# Tetrapoda

| Tetrapoda |
| --- |
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|  Irudi gehiago |
| Sailkapen zientifikoa |
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| Erreinua | Animalia |
| Filuma | Chordata
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| Superklasea  | **Tetrapoda** ezezaguna 1912ra arte
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| Azpibanaketa |
| * Amphibia klasea
* Amniota
	+ Sauropsida kladoa
		- Reptilia klasea
		- Aves klasea
	+ Synapsida kladoa
		- Mammalia klasea
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**Tetrapoda** (grezieraz *lau hanka*) animalia ornodun klado bat da, lau adarrekin. Anfibio, narrasti, ugaztun eta hegaztiak tetrapodoak dira, baita sugeak edo hankarik gabeko anfibioak ere. Sarcopterigii taldetik eboluzionatu zuten Devoniar garaian.

Lau luzakin dituzten animalia ornodunez esaten da. Tetrapodoen luzakinak agerikoak edo atrofiatuak izan daitezke. Arrainak izan ezik gainerako ornodun guztiak dira tetrapodoak, anfibioak, narrastiak, hegaztiak eta ugaztunak alegia.

Tetrapodoek Tetrapodomorpho izenez ezagutzen diren animalia erdiurtar primitiboen klado batetik aurrera eboluzionatu zuten, zeinak, era berean, antzinako arrain lobulu finetatik (sarkopterygiarrak) duela 390 milioi urte inguru eboluzionatu zuen Devoniar Erdi Aroan; haien formak arrain hegatsadunen eta lau hankako benetako tetrapodoen arteko iragaitekoak ziren. Gorputz-adar ornodunak (tetrapodoak, hitzaren zentzu zabalean) Devoniar Erdiko arrastoetatik ezagutzen dira lehenik, eta gorputzeko fosilak Devoniar Berantiarraren amaieratik hurbil bihurtu ziren, baina denak uretakoak ziren. Lehen koroa-tetrapodoak (lehorreko lokomoziorako gai diren tetrapodo zaharren azken arbaso arruntak) Karbonifero oso goiztiarrean agertu ziren, duela 350 milioi urte.

Tetrapodoen arbaso akuatikoak eta uretatik irten ondoren Lurraren lurra kolonizatzeko prozesua ez dago argi oraindik. Ur-arnasketarako zakatzetan oinarrituriko gorputzeko eta isatsaren bulkadan oinarritutako lokomozio-plan batetik animalia uretatik atera eta lehorrean mugitzeko aukera ematen duen batera igarotzea da ezagutzen den eboluzio-aldaketa sakonenetako bat. Tetrapodoek, beren arrain aurrekariengandik bereizten dituzten hainbat ezaugarri dituzte. Horien artean, elikadurarako eta mugimenduetarako buru eta lepo egitura ezberdinak daude, eskeleto apendikularrak (sorbalda eta pelbiseko gerrikoak bereziki) pisu eta lokomoziorako, begi moldakorragoak ikusteko, erdiko belarriak entzumenerako, eta bihotz eta birika eraginkorragoak oxigenoa mugitzeko eta uretatik kanpo trukatzeko.

Lehen tetrapodoak (parafiletikoa) edo *isopodoak* nagusiki uretakoak izan ziren. Anfibio modernoak, aurreko taldeetatik eboluzionatu zutenak, erdiurtarrak izan ziren eskuarki; beren bizitzako lehen faseak zapaburu bezala ezagutzen diren arrainen antzeko larba eta arrautzak dira, eta, geroago, metamorfosia jasaten dute gorputz-adarrak hazteko eta zati batean lehorrekoak eta beste zati batean uretakoak bihurtzeko. Hala ere, gaur egun, tetrapodoak diren espezie gehienak amniotak dira, eta gehienak lehorreko tetrapodoak dira, zeinen adarrak Karbonifero Berantiarrean lehen tetrapodoetatik eboluzionatu baitzuen. Anfibioen gaineko amniotetan funtsezko berrikuntza amniona da, zeinak arrautzek lehorrean eduki eta urtsua gordetzeko aukera ematen baitu, uretan geratu beharrean. (Zenbait amniotek geroago eboluzionatu zuten barne-ernalkuntzan, nahiz eta tetrapodo zuhaitzetik kanpoko espezie urtar askok tetrapodoak, adibidez *Materpiscis*, agertu aurretik eboluzionatu zuten). Tetrapodo batzuek, hala nola sugeek eta zeziliek, beren gorputz-adar batzuk edo guztiak galdu dituzte espeziazio eta eboluzio handiagoaren ondorioz, eta beste batzuek, berriz, beren arbasoen gorputz-adarren hondar gisa ezkutatu besterik ez dute egin. Beste batzuk anfibio izatera itzuli ziren edo, bestela, bizitza partzialki edo erabat urtar izatera, lehen karboniferoen garaian, beste batzuk zenozoiko bezain garai hurbilean.

