Gavialidae

Last updated

Gavialidae
Temporal range: Eocene, 38–0  Ma
Indian Gharial Crocodile Digon3.JPG
Indian gharial, Gavialis gangeticus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Clade: Archosauriformes
Order: Crocodilia
Superfamily: Gavialoidea
Family: Gavialidae
Adams, 1854
Subfamilies

Gavialidae is a family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two living species, the gharial (Gavialis gangeticus) and the false gharial (Tomistoma schlegelii), 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. [1]

Contents

Taxonomy

The family Gavialidae was proposed by Arthur Adams in 1854 for reptiles with a very long and slender muzzle, webbed feet and nearly equal teeth. [2] It is currently recognized as a crown group, [3] meaning that it only includes the last common ancestor of all extant (living) gavialids (the gharial and false gharial) and their descendants (living or extinct).

Traditionally, crocodiles and alligators were considered more closely related and grouped together in the clade Brevirostres, to the exclusion of the gharials. This classification was based on morphological studies primarily focused on analyzing skeletal traits of living and extinct fossil species. [4] However, recent molecular studies using DNA sequencing have rejected Brevirostres upon finding the crocodiles and gavialids to be more closely related than the alligators. [5] [6] [7] [3] [8] The new clade Longirostres was named by Harshman et al. in 2003. [5]

In addition, these recent molecular DNA studies consistently indicate that the false gharial (Tomistoma) (and by inference other related extinct forms) traditionally viewed as belonging to the crocodylian subfamily Tomistominae actually belong to Gavialoidea (and Gavialidae). [5] [9] [10] [6] [7] [3] [8] As its name suggests, the false gharial was once thought to be only distantly related to the gharial despite its similar appearance. The false gharial and other tomistomines were traditionally classified within the superfamily Crocodyloidea as close relatives of crocodiles, based solely on morphological evidence. [9]

A 2018 tip dating study by Lee & Yates simultaneously using morphological, molecular (DNA sequencing), and stratigraphic (fossil age) data established the inter-relationships within Crocodilia, [3] which was expanded upon in 2021 by Hekkala et al. using paleogenomics by extracting DNA from the extinct Voay . [8] The tip dating analysis resolved the extinct Thoracosaurus and similar extinct close relatives ("thoracosaurs") as outside of Gavialoidea due to the large time difference. They concluded that the only possible explanation for the morphological data placing thoracosaurs within the gharial lineage was a significant amount of homoplastic convergence between thoracosaurs and Gavialis. [3]

The below cladogram from latest study shows Gavialidae's placement within Crocodylia:

Crocodylia
Alligatoroidea

extinct basal Alligatoroids

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

(crown group)
(stem-based group)
Longirostres
Crocodyloidea

"Crocodylus" megarhinus

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

(crown group)
(stem-based group)
Gavialoidea

Kentisuchus

Maroccosuchus

Paratomistoma

Dollosuchoides

Tomistoma cairense

Gavialidae

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

Tomistoma Tomistoma schlegelii. white background.JPG

(crown group)
(stem-based group)
(crown group)
(crown group)

Here is a more detailed cladogram that shows the proposed phylogeny of Gavialidae including extinct members: [11]

Crocodyloidea

Gavialoidea

Maroccosuchus zennaroi

Megadontosuchus arduini

Dollosuchoides densmorei

Xaymacachampsa kugleri

Kentisuchus astrei

Kentisuchus spenceri

Gavialidae

Tomistoma cairense

Tomistoma coppensi

Maomingosuchus petrolica

Tomistominae

Tomistoma schlegelii, false gharial

Tomistoma lusitanicum

Gavialosuchus eggenburgensis

Melitosaurus champsoides

Tomistoma calaritanum

Tomistoma gaudense

Thecachampsa carolinensis

Thecachampsa antiqua

Gavialinae

Paratomistoma courti

Penghusuchus pani

Toyotamaphimeia machikanensis

Hanyusuchus sinensis

Eosuchus lerichei

Eosuchus minor

Ocepesuchus eoafricanus

Eothoracosaurus mississippiensis

Thoracosaurus macrorhynchus

Thoracosaurus neocesariensis

Aktiogavialis puertoricensis

Eogavialis africanum

Argochampsa krebsi

Piscogavialis jugaliperforatus

Ikanogavialis gameroi

Siquisiquesuchus venezuelensis

Dadagavialis gunai

Gryposuchus neogaeus

Gryposuchus croizati

Aktiogavialis caribesi

Gryposuchus pachakamue

Gryposuchus colombianus

Rhamphosuchus crassidens

Myanmar gavialid

Gavialis lewisi

Gavialis bengawanicus

Gavialis gangeticus, gharial

Siwalik Gavialis

Species list

Family Gavialidae

† Indicates extinct group

Extant species

ImageScientific nameCommon NameDistribution
Indian Gharial Crocodile Digon3.JPG Gavialis gangeticus Gharial India
False Gharials (Tomistoma schlegelii) (6749856797).jpg Tomistoma schlegelii False gharial, Malayan gharial, Sunda gharial and tomistomaPeninsular Malaysia, Borneo, Sumatra, and Java.

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.

<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>Tomistoma</i> Genus of crocodilians

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, 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.

<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. taiwanica 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.

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

Alligatoroidea is one of three superfamilies of crocodylians, the other two being Crocodyloidea and Gavialoidea. Alligatoroidea evolved in the Late Cretaceous period, and consists of the alligators and caimans, as well as extinct members more closely related to the alligators than the two other groups.

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>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. Magnusson, W. E. (1998). Cogger, H. G.; Zweifel, R. G. (eds.). Encyclopedia of Reptiles and Amphibians. San Diego: Academic Press. pp. 229–230. ISBN   978-0-12-178560-4.
  2. Adams, A. (1854). "II. Order – Emydosaurians (Emydosauria)". In Adams, A.; Baikie, W. B.; Barron, C. (eds.). A Manual of Natural History, for the Use of Travellers: Being a Description of the Families of the Animal and Vegetable Kingdoms: with Remarks on the Practical Study of Geology and Meteorology. London: John Van Voorst. pp. 70–71.
  3. 1 2 3 4 5 6 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.
  4. Holliday, Casey M.; Gardner, Nicholas M. (2012). Farke, Andrew A (ed.). "A new eusuchian crocodyliform with novel cranial integument and its significance for the origin and evolution of Crocodylia". PLOS ONE. 7 (1): e30471. Bibcode:2012PLoSO...730471H. doi: 10.1371/journal.pone.0030471 . PMC   3269432 . PMID   22303441.
  5. 1 2 3 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.
  6. 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.
  7. 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). 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.
  8. 1 2 3 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.
  9. 1 2 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.
  10. 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.
  11. Iijima M, Qiao Y, Lin W, Peng Y, Yoneda M, Liu J (2022). "An intermediate crocodylian linking two extant gharials from the Bronze Age of China and its human-induced extinction". Proceedings of the Royal Society B: Biological Sciences. 289 (1970): Article ID 20220085. doi: 10.1098/rspb.2022.0085 . PMC   8905159 . PMID   35259993.