Tomistoma

Last updated

Tomistoma
Temporal range: Eocene - Recent, 47.8–0  Ma [1]
Tomistoma schlegelii false gharial LA zoo 01.jpg
False gharial (Tomistoma schlegelii)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Clade: Archosauriformes
Order: Crocodilia
Family: Gavialidae
Subfamily: Tomistominae
Genus: Tomistoma
S. Müller, 1846
Species

Tomistoma is a genus of gavialid crocodilians. They are noted for their long narrow snouts used to catch fish, similar to the gharial. Tomistoma contains one extant (living) member, the false gharial (Tomistoma schlegelii), as well as potentially several extinct species: T. cairense, T. lusitanicumT. coppensi, and T. dowsoni. However, these species may need to be reclassified to different genera as studies have shown them to be paraphyletic, for example: previously assigned species T. taiwanicus from Taiwan, is reclassified to the genus Toyotamaphimeia , and T. dowsoni should be excluded from Tomistoma based on phylogenetic analysis. [2] [3] [4] [5] [6]

Unlike the gharial, the false gharial's snout broadens considerably towards the base and so is more similar to those of true crocodiles than the gharial, whose osteology indicated a distinct lineage from all other living crocodilians. [7] However, although more morphologically similar to Crocodylidae based on skeletal features, recent molecular studies using DNA sequencing consistently indicate that the false gharial and by inference other related extinct forms traditionally viewed as belonging to the crocodylian subfamily Tomistominae actually belong to Gavialoidea and Gavialidae. [8] [9] [10] [11] [12] [13] [2] [14]

Fossil dorsal plates of "Tomistoma" calaritanum Gavialidae - Tomistoma calaritanus.JPG
Fossil dorsal plates of "Tomistoma" calaritanum

Fossils of extinct Tomistoma species have been found in deposits of Paleogene, Neogene, and Quaternary ages in Uganda, Italy, Portugal, Egypt and India, but nearly all of them are likely to be distinct genera due to older age compared to the false gharial. [15]

The below cladogram of the major living crocodile groups is based on molecular studies and shows the false gharial's close relationships: [9] [12] [13] [2] [14]

Crocodilia
Alligatoridae
Caimaninae

Caiman Caiman crocodilus llanos white background.JPG

Melanosuchus Melanosuchus niger white background.jpg

Paleosuchus Dwarf Caiman white background.jpg

Alligatorinae

Alligator Alligator white background.jpg

Longirostres
Crocodylidae

Crocodylus Siamese Crocodile white background.jpg

Mecistops Crocodylus cataphractus faux-gavial d'Afrique2 white background.JPG

Osteolaemus Bristol.zoo.westafrican.dwarf.croc.arp. white background.jpg

Gavialidae

Gavialis Gavialis gangeticus (Gharial, Gavial) white background.jpg

Tomistoma Tomistoma schlegelii. white background.JPG

Here is a more detailed cladogram from a 2018 tip dating study by Lee & Yates simultaneously using morphological, molecular (DNA sequencing), and stratigraphic (fossil age) data that shows the false gharial's proposed placement within Gavialidae, including extinct members: [2]

Gavialoidea
Gavialidae

Gavialis gangeticus Gharial

Gavialis bengawanicus

Gavialis browni

Gryposuchus colombianus

Ikanogavialis

Gryposuchus pachakamue

Piscogavialis

Harpacochampsa

Toyotamaphimeia

Penghusuchus

Gavialosuchus

Tomistoma lusitanicum

Tomistoma schlegelii False gharial

(crown group)

Tomistoma cairense

Dollosuchoides

Maroccosuchus

Paratomistoma

Kentisuchus

(stem-based group)

Related Research Articles

<span class="mw-page-title-main">Gavialinae</span> Subfamily of gharial crocodylians

Gavialinae is a subfamily of large semiaquatic crocodilian reptiles, resembling crocodiles, but with much thinner snouts. Gavialinae is one of the two major subfamilies within the family Gavialidae - the other being the subfamily Tomistominae, which contains the false gharial and extinct relatives.

<span class="mw-page-title-main">Gavialidae</span> Family of gharial crocodylians

Gavialidae is a family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two living species, the gharial and the false gharial, both occurring in Asia. Many extinct members are known from a broader range, including the recently extinct Hanyusuchus. Gavialids are generally regarded as lacking the jaw strength to capture the large mammalian prey favoured by crocodiles and alligators of similar size so their thin snout is best used to catch fish, however the false gharial has been found to have a generalist diet with mature adults preying upon larger vertebrates, such as ungulates.

<i>Gavialis</i> Genus of reptiles

Gavialis is a genus of crocodylians that includes the living gharial Gavialis gangeticus and one known extinct species, Gavialis bengawanicus.G. gangeticus comes from the Indian Subcontinent, while G. bengawanicus is known from Java. Gavialis likely first appeared in the Indian Subcontinent in the Pliocene and dispersed into the Malay Archipelago through a path called the Siva–Malayan route in the Quaternary. Remains attributed to Gavialis have also been found on Sulawesi and Woodlark Island east of the Wallace Line, suggesting a prehistoric lineage of Gavialis was able to traverse marine environments and reach places possibly as far as western Oceania.

