| TomistominosRango fósil: Eoceno - Actual(desde hai 55 millóns de anos) |
| --- |
| O falso gavial, *Tomistoma schlegelii* |
| Clasificación científica |
| 

|  |  |
| --- | --- |
| Reino: | Animalia |
| Filo: | Chordata |
| Clase: | Reptilia |
| (sen clasif.): | Archosauria |
| Orde: | Crocodilia |
| Superfamilia: | Gavialoidea |
| Familia: | Gavialidae |
| Subfamilia: | **Tomistominae**Kälin, 1955 |

 |
| Xéneros |
| * *Tomistoma*
* †*Dollosuchoides*
* †*Dollosuchus*
* †*Ferganosuchus*
* †*Gavialosuchus*
* †*Kentisuchus*
* †*Maroccosuchus*
* †*Megadontosuchus*
* †*Paratomistoma*
* †*Penghusuchus*
* †*Rhamphosuchus*
* †*Toyotamaphimeia*
* †*Thecachampsa*
 |

Os **tomistominos** (**Tomistominae**) son unha subfamilia de réptiles crocodilianos que comprende unha soa especie viva, o falso gavial ou gavial malaio. Coñécense tamén moitas especies fósiles encadradas nesta subfamilia, que estenden o rango da subfamilia ata o Eoceno. A diferenza do falso gavial, que é unha especie de auga doce que vive exclusivamente no sueste asiático, os tomistominos extintos tiñan unha distribución global e vivían en estuarios e ao longo da liña costeira.

A clasificación dos tomistominos dentro de Crocodilia foi moi discutida e flutuante; tradicionalmente se pensaba que estaban dentro da familia Crocodyloidea, xa que teñen moitas semellanzas morfolóxicas cos crocodilos, pero as evidencias moleculares indican que están máis próximos aos gaviais e dentro da familia Gavialoidea.

Descrición
----------

Os tomistominos teñen fociños estreitos ou loxirrostrinos como os dos gaviais indios. O actual falso gavial vive en augas doces e utiliza o seu longo fociño e afiados dentes para capturar peixes, aínda que os gaviais indios ou verdadeiros están máis adaptados á dieta piscívora. A pesar da semellanza cos gaviais, a forma dos ósos do cranio dos tomistominos líganos máis aos crocodilos. Por exemplo, os tomistominos e crocodilos teñen barras postorbitais delgadas detrás das órbitas oculares e unha gran cavidade para o quinto dente maxilar. O óso esplenial da mandíbula inferior é longo e delgado, adquirindo unha distintiva forma de V que non se observa nos gaviais.

Historia evolutiva
------------------

Cranio de *Kentisuchus toliapicus*.
Os tomistominos apareceron no Eoceno en Europa e Norte de África. O tomistomino máis antigo coñecido é *Kentisuchus spenceri* de Inglaterra, aínda que hai un resto fósil aínda máis antigo dun posible tomistomino do Paleoceno de España. Outros tomistominos iniciais son *Maroccosuchus zennaroi* de Marrocos e *Dollosuchus dixoni* de Bélxica. Estes primeiros tomistominos habitaban no océano de Tetis, que cubría gran parte de Europa e o norte de África no Paleoxeno. Varios tomistominos temperáns atopáronse en depósitos mariños costeiros, o que suxire que vivían ao longo da liña de costa ou en estuarios. Os gaviaoloides pénsase que eran animais costeiros. Os estilos de vida mariños destas primeiras formas probablemente permitiron aos tomistominos espallarse arredor do mar de Tetis, formando unha poboación do norte en Europa e outra do sur no norte de África.

