Tomistominae

Tomistominae is a subfamily of crocodylians that includes one living species, the false gharial. Many more extinct species are known, extending the range of the subfamily back to the Eocene 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.

Tomistominae
Temporal range: Eocene - Recent,
False gharial, Tomistoma schlegelii
Scientific classification Edit this classification
(disputed)
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Crocodilia
Clade: Longirostres
Superfamily: Gavialoidea
Family: Gavialidae
Subfamily: Tomistominae
Kälin, 1955
Genera

The classification of tomistomines among Crocodylia has been in flux; while traditionally thought to be within Crocodyloidea, molecular evidence indicates that they are more closely related to true gharials as members of Gavialoidea.

Description

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 toward piscivory, 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. The splenial bone of the lower jaw is long and slender, forming a distinctive "V" shape not seen in gharials.

Evolutionary history

Skull of Kentisuchus toliapicus.

Tomistomines first appeared in the Eocene in Europe and North Africa. The oldest known tomistomine is Kentisuchus spenceri from England, although a possible tomistomine fossil from the Paleocene of Spain is even older.[1] Other early tomistomines include Maroccosuchus zennaroi from Morocco and Dollosuchus dixoni from Belgium. These early tomistomines inhabited the Tethys 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 and Oligocene, tomistomines spread across Asia. The middle Eocene species Ferganosuchus planus and Dollosuchus zajsanicus are known from Kazakhstan and Kyrgyzstan. Tomistomines reached China and Taiwan with the late Eocene species Maomingosuchus petrolica and the Miocene species Penghusuchus pani.[2] One species, "Tomistoma" tandoni, lived in India 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. The Obik 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]

Rhamphosuchus crassidens, a giant tomistomine from India.

Tomistomines crossed the Atlantic Ocean and spread into the Americas in the Oligocene, Miocene, and Pliocene. The earliest known neotropical tomistomine is Charactosuchus kuleri from Jamaica. A close relationship has been proposed between C. kuleri and D. zajsanicus from Belgium, suggesting that tomistomines migrated from Europe to the Americas through the De Geer land bridge connecting Norway to Greenland and the North American mainland or the Thule land bridge connecting Scotland, Iceland, Greenland, and the North American mainland. The genus Thecachampsa 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 of Uganda. 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 when Arabia 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 as Toyotamaphimeia machikanensis were present in Japan in the Pleistocene. 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]

Phylogeny

Tomistominae is cladistically defined as Tomistoma schlegelii (the false gharial) and all species closer to it than to Gavialis gangeticus (the gharial) or Crocodylus rhombifer (the Cuban crocodile).[3][4] This is a stem-based definition for tomistominae, and means that it includes more basal extinct tomistomine ancestors that are more closely related to the false gharial than to the gharial or crocodiles.

Hypotheses of tomistomine phylogeny
Morphological
Crocodylia 

Gavialidae

Alligatoridae

 Crocodylidae 

Crocodylinae

Tomistominae

Molecular
Crocodylia 

Alligatoridae

 Gavialoidea 

Gavialis

Tomistominae

Crocodylidae

Below is a cladogram based morphological studies comparing skeletal features that shows the members of Tomistominae as belonging to Crocodylidae:[5]

Crocodylidae

Crocodylinae

Tomistominae

Xaymacachampsa

Megadontosuchus

Kentisuchus

Maroccosuchus

Dollosuchoides

Thecachampsa

Penghusuchus

Toyotamaphimeia

Tomistoma cairense

Maomingosuchus

Tomistoma schlegelii False gharial

Gavialosuchus

Tomistoma lusitanica

Paratomistoma

Tomistoma coppensi


Based on morphological studies of extinct taxa, the tomistomines (including the living false gharial) were long thought to be classified as crocodiles and not closely related to gavialoids.[6] However, recent molecular studies using DNA sequencing have consistently indicated that the false gharial (Tomistoma) (and by inference other related extinct forms in Tomistominae) actually belong to Gavialoidea (and Gavialidae).[7][4][8][9][10][11][12]

Below is a cladogram from a 2018 tip dating study by Lee & Yates simultaneously using morphological, molecular (DNA sequencing), and stratigraphic (fossil age) data that shows the tomistomines belonging to Gavialidae, and that the members traditionally belonging to Tomistominae may in fact be paraphyletic with respect to the gharial:[11]

Longirostres

Crocodyloidea/Crocodylidae

Gavialoidea
Gavialidae
(Gavialinae?)