<span class="mw-page-title-main">False gharial</span> Species of crocodilian

The false gharial, also known by the names Malayan gharial, Sunda gharial and tomistoma, is a freshwater crocodilian of the family Gavialidae native to Peninsular Malaysia, Borneo, Sumatra and Java. It is listed as Endangered on the IUCN Red List, as the global population is estimated at around 2,500 to 10,000 mature individuals.

<i>Toyotamaphimeia</i> Extinct genus of reptiles

Toyotamaphimeia is a genus of extinct gavialid crocodylian which lived in Japan and Taiwan during the Pleistocene. A specimen recovered in 1964 at Osaka University during the construction of a new science building has been dated to around 430–380 thousand years old based on the stratum in which it was found. Toyotamaphimeia was a fairly large crocodylian measuring approximately 6.3–7.3 metres (21–24 ft) long. Two species are named, T. machikanensis from Japan and T. taiwanicus from Taiwan, both originally described as members of the genus Tomistoma.

<i>Gavialosuchus</i> Extinct genus of reptiles

Gavialosuchus is an extinct genus of gavialoid crocodylian from the early Miocene of Europe. Currently only one species is recognized, as a few other species of Gavialosuchus have since been reclassified to other genera.

Dollosuchoides, colloquially known as the Crocodile of Maransart, is an extinct monospecific genus of gavialoid crocodilian, traditionally regarded as a member of the subfamily Tomistominae. Fossils have been found in the Brussel Formation of Maransart, Belgium and date back to the middle Eocene.

Eogavialis is an extinct genus of eusuchian crocodylomorph, usually regarded as a gavialoid crocodylian. It superficially resembles Tomistoma schlegelii, the extant false gharial, and consequently material from the genus was originally referred to Tomistoma. Indeed, it was not until 1982 that the name Eogavialis was constructed after it was realised that the specimens were from a more basal form.

<i>Kentisuchus</i> Extinct genus of reptiles

Kentisuchus is an extinct genus of gavialoid crocodylian, traditionally regarded as a member of the subfamily Tomistominae. Fossils have been found from England and France that date back to the early Eocene. The genus has also been recorded from Ukraine, but it unclear whether specimens from Ukraine are referable to Kentisuchus.

Maroccosuchus zennaroi is an extinct gavialoid crocodylian from the Early Eocene of Morocco, traditionally regarded as a member of the subfamily Tomistominae.

Paratomistoma is an extinct monospecific genus of gavialoid crocodylian. It is based on the holotype specimen CGM 42188, a partial posterior skull and lower jaw discovered at Wadi Hitan, Egypt, in Middle Eocene-age rocks of the Gehannam Formation. The skull is unfused but considered morphologically mature. Paratomistoma was named in 2000 by Christopher Brochu and Philip Gingerich; the type species is P. courti in honor of Nicholas Court, who found CGM 42188. They performed a phylogenetic analysis and found Paratomistoma to be a derived member of Tomistominae, related to the false gharial. It may have been a marine or coastal crocodilian.

<i>Planocrania</i> Extinct genus of reptiles

Planocrania is an extinct genus of eusuchian crocodyliforms from what is now China. Two species are currently known to belong to the genus.

<i>Penghusuchus</i> Extinct genus of reptiles

Penghusuchus is an extinct genus of gavialid crocodylian. It is known from a skeleton found in Middle to Upper Miocene rocks of Penghu Island, off Taiwan. The taxon was described in 2009 by Shan and colleagues; the type species is P. pani. It may be related to two other fossil Asian gavialids: Toyotamaphimeia machikanensis of Japan and Hanyusuchus sinensis of South China. It was a medium-sized gavialid with an estimated total length of 4.5 metres (15 ft).

<span class="mw-page-title-main">Crocodyloidea</span> Superfamily of crocodiles

Crocodyloidea is one of three superfamilies of crocodilians, the other two being Alligatoroidea and Gavialoidea, and it includes the crocodiles. Crocodyloidea may also include the extinct Mekosuchinae, native to Australasia from the Eocene to the Holocene, although this is disputed.

Gryposuchinae is an extinct subfamily of gavialid crocodylians. Gryposuchines lived mainly in the Miocene of South America. However, "Ikanogavialis" papuensis may have survived more recently, into the Late Pleistocene/Holocene. Most were long-snouted coastal forms. The group was named in 2007 and includes genera such as Gryposuchus and Aktiogavialis, although a 2018 study indicates that the group might be paraphyletic and rather an evolutionary grade towards the gharial.