Posteriormente no Eoceno e Oligoceno, os tomistominos estendéronse por Asia. As especies do Eoceno medio *Ferganosuchus planus* e *Dollosuchus zajsanicus* atopáronse en Casaquistán e Kirguizistán. Os tomistominos chegaron a China e Taiwán coa especie de final do Eocene *Maomingosuchus petrolica* e a especie do Mioceno *Penghusuchus pani*. Unha especie, *"Tomistoma" tandoni*, viviu na India durante o Eoceno medio. Durante este tempo, o subcontinente Indio estaba separado da terra continental de Asia, o que era unha barreira para as especies que non podían tolerar a auga de mar. O mar de Obik, que separaba Europa de Asia, tamén impedía unha fácil viaxe. Os tomistominos podían cruzar estas áreas, o que indica que tiñan tolerancia pola auga salgada.

Os tomistominos cruzaron o océano Atlántico e espalláronse polas Américas no Oligoceno, Mioceno e Plioceno. O tomistomino neotropical máis antigo coñecido é *Charactosuchus kuleri* de Xamaica. Propúxose que hai unha estreita relación entre *C. kuleri* e *D. zajsanicus* de Bélxica, suxerindo que os tomistominos migraron desde Europa ás Américas a través da ponte de terra de De Geer, que conectaba Noruega e Groenlandia, e a terra continental de Norteamérica ou a ponte de terra de Thule que conectaba a Escocia, Islandia, Groenlandia e Norteamérica. O xénero de *Thecachampsa* estaba presente ao longo da costa leste de Norteamérica durante o Oligoceno, Mioceno e Plioceno.

*Rhamphosuchus crassidens*, un tomistomino xigante da India.
Os tomistominos desapareceron de Europa durante o Oligoceno pero regresaron ao final desa época. Diversificáronse e fixéronse comúns no Mioceno medio. Un tomistomino, *Tomistoma coppensi*, é coñecido desde o Mioceno tardío de Uganda. A aparición dos tomistominos en África central é pouco usual, xa que hai poucas evidencias de especies do Mioceno tardío no norte de África, unha área á que deberon viaxar desde Europa.

Os tomistominos puideron viaxar de África á Asia cando Arabia chocou co continente eurasiático no Mioceno temperán. Porén, os tomistominos do Mioceno de Asia puideron tamén ser descendentes dos tomistominos do Eoceno que estaban xa presentes no leste de Asia. Os tomistominos espalláronse polo subcontinente Indio durante esa época. Unha especie, *Rhamphosuchus crassidens*, foi un dos crocodilos máis grandes que nunca existiu, que medraba ata uns estimados 8 a 11 m. Novas especies como *Toyotamaphimeia machikanensis* estaban presentes no Xapón no Plistoceno. Porén, no sueste de Asia, hai poucas evidencias fósiles dos tomistominos que precederon ao falso gavial. Por tanto, a súa relación coas especies extintas non está clara.

Filoxenia
---------

|  |
| --- |
| **Hipóteses da filoxenia dos tomistominos** |
| 

|  |
| --- |
| Morfolóxica |
| 

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| Crocodylia  | 

|  |  |
| --- | --- |
|   | Gavialidae  |
|   |
|   | 

|  |  |
| --- | --- |
|   | Alligatoridae  |
|   |
|  Crocodylidae  | 

|  |  |
| --- | --- |
|   | Crocodylinae  |
|   |
|   | **Tomistominae**  |
|   |

