# Tetrapoda

| Tetrapoda |
| --- |
| Raspon fosila: Rani devion → Sadašnjost |
| *Salamandra salamandra* |
| Sistematika |
| Natcarstvo | Eukaryota |
| Carstvo | Animalia |
| Potcarstvo | Eumetazoa |
| Natkoljeno | Deuterostomia |
| Koljeno | Chordata |
| Potkoljeno | Vertebrata |
| Infrakoljeno | Gnathostomata |
| Natklasa | **Tetrapoda** |
| Razredi, podazredi i infrarazredi |
| * Razred Amphibia
* Podrazred Lissamphibia
* Serija Amniota
* Razred Reptilia
* Podrazred Anapsida
* Podrazred Diapsida
* Infrarazred Archosauromorpha
* Infrarazred Lepidosauromorpha
* Razred Aves
* Podrazred Carinatae
* Infrarazred Neornithes
* Razred Mammalia
* PodrazredPrototheria
* Infrarazred Australosphenida
* Podrazred Theria
* Infrarazred Metatheria
* Infrarazred Eutheria
 |

**Četvoronošci**, **tetrapode** ili **kopneni kičmenjaci** (lat. Tetrapoda, od grčkog *tetre* = četiri + *poda* = ud, noga), u biološkoj sistematici obuhvataju sve kičmenjake koji imaju četiri noge. U ovu grupu se ubrajaju vodozemci i amnioti: gmizavci, ptice i sisari. Četiri noge mogle su se tokom evolucije sekundarno ponovo izgubiti, kao što je to slučaj sa zmijama, ili su se prednje noge preoblikovale u peraja (kitovi), krila (ptice, šišmiši) ili ruke. Danas ovu grupa obuhvata oko 26.700 vrsta. Dominantni su u svim ekosistemima, a s pticama i šišmišima, osvojili su i zrak. Neke vrste kopnenih kičmenjaka ponovo su se dijelom (perajari, pingvini) ili potpuno (kitovi, morske krave) vratili životu u vodi.

Četveronošci su evoluirali od mesoperki, prije oko 395 miliona godina, tokom devonaa. Tačni vodeni preci tetrapoda, kao i proces kolonizacije kopna, nisu sigurno određeni i o tim temama još uvijek postoje polja aktivnog istraživanja i diskusije među paleontolozima.

Iako je većina današnjih vrsta kopnena, postoji samo mali broj dokaza koji pružaju podršku stavu da se i jedan od najranijih tetrapoda mogao kretati na kopnu, budući da im udovi nisu mogli podići tijelo s tla, a pronađeni tragovi ne ukazuju da su ga vukli po tlu. Tragove su po svoj prilici ostavile životinje koje su hodale po dnu plitkih vodenih bazena. Vodozemci danas općenito još uvijek žive vodo-zemno, tako što prvi stadij života provedu kao riboliki punoglavci. Nekoliko grupa tetrapoda, kao što su zmije i kitovi, izgubili su udove. Pored toga, mnogi su se vratili, djelimično ili potpuno, životu u vodi (u moderne primjere spadaju kitovi i morske krave). Moguće je da se prvi povratak na život u vodi dogodio čak tokom perioda krede, dok su se noviji pojavili u kenozoiku u vodi, kao što su kitovi i perajari, i nekolicina Lissamphibia.

Promjena sa organizacije telesnog plana stvorenog za disanje i kretanje u vodi – na plan građe koji omogućava kretanje na kopnu, jedna je od najdubljih poznatih evolucijsikh promjena. Takođe je jedna od najbolje istraženih, većinski zahvaljujući velikom broju značajnih prelaznih fosila otkrivenih u kasnom 20. veku, kao i usavršenoj filogenetičkoj analizi.

Unatoč pojavi da većinu tetrapoda čine četvoronošci, ta dva pojma nemaju isto značenje. Četvoronošci su općenita kategorija u koju spada bilo koja životinja koja hoda na četiri noge, bila ona tetrapoda ili ne. Tetrapode su taksonomska kategorija koja obuhvata razrede vodozemaca, gmizavaca, ptica i sisara, čiji se pripadnici mogu, ali ne moraju kretati na četiri noge.

