|Female and subadult gharial|
The gharial (Gavialis gangeticus), also known as the gavial, and fish-eating crocodile is a crocodilian in the family Gavialidae, native to sandy freshwater river banks in the plains of the northern part of the Indian subcontinent. It is threatened by loss of riverine habitat, depletion of fish resources, and entanglement in fishing nets. As the wild population has declined drastically since the 1930s, the gharial is listed as Critically Endangered on the IUCN Red List. It once inhabited all the major river systems of the northern Indian subcontinent. Today, its distribution is limited to only 2% of its historical range. It inhabits foremost flowing rivers with high sand banks that it uses for basking and building nests. Adults mate in the cold season. The young hatch before the onset of the monsoon.
The gharial is one of the longest of all living crocodilians. Males reach a body length of up to 6 m (20 ft) and have a distinctive boss at the end of the snout, which resembles an earthenware pot known in Hindi as ghara. Its common name is derived from this similarity. With 110 sharp, interdigitated teeth in its long, thin snout, it is well adapted to catching fish, its main diet. Fossil remains were excavated in Pliocene deposits in the Sivalik Hills and Narmada River valley. It probably evolved 42 million years ago.
- C. gavial by Pierre Joseph Bonnaterre in 1789.
- C. longirostris by Johann Gottlob Theaenus Schneider in 1801.
- C. arctirostris by François Marie Daudin in 1802.
- C. tenuirostris by Georges Cuvier in 1807 for a gharial with a narrower skull and eye sockets than C. gangeticus.
The generic name Gavialis was proposed by Nicolaus Michael Oppel in 1811 for crocodiles with a cylindrical-shaped back. He placed this genus in the family Crocodilini. Rhamphostoma was proposed by Johann Georg Wagler in 1830 who considered this genus to contain two species, Crocodilus gangeticus and C. tenuirostris. The family name Gavialidae was proposed by Arthur Adams in 1854 for reptiles with a very long and slender muzzle, webbed feet and nearly equal teeth. Gavialis gangetica was the scientific name used by Albert Günther in 1864 who considered Gavialis a monotypic taxon, but placed in the family Crocodilidae. John Edward Gray reviewed zoological specimens and also considered it monotypic in 1869, but placed it in the family Gavialidae.
Gavialis and Tomistoma have been regarded close relatives since the 1950s and were thought to have had a common ancestor before the end of the Cretaceous. Results of molecular genetic studies indicate that the gharial and the false gharial (T. schlegelii) are indeed close relatives that genetically diverged in the Eocene about 42 million years ago.
South American gharials likely dispersed in the mid Tertiary from Africa and Asia. Fossil remains of the Puerto Rican gharial Aktiogavialis puertorisensis were discovered in a cave located in San Sebastián, Puerto Rico and dated to the Oligocene. This indicates that the Caribbean constituted a link between continents.
The gharial's belly is yellowish-white, its neck long and thick. There are two rows of ridges on the central region of the back. Male gharials develop a hollow bulbous nasal protuberance at the tip of the snout upon sexual maturity. This nasal growth starts growing over the nostrils at an age of 11.5 years and measures about 5 cm × 6 cm × 3.5 cm (2.0 in × 2.4 in × 1.4 in) at an age of 15.5 years, and enables the males to emit a hissing sound that can be heard at a distance of 75 m (246 ft). It resembles an earthen pot known locally as "ghara". The nasal growth is apparently used to indicate sexual maturity, as sound resonator when bubbling under water or other sexual behaviours. The gharial is the only living crocodilian with such visible sexual dimorphism.
The gharial's snout is very long and narrow, with 27 to 29 upper and 25 or 26 lower teeth on each side. The front teeth are the largest. The first, second, and third mandibular teeth fit into notches in the upper jaw. The nasal bones are rather short and widely separated from the premaxilla. The nasal opening is smaller than the supratemporal fossae. The jugal bone is raised and the extremely long symphysis extends to the 23rd or 24th tooth. The snout is dilated at the end. It becomes proportionally thicker with age. This long snout is considered an adaptation to a primarily piscivorous diet. The long, needle-like teeth are individually socketed.
The tail is well-developed and laterally flattened. Together with the webbed feet it provides tremendous manoeuvrability in deep water. On land, a gharial can only slide on its belly and push itself forward.