 |
|   |

 |
|   |

 |
|   |

 |

 |
| 

|  |
| --- |
| **Molecular** |
| 

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| Crocodylia  | 

|  |  |
| --- | --- |
|   | Alligatoridae  |
|   |
|   | 

|  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- |
|  Gavialoidea  | 

|  |  |
| --- | --- |
|   | *Gavialis*  |
|   |
|   | **Tomistominae**  |
|   |

 |
|   |
|   | Crocodylidae  |
|   |

 |
|   |

 |
|   |

 |

 |

O falso gavial *Tomistoma* é similar ao verdadeiro gavial *Gavialis* no seu fociño lonxirrostrino, que é longo e estreito. Aínda que os crocodilos son tamén lonxirrostrinos, os seus fociños non son tan estreitos coma os de *Tomistoma* e *Gavialis*. Certos detalles da morfoloxía de *Tomistoma* e *Gavialis* suxiren que ambos está só distantemente relacionados e que os fociños longos evolucionaron independentemente en cada liñaxe como resultado de evolución converxente. Morfoloxicamente, os tomistominos parecen estar máis estreitamente relacionados cos crocodílidos, polo que poderían compartir un antepasado común máis recente cos crocodilos que co gavial. Nesta filoxenia, os gaviais separáronse do antepasado común dos tomistominos, crocodilos e aligátores hai uns 66 millóns de anos no Cretáceo tardío. Velaquí un cladograma de Piras *et al.* (2007) que sitúa os tomistominos dentro dos Crocodyloidea:


|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| Crocodylia  | 

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

|  |  |
| --- | --- |
|   | *Borealosuchus* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Pristichampsus* |
|   |
|   | 

|  |  |
| --- | --- |
|   | Alligatoroidea |
|   |
|  Crocodyloidea  | 

|  |  |
| --- | --- |
|   | *Prodiplocynodon* |
|   |
|   | *Asiatosuchus* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Brachyuranochampsa* |
|   |
|  Crocodylidae  | 

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

|  |  |
| --- | --- |
|   | *"Crocodylus" megarhinus* |
|   |
|   | Mekosuchinae |
|   |
|   | 

|  |  |
| --- | --- |
|   | Osteolaeminae |
|   |
|   | *Crocodylus* |
|   |

 |
|   |

 |
|   |
|  **Tomistominae** | 

|  |  |
| --- | --- |
|   | *Kentisuchus* |
|   |
|   | 

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

|  |  |
| --- | --- |
|   | *Tomistoma schlegelii* |
|   |
|   | *Toyotamaphimeia* |
|   |

 |
|   |
|   | *Gavialosuchus* |
|   |
|   | *Paratomistoma* |
|   |
|   | *Megadontosuchus* |
|   |
|   | *Dollosuchus* |
|   |

 |
|   |

 |
|   |

 |
|   |

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|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |
|  Gavialoidea  | 

|  |  |
| --- | --- |
|   | *Thoracosaurus* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Eogavialis* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Gryposuchus* |
|   |
|   | *Gavialis* |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |

Non obstante, recentes análises moleculares dos crocodilianos atoparon unha estreita relación entre *Tomistoma* e *Gavialis*. Segundo esta filoxenia, as dúas formas son taxons irmáns, máis estreitamente relacionados entre si que con calquera crocodiliano vivo. Se son taxons irmáns, a diverxencia das dúas formas crese que ocorreu no Eoceno ou Mioceno medio. Se *Tomistoma* e *Gavialis* se considera que están estreitamente relacionados, as semellanzas morfolóxicas entre *Tomistoma* e os crocodilos deben ser unha converxencia. Nese caso, a converxencia orixina un problema coñecido como atracción das ramas longas, onde características similares pero que evolucionaron independentemente en *Tomistoma* e crocodilos implican erradamente que estes dous grupos moi derivados están estreitamente relacionados. De feito, estes caracteres compartidos enmascaran as relacións próximas entre *Tomistoma* e *Gavialis*.

A atracción das ramas longas pode resolverse se se observan diferenzas entre os membros ancestrais de cada grupo. Se os dous grupos estaban moi relacionados, os membros temperáns deberían ter trazos primitivos do antepasado común e deberían carecer de trazos derivados que causan a atracción das ramas longas. Desgrazadamente, os fósiles de tomistominos non resolven a atracción das ramas longas cando os seus caracteres se engaden ás análises filoxenéticas, polo que aínda permanece o conflito entre as filoxenias morfolóxicas e as moleculares.

Algúns estudos combinaron caracteres morfolóxicos e moleculares nas súas análises. As análises que inclúen caracteres morfolóxicos pero aínda dependen fortemente das secuencias de ADN encontran unha estreita relación entre *Tomistoma* e *Gavialis*. As análises que usan máis caracteres dos ósos e fósiles sosteñen a relación estreita de *Tomistoma* cos crocodilos.

