Tomistominae Temporal range:Eocene - Recent,55–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N
False gharial,Tomistoma schlegelii
Scientific classification (disputed)
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Clade:	Archosauria
Order:	Crocodilia
Clade:	Longirostres
Superfamily:	Gavialoidea
Family:	Gavialidae
Subfamily:	Tomistominae Kälin, 1955
Genera
Tomistoma †Dollosuchoides? †Dollosuchus? †Ferganosuchus? †Gavialosuchus? †Gunggamarandu †Kentisuchus? †Maomingosuchus? †Maroccosuchus? †Megadontosuchus? †Paratomistoma? †Penghusuchus? †Toyotamaphimeia? †Thecachampsa?

Tomistominae is asubfamily ofcrocodylians that includes one living species, thefalse gharial. Many more extinct species are known, extending the range of the subfamily back to theEocene epoch. In contrast to the false gharial, which is a freshwater species that lives only in southeast Asia, extinct tomistomines had a global distribution and lived in estuaries and along coastlines.

The classification of tomistomines among Crocodylia has been in flux; while traditionally thought to be withinCrocodyloidea, molecular evidence indicates that they are more closely related to truegharials as members ofGavialoidea.

Tomistomines have narrow or longirostrine snouts like gharials. The living false gharial lives in fresh water and uses its long snout and sharp teeth to catch fish, although true gharials are more adapted towardpiscivory, or fish-eating. Despite the similarity with gharials, the shapes of bones in tomistomine skulls link them with crocodiles. For example, both tomistomines and crocodiles have thin postorbital bars behind the eye sockets and a large socket for the fifth maxillary tooth. Thesplenial bone of the lower jaw is long and slender, forming a distinctive "V" shape not seen in gharials.

Tomistomines first appeared in the Eocene inEurope andNorth Africa. The oldest known tomistomine isKentisuchus spenceri fromEngland, although a possible tomistomine fossil from thePaleocene ofSpain is even older.^[1] Other early tomistomines includeMaroccosuchus zennaroi fromMorocco andDollosuchus dixoni fromBelgium. These early tomistomines inhabited theTethys Ocean, which covered much of Europe and North Africa during the Paleogene. Several early tomistomines are found in coastal marine deposits, suggesting that they lived along the shoreline or in estuaries. Extinct gavialoids are also thought to have been coastal animals. The marine lifestyles of these early forms likely allowed tomistomines to spread around the Tethys, forming a northern population in Europe and a southern in North Africa.^[1]

Later in the Eocene andOligocene, tomistomines spread across Asia. The middle Eocene speciesFerganosuchus planus andDollosuchus zajsanicus are known fromKazakhstan andKyrgyzstan. Tomistomines reached China and Taiwan with the late Eocene speciesMaomingosuchus petrolica and theMiocene speciesPenghusuchus pani.^[2] One species,"Tomistoma" tandoni, lived inIndia during the middle Eocene. During this time, the Indian subcontinent was separated from mainland Asia, creating a barrier to species that could not tolerate salt water. TheObik Sea, which separated Europe from Asia, also impeded travel. Tomistomines were able to cross these areas, indicating that they had tolerance to salt water.^[1]

Tomistomines crossed the Atlantic Ocean and spread into the Americas in the Oligocene, Miocene, andPliocene. The earliest knownneotropical tomistomine isCharactosuchus kuleri fromJamaica. A close relationship has been proposed betweenC. kuleri andD. zajsanicus from Belgium, suggesting that tomistomines migrated from Europe to the Americas through theDe Geerland bridge connectingNorway toGreenland and the North American mainland or theThule land bridge connecting Scotland, Iceland, Greenland, and the North American mainland. The genusThecachampsa was present along the eastern coast of North America during the Oligocene, Miocene, and Pliocene.^[1]

Tomistomines disappeared from Europe during the Oligocene but returned by the end of the epoch. They diversified and became common in the middle Miocene. One tomistomine,Tomistoma coppensi, is known from the late Miocene ofUganda. The appearance of tomistomines in central Africa is unusual because there is little evidence of late Miocene species in North Africa, an area where they must have traveled through from Europe.^[1]

Tomistomines may have traveled from Africa into Asia whenArabia collided with the Eurasian continent in the Early Miocene. However, Asian Miocene tomistomines may also have descended from the Eocene tomistomines that were already present in eastern Asia. Tomistomines spread throughout the Indian subcontinent during this time. One species,Rhamphosuchus crassidens, was one of the largest crocodilians that ever lived, growing to an estimated 8 to 11 metres (26 to 36 ft). New species such asToyotamaphimeia machikanensis were present in Japan in thePleistocene. In southeast Asia however, there is little fossil evidence of the tomistomines that preceded the false gharial. Therefore, its relation with extinct species is unclear.^[1]

Tomistominae iscladistically defined asTomistoma schlegelii (thefalse gharial) and all species closer to it than toGavialis gangeticus (thegharial) orCrocodylus rhombifer (theCuban crocodile).^[3][4] This is astem-based definition for tomistominae, and means that it includes morebasalextinct tomistomine ancestors that are more closely related to the false gharial than to thegharial orcrocodiles.

Below is acladogram basedmorphological studies comparing skeletal features that shows the members of Tomistominae as belonging toCrocodylidae:^[5]

Based on morphological studies ofextincttaxa, the tomistomines (including the livingfalse gharial) were long thought to be classified ascrocodiles and not closely related togavialoids.^[6] However, recent molecular studies usingDNA sequencing have consistently indicated that thefalse gharial (Tomistoma) (and by inference other related extinct forms in Tomistominae) actually belong toGavialoidea (andGavialidae).^{[7][4][8][9][10][11][12]}

Below is acladogram from a 2018tip dating study by Lee & Yates simultaneously usingmorphological, molecular (DNA sequencing), andstratigraphic (fossil age) data that shows the tomistomines belonging toGavialidae, and that the members traditionally belonging to Tomistominae may in fact beparaphyletic with respect to thegharial:^[11]

• Enigmatic crocodilian taxa
• Crocodilia
• Gavialidae
• Extant Eocene first appearances

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