## Raznolikost

U tetrapode spadaju tri razreda: vodozemci, gmizavci, ptice i sisari. Sveukupni diverzitet podrazreda Lissamphibia, kao i tetrapoda gopćenito, s vremenom se eksponencijalno povećavao. Danas ih ima više od 30.000 vrsta, koje potiču od samo jedne grupe vodozemaca iz ranog do srednjeg devona. Međutim, taj proces diversifikacije prekinut je bar nekoliko puta, velikim biološkim krizama, kao što je [[permsko-trijasko Izumrli, koje su, u najblažem slučaju, imale uticaj bat na amniote. Cjelokupni sastav diverziteta u paleozoiku predvodili su vodozemci, u mezozoiku dominirali su gmizavci, nakon čega je u kenozoiku došlo do eksplozivnog porasta broja vrsta ptica i sisara. Kako je diverzitet rastao, tako se proširivao i broj niša koje su tetrapode zauzimale. Prve su bile vodeni oblici i pretežno su se hranili ribama. Na Zemlji danas postoji veliki diverzitet tetrapoda, koje žive u mnogim staništima i imaju raznoliku ishranu.

## Evolucija

### Porijeklo

Devonske ribe, uključujući i ranog morskog psa *Cladoselachia*, *Eusthenopterona* i ostale mezoperke, kao i oklopnjaču *Bothriolepis* (Joseph Smit, 1905)
Tetrapode evoluirali su od ranih košljoriba (*Osteichthyes*) koje su živjele u slatkovodnim i bočatim staništima na samom početku devona. Te ribe imale su mnoge osobine zajedničke sa svojim hrskavičavim precima, ali također i bitne razlike, najviše u pogledu ribljeg mehura/pluća, obilježja koju morski psi i raže nisu imali.

*Eusthenopteron*, prije ~385 miliona godina
Pluće/riblji mjehur nastalo je kao izraštaj stomaka, formirajući mjehur iznad probavnog trakta, ispunjen plinom. Primarna funkcija tog organa u njegovom primitivnom obliku još uvijek nije sigurno utvrđena. Vjerovatno su ga koristili kao pluća za udisanje zraka, ali mogao je imati ulogu u plovnosti. Moguće je da je imao obje funkcije.

*Tiktaalik*, prije ~375 miliona godina
Smatra se da su mesnate peraje podržane kostima nasljedno svojstvo svih košljoriba (Osteichthyes). Preci zrakoperki (Actinopterygii) evoluirali su peraja u drugačijem smjeru i razvili mehur sa isključivom ulogom u kontroli plovnosti. Tetrapodomorfni preci tetrapoda zadržali su mjehur sličan plućnomm krilu i dalje razvili mesnata peraja. Ona su imala kosti koje odgovaraju ramenoj, lakatnoj i palčanoj kosti, na prednjim udovima i bedrenoj, goljeničnoj i lisnoj kosti, na zadnjim udovima.

Iako su tetrapodomorfi većinom bili ribe otvorenih voda, grupa Elpistostegalia prilagodila se životu u plićacima. Razvili su horizontalno ravna tijela za kretanje u plitkim vodama, a ramena i karlična peraja preuzela su ulogu glavnih organa za kretanje. Budući da bi plićaci povremeno bili pogođeni nedostatkom kisika, sposobnost udisanja atmosferskog zraka, pomoću ribljeg mehura, postala je jako bitna.

*Acanthostega*, prije ~365 miliona godina
Prve tetrapode vjerovatno su, krajem devona, evoluirale od tetrapodomorfnih riba iz plitkih vodenih staništa.

## Kladistika

### Matična grupa

Matične tetrapode su sve životinje koje su bliže povezane sa tetrapodama nego s plućima, ali isključujući krunsku tetrapodnu grupu. Kladogram u nastavku ilustrira odnos matičnih tetrapoda , prema, 2012:

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| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| Rhipidistia |

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Dipnomorpha  |
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| Tetrapodomorpha  |

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|   | *Kenichthys* |
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Rhizodontidae  |
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| Canowindridae |

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|   | *Marsdenichthys* |
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|   | *Canowindra* |
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|   | *Koharalepis* |
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|   | *Beelarongia* |
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| Megalichthyiformes |

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|   | *Gogonasus*  |
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|   | *Gyroptychius* |
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|   | *Osteolepis*  |
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|   | *Medoevia* |
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Megalichthyidae |
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| Eotetrapodiformes |

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| Tristichopteridae |

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|   | *Spodichthys* |
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|   | *Tristichopterus* |
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|   | *Eusthenopteron*  |
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|   | *Jarvikina* |
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|   | *Cabbonichthys* |
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|   | *Mandageria* |
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|   | *Eusthenodon* |
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|   | *Tinirau* |
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|   | *Platycephalichthys* |
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| Elpistostegalia |