Gharial hatchlings are pale olive on the back and become darker with age. Dark cross-bands and speckles are visible on head, body and tail. Scutes on head, neck and back form a single continuous plate composed of 21 to 22 transverse series, and four longitudinal series. Scutes on the back are bony, but softer and feebly keeled on the sides. The outer edges of the forearms, legs, and feet are crested, and fingers and toes partly webbed.
The average size of mature gharials is 3.5–4.5 m (11–15 ft). Hatchlings range from 35–39.2 cm (13.8–15.4 in) in body length with a weight of 82–130 g (2.9–4.6 oz). Young gharials reach a length of 100 cm (39 in) in 18 months. Females grow up to a body length of 4.2 m (14 ft) and males more than 5.7 m (19 ft). Adults weigh 160 kg (350 lb) on average.
Distribution and habitat
The gharial once thrived in all the major river systems of the northern Indian subcontinent, from the Indus River in Pakistan and the Ganges to the Irrawaddy River in Myanmar. By 1976, its range had decreased to only 2% of its historical range, and fewer than 200 gharials were estimated to survive. In 2017, the global population was estimated at maximum 900 individuals, including 500 mature adults.
In Nepal, small populations are present and slowly recovering in tributaries of the Ganges, such as the Narayani-Rapti river system in Chitwan National Park and the Karnali-Babai river system in Bardia National Park. In spring 2017, the Babai River was surveyed over a stretch of 102 km (63 mi) using an unmanned aerial vehicle, which was flown at an altitude of 80 m (260 ft). 33 gharials were counted.
In India, gharial populations are present in the
- Ramganga River in Corbett National Park, where five gharials were recorded in 1974. Captive-bred gharials were released since the late 1970s. The population is breeding since 2008, and increased to about 42 adults by 2013. Surveys in 2015 revealed a population of 90 gharials including 59 breeding adults.
- Girwa River in Katarniaghat Wildlife Sanctuary where the small breeding population was reinforced with captive reared gharials since 1979. A total of 909 gharials were released until 2006, but only 16 nesting females were recorded in the same year.
- Gandaki River downstream the Triveni barrage west of Valmiki Tiger Reserve and adjacent to Sohagi Barwa Sanctuary. The population increased from 15 gharials in 2010 to 54 individuals recorded in March 2015 on a stretch of 320 km (200 mi). 35 of these gharials were wild-born.
- Chambal River in National Chambal Sanctuary where 107 gharials were recorded in 1974. Captive-bred gharials were released since 1979, and the population increased to 1,095 gharials in 1992. Until 2006, a total of 3,776 gharials were released, but only 68 nests were counted in this year. Between December 2007 and March 2008, 111 gharials were found dead. A total of 948 gharials were counted during surveys in 2013.
- Ganges, where 494 gharials were released between 2009 and 2012 in Hastinapur Wildlife Sanctuary.
- Son River where 164 captive-reared gharials were released between 1981 and 2011.
- Mahanadi River in Odisha's rainforest biome Satkosia Gorge Sanctuary where gharials were released since 1977.
Between 1934 and 1988, a few individuals were sighted in the Barak River and tributaries in Assam, Mizoram and Manipur, but surveys were not carried out. Between 1979 and 1993, several individuals were sighted in the Brahmaputra River, where the population had declined due to commercial fishing, poaching, encroachment by local people in gharial breeding grounds, and siltation of river beds following deforestation.
In the 1930s, the gharial was still considered common in the Indus River. By the early 1980s, it was considered almost extinct in the Indus. During surveys in 2008 and 2009, no gharial was sighted in the river.
Behaviour and ecology
The gharial is the most thoroughly aquatic of the living crocodilians. Young gharials move forward by pushing the diagonally opposite legs synchronously, whereby the hind feet step close to where the front feet were. At a young age, they can also gallop but do so only in emergency situations. When they reach a weight of about 1.5 kg (3.3 lb), their locomotion changes to pushing forward with hind and front legs simultaneously. Adult gharial do not have the ability to walk on land in the semi-upright stance as other crocodilians, but leave the water only for basking close to the water’s edge. When on the beach, they often turn round so as to face the water.
