watch?v=gZpsVSVRsZk Overview of evolution] – [[Carl Sagan]]
 
{{diversity of fish}}
{{evolution of fish|state=expanded}}
{{evolution}}
{{fins, limbs and wings}}