<span class="mw-page-title-main">Gavialoidea</span> Superfamily of large reptiles

Gavialoidea is one of three superfamilies of crocodylians, the other two being Alligatoroidea and Crocodyloidea. Although many extinct species are known, only the gharial Gavialis gangeticus and the false gharial Tomistoma schlegelii are alive today, with Hanyusuchus having become extinct in the last few centuries.

<span class="mw-page-title-main">Tomistominae</span> Subfamily of reptiles

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.

<span class="mw-page-title-main">Brevirostres</span> Taxon of reptiles

Brevirostres is a paraphyletic group of crocodilians that included alligatoroids and crocodyloids. Brevirostres are crocodilians with small snouts, and are distinguished from the long-snouted gharials. It is defined phylogenetically as the last common ancestor of Alligator mississippiensis and Crocodylus niloticus and all of its descendants. This classification was based on morphological studies primarily focused on analyzing skeletal traits of living and extinct fossil species, and placed the gharials outside the group due to their unique skull structure, and can be shown in the simplified cladogram below:

Tomistoma cairense is an extinct species of gavialoid crocodilian from the Lutetian stage of the Eocene era. It lived in North East Africa, especially Egypt. Remains of T. cairense have been found in the Mokattam Formation, in Mokattam, Egypt. Tomistoma cairense did not have a Maxilla process within their lacrimal gland, whereas all extant (living) crocodilians do.

<span class="mw-page-title-main">Longirostres</span> Clade of crocodilians

Longirostres is a clade of crocodilians that includes the crocodiles and the gavialids, to the exclusion of the alligatoroids. Defined in 2003 by Harshman et al., Longirostres is a crown group defined phylogenetically as including the last common ancestor of Crocodylus niloticus and Gavialis gangeticus and all of its descendants.

References

  1. Rio, Jonathan P.; Mannion, Philip D. (6 September 2021). "Phylogenetic analysis of a new morphological dataset elucidates the evolutionary history of Crocodylia and resolves the long-standing gharial problem". PeerJ . 9: e12094. doi: 10.7717/peerj.12094 . PMC   8428266 . PMID   34567843.
  2. 1 2 3 4 Lee, M. S. Y.; Yates, A. M. (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.
  3. Iijima, M.; Momohara, A.; Kobayashi, Y.; Hayashi, S.; Ikeda, T.; Taruno, H.; Watanabe, K.; Tanimoto, M.; Furui, S. (2018). "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.
  4. Ito, Ai; Aoki, Riosuke; Hirayama, Ren; Yoshida, Masataka; Kon, Hiroo; Endo, Hideki (April 2018). "The Rediscovery and Taxonomical Reexamination of the Longirostrine Crocodylian from the Pleistocene of Taiwan". Paleontological Research. 22 (2): 150–155. doi:10.2517/2017PR016. ISSN   1342-8144. S2CID   134961600.
  5. Burke, Paul M. J.; Mannion, Philip D. (2023-03-16). "Neuroanatomy of the crocodylian Tomistoma dowsoni from the Miocene of North Africa provides insights into the evolutionary history of gavialoids". Journal of Anatomy. 243 (1): 1–22. doi:10.1111/joa.13846. ISSN   0021-8782. PMC   10273334 . PMID   36929596.
  6. Cho, Yi-Yang; Tsai, Cheng-Hsiu (2023-06-29). "Crocodylian princess in Taiwan: Revising the taxonomic status of Tomistoma taiwanicus from the Pleistocene of Taiwan and its paleobiogeographic implications". Journal of Paleontology. 97 (4): 927–940. Bibcode:2023JPal...97..927C. doi: 10.1017/jpa.2023.36 . ISSN   0022-3360.
  7. Piras, P., Colangelo, P., Adams, D. C., Buscalioni, A., Cubo, J., Kotsakis, T., & 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.
  8. Densmore, L. D.; Owen, R. D. (1989). "Molecular Systematics of the Order Crocodilia". American Zoologist. 29 (3): 831–841. doi: 10.1093/icb/29.3.831 .
  9. 1 2 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". Systematic Biology. 52 (3): 386–402. doi: 10.1080/10635150309323 . PMID   12775527.
  10. Gatesy, J.; Amato, G.; Norell, M.; DeSalle, R.; H. C. (2003). "Combined support for wholesale taxic atavism in gavialine crocodylians". Systematic Biology. 52 (3): 403–422. doi: 10.1080/10635150309329 . PMID   12775528.
  11. Willis, R. E.; McAliley, L. R.; Neeley, E. D.; Densmore Ld, L. D. (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.
  12. 1 2 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.
  13. 1 2 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). "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.
  14. 1 2 Hekkala, E.; Gatesy, J.; Narechania, A.; Meredith, R.; Russello, M.; Aardema, M. L.; Jensen, E.; Montanari, S.; Brochu, C.; Norell, M.; Amato, G. (2021). "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 . PMC   8079395 . PMID   33907305.
  15. "Fossilworks: Tomistoma". fossilworks.org.