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Tomistoma schlegelii

Tomistoma schlegelii (*)

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Archosauromorpha
Cladus: Crurotarsi
Divisio: Archosauria
Subdivisio: Pseudosuchia
Superordo: Crocodylomorpha
Ordo: Crocodilia
Subordo: Eusuchia

Familia: Gavialidae
Subfamilia: Tomistominae
Genus: Tomistoma
Species: Tomistoma schlegelii
Name

Tomistoma schlegelii Müller, 1838

Type locality: Karau river and Lake Lamoeda on river Doeson, Southern Borneo

Holotype: Syntypes RMNH 39581, RMNH 35444, RMNH 35445, RMNH 7934, RMNH 7935, RMNH 35449, RMNH 3198, RMNH 3200
Synonyms

Crocodilus (Gavialis) schlegelii Müller, 1838
Tomistoma schlegelii — King & Burke, 1989
Tomistoma schlegelii — Cox et al., 1998
Tomistoma schlegelii — Chan-Ard et al., 1999

References

S. Müller: Waarnemingen over de Indische Krokodillen en Beschrijving Van Enne Nieuwe Soort. Tijdschrift voor Natuurlijke Geschiedenis en Physiologie. Amsterdam and Leyden. 5: 61–87, pl. 3. (1838)
S. Müller: Ueber den Character der Thierwelt auf den Inseln des indischen Archipels, ein Beitrag zur zoologischen Geographie. Archiv für Naturgeschichte 12(1): 109–128. (1846)
Tomistoma schlegelii at the New Reptile Database. Accessed on 16 August 2009.
IUCN: Tomistoma schlegelii Müller, 1838 (Endangered)
Tomistoma schlegelii Müller, 1838 – Taxon details on Integrated Taxonomic Information System (ITIS).

Vernacular names
Deutsch: Sunda-Gavial
English: False gharial
suomi: Sundagaviaali, valegaviaali
français: Faux-gavial de Malaisie
עברית: גאביאל חקיין
magyar: Szunda-krokodil
Bahasa Indonesia: Buaya sepit
日本語: マレーガビアル
Bahasa Melayu: Buaya Jejulong
polski: Krokodyl gawialowy, krokodyl sundajski
português: Falso-gavial, Gavial-da-malásia
русский: Псевдогавиал
ไทย: ตะโขง, จระเข้ปากกระทุงเหว
Tiếng Việt: Cá sấu Ấn Độ giả
中文: 馬來鱷

Tomistoma schlegelii

The false gharial (Tomistoma schlegelii), also known as the Malayan gharial, the Sunda gharial, and by the genus name Tomistoma,[3][4][5][6][7] is a freshwater crocodilian in the family Gavialidae. It is native to Peninsular Malaysia, Borneo, Sumatra, and Java. It is listed as Vulnerable on the IUCN Red List, as the global population is estimated at around 2,500 to 10,000 mature individuals.[2]

The specific name schlegelii honors German herpetologist Hermann Schlegel.[8][9]

Taxonomy

The false gharial belongs to the genus Tomistoma, which also potentially contains several extinct species: T. cairense, T. lusitanicum, T. taiwanicus, and T. coppensi. However, these species may need to be reclassified to different genera as studies have shown them to be paraphyletic.[10][11]

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.[12] 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.[13][14][15][16][17][18][10][19]
Fossil dorsal plates of "Tomistoma" calaritanus

Fossils of extinct Tomistoma species have been found in deposits of Paleogene, Neogene, and Quaternary ages in Taiwan, 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.[20]

The below cladogram of the major living crocodile groups is based on molecular studies and shows the false gharial's close relationships:[14][17][18][10][19]

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:[10]

Gavialoidea
Gavialidae

Gavialis gangeticus Gharial

Gavialis bengawanicus

Gavialis browni

Gryposuchus colombianus

Ikanogavialis

Gryposuchus pachakamue

Piscogavialis

Harpacochampsa

Toyotamaphimeia

Penghusuchus

Gavialosuchus

Tomistoma lusitanica

Tomistoma schlegelii False gharial

(crown group)

Tomistoma cairense

Dollosuchoides

Maroccosuchus

Paratomistoma

Kentisuchus

(stem-based group)