The gharial is a thermoconformer and seeks to cool down during hot times and to warm up when ambient temperature is cool. Gharials bask daily in winter, foremost in the mornings, and prefer sandy and moist beaches. They change their basking pattern with increasing daily temperatures, and start basking earlier in the mornings, move back into the river when it is hot, and return to the beach later in the afternoon. Large groups of young, subadult and adult gharials form in December and January to bask together. Adult males and females associate by mid February.
Young gharials hide and forage in shallow water. Gharials up to a body length of 120 cm (47 in) prefer a water level of 1–3 m (3.3–9.8 ft). With body size increasing, they move to deeper water. Gharials up to 180 cm (71 in) hunt and hide in 2–3 m (6.6–9.8 ft) deep water. Adult gharials prefer water deeper than 4 m (13 ft).
Though it is rarely seen, gharials are known to perform the characteristic "death roll" found in almost all extant crocodilians. This was also found to be true for other narrow-snouted crocodilians, and is likely linked to inter- and intraspecific competition.
The gharial is efficient in and well adapted to hunting fish under water, because of its sharp interdigitated teeth and long narrow snout that meets little resistance in the water. It does not chew its prey, but swallows it whole. Juvenile gharials were observed to jerk their heads back to manoeuvre fish into their gullets, sliding them in head first. Young gharials feed on insects, tadpoles, small fish and frogs. Adults also feed on small crustaceans. Remains of Indian softshell turtle (Nilssonia gangetica) was also found in gharial stomachs. Gharials tear apart large fish and pick up and swallow stones as gastroliths, probably to aid digestion or regulate buoyancy. Jewellery found in gharial stomachs may have been the reason for the myth that they ate humans.
Females mature at a body length of around 2.6 m (8.5 ft). Captive females breed at a body length of 3 m (9.8 ft). Male gharials mature at 15–18 years of age, when they reach a body length of around 4 m (13 ft) and once the ghara is developed.
Courting and mating starts by mid-February. In the dry season, reproductive females routinely move 80–120 km (50–75 mi) and join female breeding groups to dig nests together. These nests are 50 to 60 cm (20 to 24 in) deep holes in riverside sand or silt bank and 1 to 5 m (3.3 to 16.4 ft) away from the waterline. They lay 20–95 eggs. The eggs are the largest of all crocodilians and weigh an average of 160 g (5.6 oz). Each egg is 85–90 mm (3.3–3.5 in) long by 65–70 mm (2.6–2.8 in) wide. After 71 to 93 days of incubation, young gharials hatch in July just before the onset of the rainy season. Their sex is most likely determined by temperature. Females dig up the hatchlings in response to hatching chirps, but do not assist them to reach the water. They stay at nesting sites until monsoon floods arrive and return after monsoon.
Captive male gharials observed in the 1980s did not participate in guarding nests. A captive male gharial was observed to show an interest in hatchlings and was allowed by the female to carry hatchlings on his back. VHF radio tracking of a junior male gharial in the Chambal River revealed that he was the dominant male guarding nests at a communal nesting site for two years.
The gharial is sympatric with the mugger crocodile (Crocodylus palustris) in parts of its range. The gharial basks close to water on shallow, sandy beaches and lays eggs only in sandy soil near water. The mugger basks on sandy beaches, but also climbs steep embankments and rocks, and moves farther away for both basking and nest building.
The gharial population is estimated to have declined from 5,000-10,000 individuals in 1946 to fewer than 250 individuals alive in 2006, a decline of 96–98% within three generations. Gharials were killed by fishermen, hunted for skins, trophies and indigenous medicine, and their eggs collected for consumption. Today, the remaining individuals form several fragmented subpopulations. Hunting is no longer considered a significant threat. However, the wild population declined from an estimated 436 adult gharials in 1997 to fewer than 250 mature individuals in 2006 because:
- fishing and the use of gill nets increased in most of the present gharial habitat, even in protected areas;
- riverine habitat decreased as dams, barrages, irrigation canals and artificial embankments were built; siltation and sand-mining changed river courses; land is used for riparian agriculture and grazing by livestock.
When dead gharials were found in the Chambal River between December 2007 and March 2008, it was initially suspected that they had died either because of toxicants or because of the illegal use of fish nets, in which they became trapped and subsequently drowned. Later post mortem pathological testing of tissue samples from the dead gharials revealed high levels of heavy metals such as lead and cadmium, which together with stomach ulcers and protozoan parasites reported in most necropsies were thought to have caused their deaths.