Gavialis gangeticus Gharial

Gavialis bengawanicus

Gavialis browni

Gryposuchus colombianus

Ikanogavialis

Gryposuchus pachakamue

Piscogavialis

Harpacochampsa

Toyotamaphimeia

Penghusuchus

Gavialosuchus

(stem-based group)
(Tomistominae?)

Tomistoma lusitanica

Tomistoma schlegelii False gharial

(stem-based group)
(crown group)

Tomistoma cairense

Dollosuchoides

Maroccosuchus

Paratomistoma

Kentisuchus

(stem-based group)
Traditional Tomistominae
(crown group)

References

  1. 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.
  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. Bibcode:2009CaJES..46..529S. doi:10.1139/E09-036.
  3. Brochu, C. A. (2003). "Phylogenetic approaches toward crocodylian history" (PDF). Annual Review of Earth and Planetary Sciences. 31 (31): 357–97. Bibcode:2003AREPS..31..357B. doi:10.1146/annurev.earth.31.100901.141308. Archived from the original (PDF) on 2015-04-02. Retrieved 2011-06-25.
  4. 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:10.1080/10635150309329. PMID 12775528.
  5. Iijima, Masaya; Momohara, Arata; Kobayashi, Yoshitsugu; Hayashi, Shoji; Ikeda, Tadahiro; Taruno, Hiroyuki; Watanabe, Katsunori; Tanimoto, Masahiro; Furui, Sora (2018-05-01). "Toyotamaphimeia cf. machikanensis (Crocodylia, Tomistominae) from the Middle Pleistocene of Osaka, Japan, and crocodylian survivorship through the Pliocene-Pleistocene climatic oscillations". Palaeogeography, Palaeoclimatology, Palaeoecology. 496: 346–360. Bibcode:2018PPP...496..346I. doi:10.1016/j.palaeo.2018.02.002. ISSN 0031-0182.
  6. Brochu, C.A.; Gingerich, P.D. (2000). "New tomistomine crocodylian from the Middle Eocene (Bartonian) of Wadi Hitan, Fayum Province, Egypt". University of Michigan Contributions from the Museum of Paleontology. 30 (10): 251–268.
  7. Harshman, J.; Huddleston, C. J.; Bollback, J. P.; Parsons, T. J.; Braun, M. J. (2003). "True and false gharials: A nuclear gene phylogeny of crocodylia" (PDF). Systematic Biology. 52 (3): 386–402. doi:10.1080/10635150309323. PMID 12775527. Archived from the original (PDF) on 2022-10-09. Retrieved 2021-06-30.
  8. 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. doi:10.1016/j.ympev.2007.02.005. PMID 17433721.
  9. Gatesy, J.; Amato, G. (2008). "The rapid accumulation of consistent molecular support for intergeneric crocodylian relationships". Molecular Phylogenetics and Evolution. 48 (3): 1232–1237. doi:10.1016/j.ympev.2008.02.009. PMID 18372192.
  10. Erickson, G. M.; Gignac, P. M.; Steppan, S. J.; Lappin, A. K.; Vliet, K. A.; Brueggen, J. A.; Inouye, B. D.; Kledzik, D.; Webb, G. J. W. (2012). Claessens, Leon (ed.). "Insights into the ecology and evolutionary success of crocodilians revealed through bite-force and tooth-pressure experimentation". PLOS ONE. 7 (3): e31781. Bibcode:2012PLoSO...731781E. doi:10.1371/journal.pone.0031781. PMC 3303775. PMID 22431965.
  11. Michael S. Y. Lee; Adam M. Yates (27 June 2018). "Tip-dating and homoplasy: reconciling the shallow molecular divergences of modern gharials with their long fossil". Proceedings of the Royal Society B. 285 (1881). doi:10.1098/rspb.2018.1071. PMC 6030529. PMID 30051855.
  12. Hekkala, E.; Gatesy, J.; Narechania, A.; Meredith, R.; Russello, M.; Aardema, M. L.; Jensen, E.; Montanari, S.; Brochu, C.; Norell, M.; Amato, G. (2021-04-27). "Paleogenomics illuminates the evolutionary history of the extinct Holocene "horned" crocodile of Madagascar, Voay robustus". Communications Biology. 4 (1): 505. doi:10.1038/s42003-021-02017-0. ISSN 2399-3642. PMC 8079395. PMID 33907305.
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