Characteristics
Close-up of a false gharial at the Tierpark Berlin
Skull of a false gharial at the Zoological Museum of the Zoological Institute of the Russian Academy of Sciences, St. Petersburg

The false gharial is dark reddish-brown above with dark brown or black spots and cross-bands on the back and tail. Ventrals are grayish-white, with some lateral dark mottling. Juveniles are mottled with black on the sides of the jaws, body, and tail. The smooth and unornamented snout is extremely long and slender, parallel sided, with a length of 3.0 to 3.5 times the width at the base. All teeth are long and needle-like, interlocking on the insides of the jaws, and are individually socketed. The dorsal scales are broad at midbody and extend onto the sides of the body. The digits are webbed at the base. Integumentary sensory organs are present on the head and body scalation. Scales behind the head are frequently a slightly enlarged single pair. Some individuals bear a number of adjoining small keeled scales. Scalation is divided medially by soft granular skin. Three transverse rows of two enlarged nuchal scales are continuous with the dorsal scales, which consist of 22 transverse rows of six to eight scales, are broad at midbody and extend onto the sides of the body. Nuchal and dorsal rows equals a total of 22 to 23 rows. It has 18 double-crested caudal whorls and 17 single-crested caudal whorls. The flanks have one or two longitudinal rows of six to eight very enlarged scales on each side.[21]

The false gharial has one of the slimmest snouts of any living crocodilian, comparable to that of the slender-snouted crocodile and the freshwater crocodile in slenderness; only that of the gharial is noticeably slimmer.[12] The false gharial is a large crocodilian. Males can reportedly grow up to 5 m (16 ft) in length.[22] Three mature males kept in captivity measured 3.6 to 3.9 m (11 ft 10 in to 12 ft 10 in) and weighed 190 to 210 kg (420 to 460 lb), while a female measured 3.27 m (10 ft 9 in) and weighed 93 kg (205 lb).[23] Females have been recorded at lengths of up to 4 m (13 ft 1 in).[24] The false gharial apparently has the largest skull of any extant crocodilian, in part because of the great length of the slender snout. Out of the eight longest crocodilian skulls from existing species that could be found in museums around the world, six of these belonged to false gharials. The longest crocodilian skull belonging to an extant species was of this species and measured 84 cm (33 in) in length, with a mandibular length of 104 cm (41 in). Most of the owners of these enormous skulls had no confirmed (or even anecdotal) total measurements for the animals, but based on the known skull-to-total length ratio for the species they would measure approximately 5.5 to 6.1 m (18 ft 1 in to 20 ft 0 in) in length.[25]
Distribution and habitat

False gharials are native to Peninsular Malaysia, Sarawak, and Indonesia (Sumatra, and Borneo), but were extirpated in Singapore, Vietnam, and Thailand. It is unclear if they remain in Java. Apart from rivers, they inhabit swamps and lakes.[2] The species is almost entirely found today in peat swamps and lowland swamp forests.[22] In the 1990s, information and sightings were available from 39 localities in 10 different river drainages, along with the remote river systems of Borneo.[citation needed]

Prior to the 1950s, Tomistoma occurred in freshwater ecosystems along the entire length of Sumatra east of the Barisan Mountains. The current distribution in eastern Sumatra has been reduced by 30-40% due to hunting, logging, fires, and agriculture.[26]
Ecology and behaviour
Diet

Until recently, very little was known about the diet or behaviour of the false gharial in the wild. Details are slowly being revealed. In the past, the false gharial was thought to have a diet of only fish and very small vertebrates, but more recent evidence indicates that it has a generalist diet despite its narrow snout. In addition to fish and smaller aquatic animals, mature adults prey on larger vertebrates, including proboscis monkeys, long-tailed macaques, deer, water birds, and reptiles.[27] There is an eyewitness account of a false gharial attacking a cow in East Kalimantan.[22]

The false gharial may be considered an ecological equivalent to Neotropical crocodiles such as the Orinoco and American crocodiles, which both have slender snouts but a broad diet.[12]
Reproduction

False gharials are mound-nesters. Females lay small clutches of 13 to 35 eggs per nest, and appear to produce the largest eggs of extant crocodilians. Sexual maturity in females appears to be attained around 2.5 to 3 m (8.2 to 9.8 ft), which is large compared to other crocodilians.[26]