Since the late 1970s, the gharial conservation approach was focused on reintroduction. Protected areas in India and Nepal used to be restocked with captive bred juvenile gharials. More than 5,000 gharials were released until 2006. In Nepal, wild eggs were collected and hatched since 1978, and a total of 1,365 gharials released in the rivers between 1981 and 2018. Releasing captive-reared gharials did not contribute significantly to re-establishing viable populations. In 2017, members of the Crocodile Specialist Group therefore recommended to foster engagement of local communities in gharial conservation programs.
In situ initiatives
The Indian Crocodile Conservation Project was set up in 1975 under the auspices of the Government of India, initially in Odisha's Satkosia Gorge Sanctuary. The project was implemented with financial aid of the United Nations Development Fund and Food and Agriculture Organization. A gharial breeding center was built in Nandankanan Zoological Park, so to later restock habitats with low numbers of gharials. A male gharial was flown in from Frankfurt Zoological Garden to become one of the founding animals of the breeding program. In subsequent years, several protected areas were established. An acute shortage of gharial eggs was overcome by their purchase from Nepal. Sixteen crocodile rehabilitation centers and five crocodile sanctuaries including the National Chambal Sanctuary and Katerniaghat Wildlife Sanctuary were established in India between 1975 and 1982. By 2004, 12,000 gharial eggs had been collected from wild and captive-breeding nests, and over 5,000 gharials reared to about a meter or more in length and released into the wild. But in 1991, funds were withdrawn for the captive-breeding and egg-collection programs. In 1997–1998, over 1,200 gharials and over 75 nests were located in the National Chambal Sanctuary, but no surveys were carried out between 1999 and 2003.
In December 2010, the then Indian Minister for Environment and Forests, Jairam Ramesh, visited the Madras Crocodile Bank Trust with Romulus Whitaker, and announced the formation of a National Tri-State Chambal Sanctuary Management and Coordination Committee for gharial conservation on 1,600 km2 (620 sq mi) of the National Chambal Sanctuary along the Chambal River in Madhya Pradesh, Rajasthan and Uttar Pradesh. The committee will comprise representatives of three states' Water Resources Ministries, states' Departments of Irrigation and Power, Wildlife Institute of India, Madras Crocodile Bank Trust, the Gharial Conservation Alliance, Development Alternatives, Ashoka Trust for Research in Ecology and the Environment, Worldwide Fund for Nature, and the divisional forest officers of the three states. The committee planned strategies for protection of gharials and their habitat, involving further research on gharial ecology and socioeconomic evaluation of dependent riparian communities. Funding for this new initiative was mobilized as a subscheme of the ‘Integrated Development of Wildlife Habitats’ with a yearly amount of 50–80 million Indian rupees (US$1 million to 1.7 million) for five years.
Gharials are bred in captivity in the National Chambal Sanctuary and in the Gharial Breeding Centre in Nepal's Chitwan National Park, where the eggs are hatched and then the gharials are grown for two to three years and average about one metre in length, when released.
In Europe, gharials are kept in Prague Zoo and Protivin Crocodile Zoo in the Czech Republic, and in the German Berlin Zoo. The European studbook of the European Association of Zoos and Aquaria is kept in Prague Zoo since 2017. La Ferme aux Crocodiles in France received six juveniles in 2000 from the Gharial Breeding Centre in Nepal.
In the United States, gharials are kept in Busch Gardens Tampa, Cleveland Metroparks Zoo, Fort Worth Zoo, Honolulu Zoo, San Diego Zoo, National Zoological Park, San Antonio Zoo and Aquarium and St. Augustine Alligator Farm Zoological Park. Bronx Zoo and Los Angeles Zoo received gharials in 2017.
- Brochu, C. A. (1997). "Morphology, fossils, divergence timing, and the phylogenetic relationships of Gavialis". Systematic Biology. 46 (3): 479–522. doi:10.1093/sysbio/46.3.479. PMID 11975331.
- Lang, J.; Chowfin, S.; Ross, J. P. (2019). "Gavialis gangeticus". The IUCN Red List of Threatened Species. IUCN. 2019: e.T8966A3148543. doi:10.2305/IUCN.UK.2019-1.RLTS.T8966A3148543.en.