### Relacións dentro de Tomistominae

Velaquí un cladograma de Shan *et al.* (2009):


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| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| **Crocodyloidea** | 

|  |  |
| --- | --- |
|   | *"Asiatosuchus" germanicus* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Prodiplocynodon* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *"Crocodylus" affinis* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Asiatosuchus grangeri* |
|   |
|   | 

|  |  |
| --- | --- |
|   | Dormaal crocodyloid |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Brachyuranochampsa* |
|   |
|   | *"Crocodylus" acer* |
|   |
|   | 

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
|  **Tomistominae** | 

|  |  |
| --- | --- |
|   | *Kentisuchus* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Megadontosuchus* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Dollosuchoides* |
|   |
|   | *"Gavialosuchus" americanus* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *"Tomistoma" cairense* |
|   |
|   | 

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

|  |  |
| --- | --- |
|   | *Paratomistoma* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Toyotamaphimeia* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Penghusuchus* |
|   |
|   | *Maomingosuchus petrolica* |
|   |

 |
|   |

 |
|   |

 |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Gavialosuchus eggenburgensis* |
|   |
|   | *Tomistoma lusitanica* |
|   |
|   | *Tomistoma schlegelii* |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |
|   | 

|  |  |
| --- | --- |
|   | Mekosuchinae |
|   |
|   | 

|  |  |
| --- | --- |
|   | *"Crocodylus" megarhinus* |
|   |
|   | 

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

|  |  |
| --- | --- |
|   | *Mecistops* |
|   |
|   | *Crocodylus* |
|   |

 |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Rimasuchus* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Voay* |
|   |
|   | *Osteolaemus* |
|   |

 |
|   |

 |
|   |

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|   |

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|   |

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|   |

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|   |

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|   |

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|   |

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|   |

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|   |

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|   |

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|   |

O cladograma seguinte é o da análise de Christopher A. Brochu e Glenn W. Storrs (2012).


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|  Crocodyloidea  | 

|  |  |
| --- | --- |
|   | *"Asiatosuchus" germanicus* |
|   |
|   | *Prodiplocynodon langi* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Asiatosuchus grangeri* |
|   |
|   | *"Crocodylus" affinis* |
|   |
|   | *"Crocodylus" depressifrons* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Brachyuranochampsa eversolei* |
|   |
|   | *"Crocodylus" acer* |
|   |
|  Crocodylidae  | 

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|  **Tomistominae**  | 

|  |  |
| --- | --- |
|   | *Kentisuchus spenceri* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Dollosuchoides densmorei* |
|   |
|   | *Megadontosuchus arduini* |
|   |
|   | 

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

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

|  |  |
| --- | --- |
|   | *Gavialosuchus eggenburgensis* |
|   |
|   | *Toyotamaphimeia machikanensis* |
|   |

 |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Tomistoma lusitanica* |
|   |
|   | *Tomistoma schlegelii* |
|   |

 |
|   |

 |
|   |
|   | 

|  |  |
| --- | --- |
|   | *"Tomistoma" cairense* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Thecachampsa antiqua* |
|   |
|   | *Thecachampsa americana* |
|   |
|   | *Thecachampsa carolinense* |
|   |

 |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Penghusuchus pani* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Paratomistoma courti* |
|   |
|   | *Maomingosuchus petrolica* |
|   |

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|   |

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|   |

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|   |

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|   |

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|   |

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|   |
|  Crocodylinae  | 

|  |  |
| --- | --- |
|   | *"Crocodylus" megarhinus* |
|   |
|  Mekosuchinae  | 

|  |  |
| --- | --- |
|   | *Kambara implexidens* |
|   |
|   | *Australosuchus clarkae* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Trilophosuchus rackhami* |
|   |
|   | *Quinkana* |
|   |

 |
|   |

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|   |
|   | 

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

|  |  |
| --- | --- |
|   | *"Crocodylus" pigotti* |
|   |
|   | *"Crocodylus" gariepensis* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Euthecodon arambourgii* |
|   |
|   | *Euthecodon brumpti* |
|   |