It is not known when they breed in the wild or when the nesting season is. Once the eggs are laid, and construction of the mound is completed, the female abandons her nest. Unlike most other crocodilians, the young receive no parental care and are at risk of being eaten by predators, such as mongooses, tigers, leopards, and wild dogs. The young hatch after 90 days and are left to fend for themselves.[citation needed]
Conflict

In 2008, a 4-m female false gharial attacked and ate a fisherman in central Kalimantan; his remains were found in the gharial's stomach. This was the first verified fatal human attack by a false gharial.[27] However, by 2012, at least two more verified fatal attacks on humans by false gharials had occurred indicating perhaps an increase of human-false gharial conflict possibly correlated to the decline of habitat, habitat quality, and natural prey numbers.[28]
Threats

The false gharial is threatened with extinction throughout most of its range due to the drainage of its freshwater swamplands and clearance of surrounding rainforests. The species is also hunted frequently for its skin and meat, and the eggs are often harvested for human consumption.[27]
Conservation

The false gharial is listed on CITES Appendix I.[2] Currently population surveys indicate that while the false gharial is not for the most extirpated from areas it used to inhabit, the distribution of individuals is much more spotty than the previously more connective distribution, putting the animals at risk of genetic isolation.[29] In large part, the isolation of false gharials is due to extremely extensive habitat destruction and disturbance within the species' area of distribution, few areas outside of legally protected areas are likely to bear viable breeding populations.[30]

Steps have been taken by the Malaysian and Indonesian governments to prevent its extinction in the wild. There are reports of some populations rebounding in Indonesia, yet with this slight recovery, mostly irrational fears of attacks have surfaced amongst the local human population.[27] Yayasan Ulin (The Ironwood Foundation) is currently attempting to manage a wetland area in East Kalimantan, which is known to contain the false gharials.[31]
References

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.
Bezuijen, M.R.; Shwedick, B.; Simpson, B.K.; Staniewicz, A. & Stuebing, R. (2014). "Tomistoma schlegelii". IUCN Red List of Threatened Species. 2014: e.T21981A2780499.
"Common name".
"Common name 2". 5 June 2019.
"Common name 3".
"Common name 4" (PDF).
"Common name 5".
Müller, S. (1838). "Waarnemingen over de Indische Krokodillen en Beschrijving van eene nieuwe Soort". Tijdschrift voor Natuurlijke Geschiedenis en Physiologie (in Dutch). 5: 61–87.
Beolens, B.; Watkins, M. & Grayson, M. (2011). "Tomistoma schlegelii". The Eponym Dictionary of Reptiles. Baltimore: Johns Hopkins University Press. p. 235. ISBN 978-1-4214-0135-5.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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Bezuijen, M.R., Shwedick, B.M., Sommerlad, R., Stevenson, C. and Steubing, R.B. (2010). "Tomistoma Tomistoma schlegelii". In S.C. Manolis; C. Stevenson (eds.). Crocodiles. Status Survey and Conservation Action Plan. Darwin: Crocodile Specialist Group. pp. 133−138.
Mathew, A., Ganesan, M., Majid, R. A., & Beastall, C. (2011). Breeding of false gharial, Tomistoma schlegelii, at Zoo Negara, Malaysia (PDF). Zoo Negara.
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Bezuijen, M.R., Webb, G.J.W., Hartoyo, P., Samedi, Ramono, W.S., Manolis, S.C. (1998) The False Gharial (Tomistoma schlegelii) in Sumatra. In: Crocodiles. Proceedings of the 14th Working Meeting of the Crocodile Specialist Group, IUCN. The World Conservation Union, Gland, Switzerland and Cambridge, UK. Pp. 10–31.
Rachmawan, D.; Brend, S. (2009). "Human-Tomistoma interactions in central Kalimantan, Indonesian Borneo" (PDF). Crocodile Specialist Group Newsletter. 28 (1): 9–11.
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Stuebing, R. B., Bezuijen, M. R., Auliya, M., & Voris, H. K. (2006). The current and historic distribution of Tomistoma schlegelii (the False Gharial)(Müller 1838)(Crocodylia, Reptilia). The Raffles Bulletin of Zoology, 54(1), 181-197.
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"Siamese Crocodile Conservation Case Study | the Mohamed bin Zayed Species Conservation Fund".

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