- "Gharial". Catalogue of Life. ITIS. Species 2000.CS1 maint: others (link)
- Gmelin, J. F. (1789). "Lacerta gangetica ". Caroli a Linné. Systema naturae per regna tria naturae : secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis (in Latin). Tomus I. Pars III. Leipzig: G. E. Beers. pp. 1057–1058.
- Bonnaterre, P. J. (1789). "Le Gavial ". Tableau encyclopédique et méthodique des trois règnes de la nature. Erpétologie (in French). Paris: Chez Panckoucke. pp. 34–35..
- Schneider, J. G. T. (1801). "Longirostris ". Historiae amphibiorum naturalis et literariae fasciculus secundus (in Latin). Jena: F. Frommann. pp. 160–161.
- Daudin, F. M. (1802). "Le crocodile à bec étroit ou le grand Gavial ". Histoire Naturelle, Générale et Particulière des Reptiles; ouvrage faisant suit à l'Histoire naturelle générale et particulière, composée par Leclerc de Buffon; et rédigee par C.S. Sonnini, membre de plusieurs sociétés savantes (in French). Tome 2. Paris: F. Dufart. pp. 393–396.
- Cuvier, G. (1807). "Sur les différentes espèces de crocodiles vivans et sur leurs caractères distinctifs ". Annales du Muséum National d'Histoire Naturelle (in French). 10: 8–66.
- Oppel, N. M. (1811). "Familia. Crocodilini". Die Ordnungen, Familien und Gattungen der Reptilien als Prodrom einer Naturgeschichte derselben (in German). München: J. Lindauer. p. 19.
- Wagler, J. (1830). "Rhamphostoma ". Natürliches System der Amphibien, mit vorangehender Classification der Säugethiere und Vögel. Ein Beitrag zur vergleichenden Zoologie (in German). München: J. G. Cotta'scche Buchhandlung. p. 141.
- 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.
- Günther, A. (1864). "Gavialis, Geoffr.". The reptiles of British India. London: Robert Hardwicke. p. 63.
- Gray, J. E. (1869). "Synopsis of the species of recent Crocodilians or Emydosaurians, chiefly founded on the specimens in the British Museum and the Royal College of Surgeons". The Transactions of the Zoological Society of London. 6 (4): 125–169. doi:10.1111/j.1096-3642.1867.tb00575.x.
- Janke, A.; Gullberg, A.; Hughes, S.; Aggarwal, R. K.; Arnason, U. (2005). "Mitogenomic analyses place the gharial (Gavialis gangeticus) on the crocodile tree and provide pre-K/T divergence times for most crocodilians". Journal of Molecular Evolution. 61 (5): 620–626. Bibcode:2005JMolE..61..620J. CiteSeerX 10.1.1.561.2172. doi:10.1007/s00239-004-0336-9. PMID 16211427.
- Brochu, C. A. (2003). "Phylogenetic approaches toward crocodylian history". Annual Review of Earth and Planetary Sciences. 31: 357–397. Bibcode:2003AREPS..31..357B. doi:10.1146/annurev.earth.31.100901.141308.
- Vélez-Juarbe, J.; Brochu, C. A.; Santos, H. (2007). "A gharial from the Oligocene of Puerto Rico: transoceanic dispersal in the history of a non-marine reptile". Proceedings of the Royal Society. 274 (1615): 1245–1254. doi:10.1098/rspb.2006.0455. PMC 2176176. PMID 17341454.
- Patnaik, R.; Schleich, H. H. (1993). "Fossil crocodile remains from the Upper Siwaliks of India". Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie (33): 91–117.
- 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.
- Brazaitis, P. (1973). "Family Gavialidae Gavialis gangeticus Gmelin". Zoologica : Scientific Contributions of the New York Zoological Society. 3: 80−81.
- Biswas, S.; Acharjyo, L. N.; Mohapatra, S. (1977). "A note on the protuberance or knob on the snout of male gharial, Gavialis gangeticus (Gmelin)". Journal of the Bombay Natural History Society. 74 (3): 536–537.
- Stevenson, C.; Whitaker, R. (2010). "Gharial Gavialis gangeticus" (PDF). In Manolis, S. C.; Stevenson, C. (eds.). Crocodiles. Status Survey and Conservation Action Plan (Third ed.). Darwin: Crocodile Specialist Group. pp. 139–143.