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|   |

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|   |
|  Osteolaeminae  | 

|  |  |
| --- | --- |
|   | *Rimasuchus lloydi* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Voay robustus* |
|   |
|   | 

|  |  |
| --- | --- |
|   | *Osteolaemus osborni* |
|   |
|   | *Osteolaemus tetraspis* |
|   |

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|   |

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|   |

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|   |
|   | 

|  |  |
| --- | --- |
|   | *Mecistops cataphractus* |
|   |
|   | *Crocodylus spp.* |
|   |

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|   |

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Notas
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1. ↑  Gatesy, Jorge; Amato, G.; Norell, M.; DeSalle, R.; Hayashi, C. (2003). "Combined support for wholesale taxic atavism in gavialine crocodylians" (PDF). *Systematic Biology* **52** (3): 403–422. doi:10.1080/1063515035019703. Arquivado dende o orixinal (PDF) o 22 de xuño de 2019. Consultado o 03 de setembro de 2017.
2. 1 2  Willis, R. E.; McAliley, L. R.; Neeley, E. D.; Densmore Ld, L. D. (June 2007). "Evidence for placing the false gharial (*Tomistoma schlegelii*) into the family Gavialidae: Inferences from nuclear gene sequences". *Molecular Phylogenetics and Evolution* **43** (3): 787–794. PMID 17433721. doi:10.1016/j.ympev.2007.02.005.
3. 1 2 3 4 5 6 7  Piras, P.; Delfino, M.; Del Favero, L.; Kotsakis, T. (2007). "Phylogenetic position of the crocodylian *Megadontosuchus arduini* and tomistomine palaeobiogeography" (PDF). *Acta Palaeontologica Polonica* **52** (2): 315–328. Arquivado dende o orixinal (PDF) o 03 de marzo de 2016. Consultado o 03 de setembro de 2017.
4. 1 2  Shan, Hsi-yin; Wu, Xiao-chun; Cheng, Yen-nien; Sato, Tamaki (2009). "A new tomistomine (Crocodylia) from the Miocene of Taiwan". *Canadian Journal of Earth Sciences* **46** (7): 529–555. doi:10.1139/E09-036.
5. 1 2  Piras, P.; Colangelo, P.; Adams, D.C.; Buscalioni, A.; Cubo, J.; Kotsakis, T.; Meloro, C.; Raia, P. (2010). "The *Gavialis*-*Tomistoma* debate: the contribution of skull ontogenetic allometry and growth trajectories to the study of crocodylian relationships". *Evolution & Development* **12** (6): 568–579. doi:10.1111/j.1525-142X.2010.00442.x.
6. 1 2 3  Brochu, C. A. (2003). "Phylogenetic approaches toward crocodylian history" (PDF). *Annual Review of Earth and Planetary Sciences* **31**: 357–97. doi:10.1146/annurev.earth.31.100901.141308. Arquivado dende o orixinal (PDF) o 02 de abril de 2015. Consultado o 03 de setembro de 2017.
7. ↑  Brochu, C. A.; Storrs, G. W. (2012). "A giant crocodile from the Plio-Pleistocene of Kenya, the phylogenetic relationships of Neogene African crocodylines, and the antiquity of *Crocodylus* in Africa". *Journal of Vertebrate Paleontology* **32** (3): 587. doi:10.1080/02724634.2012.652324.

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| Control de autoridades | 
* Wd: Q2448703
* MAG: 2779212090

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| --- | --- |
| Identificadores taxonómicos | 
* EOL: 57690423
* Fossilworks: 63118
* NCBI: 229546

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| Identificadores | 
* UMLS: C3334885

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