Amniota talde bat narrastietan zatitu zen: lepidosauroak, dinosauroak (hegaztiak barne), krokodiloak, dortokak eta ahaide iraungiak barne hartzen dituena; beste amniote-talde bat, berriz, ugaztunetara eta haien ahaide iraungietara zatitu zen. Amnioten artean, hegaz egiteko garatu ziren tetrapodoak badaude, hala nola dinosauroen artetik datozen hegaztiak: archosauroen pterosauroak eta ugaztunen artetik datozen saguzarrak.

## Filogenia

Tetrapodoen kladograma, garafikoki erakutsia.
### Erroko tetrapodoak

Erroko tetrapodoak Tetrapodatik Dipnomorpha arrainetatik baino gertuago dauden animaliak dira, baina tetrapodoen koroa-taldea kenduta. Kladograma honek erroko tetrapodoen arteko harremana erakusten du.

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### Koroa-taldea

Tetrapodoen koroa-taldea gaur egun bizirk dauden tetrapodo guztien (urlehortar, narrasti, hegazti eta ugaztun) azken arbaso komunaren eta bere ondorengo guztiak biltzen dituen taldea da.

Talde honetan hainbat desagertutako tetrapodo sartzea edo ez gaur egungo anfibioek edo Lissamphibiaren arteko harremanak zehaztearen araberakoa da. Gaur egun hiru hipotesi nagusi daude Lissamphibiari dagokionez. Temnospondyli hipotesian (TH), lissanbifioak Temnospondyli dissorofoideekin harremanduta daude, eta honek temnospondiloak tetrapodoen barnean kokatzea ekarriko luke. Lepospondyli hipotesian (LH), lisanfibioak talde honen ahizpa-taxoia lirateke, lepospondiloak tetrapodo eginez eta temnospondiloak erroko tetarpodo eginez. Hipotesi polifiletikoan (PH) igelak eta uhandreak bere kabuz eboluzionatu zuten temnospondilo dissorofoideetatik eta Apoda taldea Microsauria taldeko lepospondiloengandik; azken honetan bai lepospondiloak bai temnospondiloak tetrapodo izango lirateke.

#### Temnospondyli hipotesia (TH)

Hipotesi hau hainbat bariante kontuan hartuta egiten da, gehienetan lissanfibioak temnosponyli dissorofoideengandik etortzen direlarik, askota anfibamidae eta branchiosauridae aipatuz.

Temnospondyli hipotesia gaur egun babesle gehien dituena da, besteak beste Ruta *et al* (2003a,b), Ruta eta Coates (2007), Coates *et al* (2008), Sigurdsen eta Green (2011), eta Schoch (2013,2014).

Kladograma honen jatorria Coates, Ruta eta Friedman (2008) da.

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| **Tetrapoda koroa-taldea**  |

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| *Temnospondyli* |

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|  | **Lissamphibia koroa taldea**  |
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|  | **Amniota koroa taldea**  |
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Microsauria  |
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Nectridea  |
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Aistopoda  |
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Lissamphibia talde totala
Amniota talde totala |

#### Lepospondyli hipotesia

Kladograma hau Laurinen *How Vertebrates Left the Water* liburutik moldatu da:

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| **Stegocephalia ("Tetrapoda")**  |

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|  | *Acanthostega*  |
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|  | *Seymouriamorpha*  |
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| *Reptiliomorpha* |

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|  | *Aistopoda*  |
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|  | *Lysorophia*  |
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|  | **Lissamphibia**  |
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Erroko tetrapodoak
Amniota talde totala
Lissamphibia talde totala |

### Hipotesis polifiletikoa (PH)

Hipotesi honen arabera batrazioak (igelak eta uhandreak) temnospondilo dissoropfoideengatik eboluzionatu zuten, eta apodoak mikrosaurio lepospondiloetatik. Bi bariante daude, bata Carrollek eginekoa, eta bestea Andersonek.

Kladograma hau Schoch, Frobisch, (2009) lanetik moldatu da.

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| **Tetrapoda**  |

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stem tetrapods |
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| *Temnospondyli* |

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basal temnospondyls |
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| *Dissorophoidea* |

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|  | *Amphibamidae* |
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|  | **Igelak**  |
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|  | *Branchiosauridae*  |
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|  | **Uhandreak**  |
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| *Lepospondyli* |

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|  | *Lysorophia*  |
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|  | **Apoda**  |
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|  | *Microsauria*  |
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|  | *Seymouriamorpha*  |
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|  | *Diadectomorpha*  |
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|  | *Amniota*  |
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## Erreferentziak