- Martin, B. G. H.; Bellairs, A. d’A. (1977). "The narial excrescence and pterygoid bulla of the gharial, Gavialis gangeticus (Crocodilia)". Journal of Zoology. 182 (4): 541–558. doi:10.1111/j.1469-7998.1977.tb04169.x.
- Whitaker, R.; Members of the Gharial Multi-Task Force; Madras Crocodile Bank (2007). "The Gharial: Going Extinct Again" (PDF). Iguana. 14 (1): 24–33. Archived from the original (PDF) on 2011-07-26.
- Boulenger, G. A. (1890). "Gavialis gangeticus". Fauna of British India. Reptilia and Batrachia. London: Taylor and Francis. p. 3.
- Boulenger, G. A. (1889). "Gavialis". Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles in the British Museum (Natural History) (New ed.). London: Trustees of the British Museum (Natural History). pp. 275–276.
- Maskey, T. M.; Percival, H. F. (1994). "Status and Conservation of Gharial in Nepal". Proceedings of the 12th Working Meeting of the Crocodile Specialist Group. Thailand.
- Whitaker, R.; Basu, D. (1982). "The Gharial (Gavialis gangeticus): A review". Journal of the Bombay Natural History Society. 79 (3): 531–548.
- Pitman, C. R. S. (1925). "The length attained by and the habits of the Gharial (G. gangeticus)". Journal of the Bombay Natural History Society. 30 (3): 703.
- Priol, P. (2003). Gharial field study report. Kathmandu: A report submitted to Department of National Parks and Wildlife Conservation.
- Ballouard, J.M.; Priol, P.; Oison, J.; Ciliberti, A. & Cadi, A. (2010). "Does reintroduction stabilize the population of the critically endangered gharial (Gavialis gangeticus, Gavialidae) in Chitwan National Park, Nepal?". Aquatic Conservation: Marine and Freshwater Ecosystems. 20 (7): 756–761. doi:10.1002/aqc.1151.
- Thapa, G. J.; Thapa, K.; Thapa, R.; Jnawali, S. R.; Wich, S. A.; Poudyal, L. P.; Karki, S. (2018). "Counting crocodiles from the sky: monitoring the critically endangered gharial (Gavialis gangeticus) population with an unmanned aerial vehicle (UAV)". Journal of Unmanned Vehicle Systems. 6 (2): 71–82. doi:10.1139/juvs-2017-0026.
- Chowfin, S. (2010). "Crocodilian and freshwater research and conservation project, Uttarakhand, India" (PDF). Crocodile Specialist Group Newsletter. 29 (3): 19.
- Chowfin, S. M.; Leslie, A. J. (2013). "A preliminary investigation into nesting and nest predation of the critically endangered, gharial (Gavialis gangeticus) at Boksar in Corbett Tiger Reserve, Uttarakhand, India" (PDF). World Crocodile Conference. Proceedings of the 22nd Working Meeting of the IUCN-SSC Crocodile Specialist Group. Gland: IUCN Crocodile Specialist Group. pp. 26−28.
- Chowfin, S. M.; Leslie, A. J. (2016). "The Gharial (Gavialis gangeticus) in Corbett Tiger Reserve". In Crocodile Specialist Group (ed.). Crocodiles. Proceedings of the 24th Working Meeting of the Crocodile Specialist Group Skukuza, South Africa, 23-26 May 2016. Gland: IUCN. pp. 120–124.
- Rao, R. J.; Choudhury, B. C. (1992). "Sympatric distribution of gharial and mugger in India". Journal of the Bombay Natural History Society. 89: 312–315.
- Choudhary, S. K. (2010). Multi-species Survey in River Gandak, Bihar with focus on Gharial and Ganges River Dolphin. Bhagalpur: T. M. Bhagalpur University.
- Choudhury, B. C.; Behera, S. K.; Sinha, S. K.; Chandrashekar, S. (2016). "Restocking, Monitoring, Population Status, New Breeding Record and Conservation Actions for Gharial in the Gandak River, Bihar, India". In Crocodile Specialist Group (ed.). Crocodiles. Proceedings of the 24th Working Meeting of the Crocodile Specialist Group Skukuza, South Africa, 23-26 May 2016. Gland: IUCN. p. 124.