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|   | *Panderichthys*  |
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| Stegocephalia |

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|   | *Tiktaalik*  |
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|   | *Elpistostege* |
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|   | *Elginerpeton*  |
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|   | *Ventastega* |
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|   | *Acanthostega*  |
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|   | *Ichthyostega*  |
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Whatcheeriidae  |
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Colosteidae  |
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|   | *Crassigyrinus*  |
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Baphetidae |
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Tetrapoda  |
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### Krunska grupa

Krunske tetrapode definirane su kao najbliži drevni zajednički predak svih živih tetrapoda (vodozemaca, gmizavaca, ptica i sisara) zajedno sa svim potomcima tog pretka.

Uključivanje određenih izumrlih grupa ovisi o odnosima modernih vodozemaca ili Lissamphibia n. Sada postoje tri glavne hipoteze o porijeklu lisamfibija. U temnospondilnoj hipotezi (TH), lisasamfibi su najuže povezani s disorofoidnim Temnospondilama, koji bi temnospondile uklhučili u tetrapode. U hipotezi o lepospondilu (LH), lisamfibije su sestrinski takson lisorofskih Lepospondila , čineći lepospondile tetrapodima i temnospondile matičnim tetrapodama. U polifiletskoj hipotezi (PH), žabe i daždevnjaci su evoluirali iz dizorofoidnih temnospondila, dok cecilije ishode iz lepospondilskih mikrosaura, čineći i lepospondile i temnospondile pravim tetrapodama.

### Temnospondilska hipoteza (TH)

Ova hipoteza dolazi u brojnim varijantama, od kojih većinu imaju lisasamfibi koji proizlaze iz dizorofoidnih temnospondila, obično s fokusom na amfibamidima i braniosauridima.

Ova hipoteza sada je preferirana ili je većinsko stanovište, koje podržavaju Ruta *et al* (2003a,b), Ruta and Coates (2007), Coates *et al* (2008), Sigurdsen and Green (2011) i Schoch (2013, 2014).

Kladogram modificiran prema Coates, Ruta and Friedman (2008).

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| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| **Krunska grupa Tetrapoda**  |

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| --- | --- | --- | --- | --- | --- |
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| *Temnospondyli* | **Krunska grupa Lissamphibia**  |
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 | Ukupna grupa Lissamphibia |
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|   | *Embolomeri*  |
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Gephyrostegidae  |
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Seymouriamorpha  |
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Diadectomorpha  |
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|   | **Krunska grupa Amniota**  |
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Microsauria  |
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Lysorophia  |
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Adelospondyli |
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Nectridea  |
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Aistopoda |
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 | Ukupna grupa Amniota |
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### Lepospondilska hipoteza (LH)

Kladogram, modificiran prema Laurin, *How Vertebrates Left the Water* (2010).

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|   **Stegocephalia**   |

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Izumrli *Acanthostega*  |
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|   | *Ichthyostega*  |
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Izumrli *Temnospondyli*  |
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Izumrli *Embolomeri*  |
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Izumrli *Seymouriamorpha*  |
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|   *Reptiliomorpha*   |

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|   |   **Amniota**      |
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|   | † *Diadectomorpha*  |
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|   *Amphibia*   |

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Izumrli *Aistopoda*   |
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Izumrli *Adelogyrinidae*   |
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Izumrli *Nectridea*  |
|   |

 |
|   |
|   |

|  |  |
| --- | --- |
|   |
Izumrli *Lysorophia*  |
|   |
|   |

|  |  |
| --- | --- |
|   |   **Lissamphibia**          |
|   |

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|   |

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|   |

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|   |

 | Šablon:Clade labels |
| (“Tetrapoda”) |

### Hipoteza polifiletskog porijrkla (PH)

U ovoj hipotezi batrahijata (žabe i daždevnjaci) proizliaze iz disorofoidnih temnospondila, a cecilije iz lepospondilskih mikrosaura. Postoje dvije varijante, jednu je razvio Carroll, a drugu Anderson.

Kladogram modificiran prema Schoch, Frobisch, (2009).