- Hussain, S. A. (1999). "Reproductive success, hatchling survival and rate of increase of gharial Gavialis gangeticus in National Chambal Sanctuary, India". Biological Conservation. 87 (2): 261−268. doi:10.1016/S0006-3207(98)00065-2.
- Nawab, A.; Basu, D. J.; Yadav, S. K.; Gautam, P. (2013). "Impact of Mass Mortality of Gharial Gavialis gangeticus (Gmelin, 1789) on its Conservation in the Chambal River in Rajasthan". In Sharma, B. K.; Kulshreshtha, S.; Rahmani, A. R. (eds.). Faunal Heritage of Rajasthan, India. Springer International Publishing. pp. 221–229. doi:10.1007/978-3-319-01345-9_9. ISBN 978-3-319-01344-2.
- Rao, R.J.; Tagor, S.; Singh, H.; Dasgupta, N. (2013). "Monitoring of Gharial (Gavialis gangeticus) and its habitat in the National Chambal Sanctuary, India" (PDF). World Crocodile Conference. Proceedings of the 22nd Working Meeting of the IUCN-SSC Crocodile Specialist Group. Gland: IUCN Crocodile Specialist Group. pp. 66−73.
- Yadav, S. K.; Nawab, A.; Afifullah Khan, A. (2013). "Conserving the Critically Endangered Gharial Gavialis gangeticus in Hastinapur Wildlife Sanctuary, Uttar Pradesh: Promoting better coexistence for conservation" (PDF). World Crocodile Conference. Proceedings of the 22nd Working Meeting of the IUCN-SSC Crocodile Specialist Group. Gland: IUCN Crocodile Specialist Group. pp. 78−82.
- Nair, T.; Katdare, S. (2013). "Dry-season assessment of gharials (Gavialis gangeticus) in the Betwa, Ken and Son Rivers, India" (PDF). World Crocodile Conference. Proceedings of the 22nd Working Meeting of the IUCN-SSC Crocodile Specialist Group. Gland: IUCN Crocodile Specialist Group. pp. 53−65.
- Bustard, H. R. (1983). "Movement of wild Gharial, Gavialis gangeticus (Gmelin) in the River Mahanadi, Orissa (India)". British Journal of Herpetology. 6: 287–291.
- Choudhury, A. U. (1997). "Records of the gharial Gavialis gangeticus (Gmelin) from the Barak river system of north-eastern India". Journal of the Bombay Natural History Society. 94 (1): 162–164.
- Choudhury, A. U. (1998). "Status of the gharial Gavialis gangeticus in the main Brahmaputra river". Journal of the Bombay Natural History Society. 95 (1): 118–120.
- Hasan, K.; Alam, S. (2016). "Chapter 3: Findings". Gharials of Bangladesh. Dhaka: IUCN Bangladesh Country Office. pp. 29–65.
- Rao, C. J. (1933). "Gavial on the Indus". Journal of the Sind Natural History Society. 1 (4): 37.
- Bustard, H. R.; Choudhury, B. C. (1983). "The distribution of the Gharial". Journal of the Bombay Natural History Society. 79 (2): 427–429.
- Biswas, S. (1970). "A Preliminary Survey of the Gharial in the Kosi River". Indian Forester. 96 (9): 705–710.
- Bustard, H. R.; Singh, L. A. K. (1977). "Studies on the Indian gharial Gavialis gangeticus (Gmelin) (Reptilia, Crocodilia) change in terrestrial locomotory pattern with age". Journal of the Bombay Natural History Society. 74 (3): 534−535.
- Lang, J. W. (1987). "Crocodilian behaviour: implications for management". In Webb, G. J. W.; Manolis, S. C.; Whitehead, P. J. (eds.). Wildlife Management: Crocodiles and Alligators. Sydney: Surrey Beatty and Sons. pp. 273−294.
- Lang, J. W.; Kumar, P. (2013). "Behavioral ecology of Gharial on the Chambal River, India" (PDF). World Crocodile Conference. Proceedings of the 22nd Working Meeting of the IUCN-SSC Crocodile Specialist Group. Gland: IUCN Crocodile Specialist Group. pp. 42−52.
- Hussain, S. A. (2009). "Basking site and water depth selection by Gharial Gavialis gangeticus Gmelin 1789 (Crocodylia, Reptilia) in National Chambal Sanctuary, India and its implication for river conservation". Aquatic Conservation: Marine and Freshwater Ecosystems. 19 (2): 127–133. doi:10.1002/aqc.960.