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   **Tetrapoda**   |

|  |  |
| --- | --- |
|   |   Matične tetrapode   |
|   |
|   |

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   |

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   *Temnospondyli*   |

|  |  |
| --- | --- |
|   |   Bazne temnospondile   |
|   |
|   *Dissorophoidea*   |

|  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- |
|   |

|  |  |
| --- | --- |
|   |
Izumrli *Amphibamidae*   |
|   |
|   |   **Žabe**  |
|   |

 |
|   |
|   |

|  |  |
| --- | --- |
|   |
Izumrli *Branchiosauridae*  |
|   |
|   |   **Salemandre**  |
|   |

 |
|   |

 |
|   |

 |
|   |
|   |

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   *Lepospondyli*   |

|  |  |
| --- | --- |
|   | † *Lysorophia*  |
|   |
|   |

|  |  |
| --- | --- |
|   |   **Caecilians**  |
|   |
|   | † *Microsauria*  |
|   |

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|  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|   |

|  |  |
| --- | --- |
|   | † *Seymouriamorpha*  |
|   |
|   |

|  |  |
| --- | --- |
|   |
Izumrli *Diadectomorpha*  |
|   |
|   |   *Amniota*  |
|   |

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|   |

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|   |

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|   |

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|   |

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|   |

## Reference

## Dopunska literatura

* Laurin, M. (2010). *How Vertebrates Left the Water*. Berkeley, California, USA.: University of California Press. ISBN 978-0-520-26647-6.CS1 održavanje: ref=harv (link)

* -{Robert L. Carroll: *Paläontologie und Evolution der Wirbeltiere*, Thieme, Stuttgart. Tetrapoda}-
* -{Volker Storch, Ulrich Welsch: *Systematische Zoologie*, Fischer. Tetrapoda}-
* Long JA, Young GC, Holland T, Senden TJ, Fitzgerald EM (2006). "An exceptional Devonian fish from Australia sheds light on tetrapod origins". *Nature*. **444** (7116): 199–202. Bibcode:2006Natur.444..199L. doi:10.1038/nature05243. PMID 17051154.
* Benton, Michael (5. 2. 2009). *Vertebrate Palaeontology* (3 izd.). John Wiley & Sons. str. 1. ISBN 978-1-4051-4449-0. Pristupljeno 10. 6. 2015.CS1 održavanje: ref=harv (link)
* Clack, J.A. (2012). *Gaining ground: the origin and evolution of tetrapods* (2nd izd.). Bloomington, Indiana, USA.: Indiana University Press. ISBN 9780253356758.CS1 održavanje: ref=harv (link)
* Laurin, Michel (2010). *How Vertebrates Left the Water*. University of California Press. ISBN 978-0-520-26647-6. Pristupljeno 26. 5. 2015.CS1 održavanje: ref=harv (link)
* McGhee, George R., Jr. (2013). *When the Invasion of Land Failed: The Legacy of the Devonian Extinctions*. Columbia University Press. ISBN 978-0-231-16057-5. Pristupljeno 2. 5. 2015.CS1 održavanje: ref=harv (link)
* Steyer, Sebastien (2012). *Earth Before the Dinosaurs*. Indiana University Press. str. 59. ISBN 978-0-253-22380-7. Pristupljeno 1. 6. 2015.CS1 održavanje: ref=harv (link)
* Clack, Jennifer A. (2009). "The Fin to Limb Transition: New Data, Interpretations, and Hypotheses from Paleontology and Developmental Biology". *Annual Review of Earth and Planetary Sciences*. **37** (1): 163–179. Bibcode:2009AREPS..37..163C. doi:10.1146/annurev.earth.36.031207.124146.
* Hall, Brian K., ured. (2007). *Fins Into Limbs: Evolution, Development, and Transformation*. Chicago: University of Chicago Press. ISBN 978-0-226-31340-5.CS1 održavanje: ref=harv (link)
* Benton, Michael (2005). *Vertebrate Palaeontology (Benton)* (3rd izd.). Blackwell Publishing.CS1 održavanje: ref=harv (link)

## Vanjski linkovi

* Michel Laurin: *Terrestrial Vertebrates* The Tree of Life Web Project Arhivirano 1. 3. 2011. na Wayback Machine
* Jennifer A. Clack: *The Definition of the Taxon Tetrapoda* The Tree of Life Web Project Arhivirano 18. 4. 2009. na Wayback Machine
* Michel Laurin, Marc Girondot and Armand de Ricqlès: *Early Tetrapod evolution*, PDF
* Marcello Ruta,, Jonathan E. Jeffery, Michael I. Coates: *A supertree of early tetrapods.* PDF
* What's a "Tetrapod"? Palaeos.com
* Introduction to the Tetrapoda University of California Museum of Paleontology
* Getting a Leg Up on Land Scientific America
* Devonian Times
* Fische auf dem Trocknen („Ribe na suvom”)