- Drumheller, S. K.; Darlington, J.; Vliet, K. A. (2019). "Surveying death roll behavior across Crocodylia". Ethology Ecology & Evolution. 31 (4): 329–347. doi:10.1080/03949370.2019.1592231.
- Bustard, H. R. (1983). "Size at first breeding in the Gharial [Gavialis gangeticus (Gmelin)] (Reptilia, Crocodilia) in captivity". Journal of the Bombay Natural History Society. 79 (1): 206−207.
- Smith, M. A. (1931). "Gavialis". The Fauna of British India, Including Ceylon and Burma. Reptilia and Amphibia. Volume I.—Loricata, Testudines. London: Secretary of State for India in Council. pp. 37–40.
- Bustard, H. R. (1982). "Behaviour of the male Gharial during the nesting and post-hatching period". Journal of the Bombay Natural History Society. 79 (3): 677–680.
- Lang, J. W.; Kumar, P. (2016). "Chambal Gharial Ecology Project - 2016 Update". In Crocodile Specialist Group (ed.). Crocodiles. Proceedings of the 24th Working Meeting of the Crocodile Specialist Group Skukuza, South Africa, 23-26 May 2016. Gland: IUCN. pp. 136–148.
- Rao, R. J.; Choudhury, B. C. (1992). "Sympatric distribution of Gharial Gavialis gangeticus and Mugger Crocodylus palustris in India". Journal of the Bombay Natural History Society. 89 (3): 313–314.
- Choudhary, S.; Choudhury, B. C.; Gopi, G. V. (2018). "Spatio-temporal partitioning between two sympatric crocodilians (Gavialis gangeticus & Crocodylus palustris) in Katarniaghat Wildlife Sanctuary, India". Aquatic Conservation: Marine and Freshwater Ecosystems. 28 (5): 1067–1076. doi:10.1002/aqc.2911.
- Whitaker, R.; Basu, D.; Huchzermeyer, F. (2008). "Update on gharial mass mortality in National Chambal Sanctuary" (PDF). Crocodile Specialist Group Newsletter. 27 (1): 4–8.
- Khadka, B.B. (2018). "119 Juvenile Gharials released into the Rapti River, Chitwan National Park, Nepal" (PDF). Crocodile Specialist Group Newsletter. 37 (1): 12−13.
- Webb, G. (2018). "Editorial" (PDF). Crocodile Specialist Group Newsletter. 37 (1): 3−4.
- Bustard, H. R. (1999). "Indian Crocodile Conservation Project". Envis Wildlife and Protected Areas. 2 (1): 5–9.
- Mallapur, G. (2010). "Gharial Conservation gets a leg-up!". Madras: The Madras Crocodile Bank Trust and Center for Herpetology. Archived from the original on 2012-11-19.
- Choudhury, B. C. (1999). "Crocodile Breeding in Indian Zoos". Envis Wildlife and Protected Areas. 2 (1): 100–103.
- Ziegler, T. (2018). "Europe" (PDF). In Crocodile Specialist Group (ed.). Crocodile Specialist Group Steering Committee Meeting, Universidad Nacional del Litoral, Santa Fe, Argentina (6 May 2018). pp. Agenda Item: SC. 2.7.
- Fougeirol, L. (2009). "Le gavial du Gange, un rêve" (in French). www.luc-fougeirol.com. Archived from the original on 2011-02-06. Retrieved 2017-12-26.
- Bronx Zoo (2017). "Indian Gharials Return to the Zoo". Wildlife Conservation Society.
- L.A. Zoo (2017). "L.A. Zoo Becomes One of Nine Zoos in the Western Hemisphere to House Gharials Flown in from India". Los Angeles Zoo and Botanical Gardens.
- Parpola, A. (2011). "Crocodile in the Indus Civilization and later South Asian traditions" (PDF). In Osada, H.; Endo, H. (eds.). Linguistics, Archaeology and the Human Past. Kyoto, Japan: Indus Project Research Institute for Humanity and Nature. pp. 1–57. ISBN 978-4-902325-67-6.
- Behera, S. K.; Singh, H.; Sagar, V. (2014). "Indicator Species (Gharial and Dolphin) of Riverine Ecosystem: An Exploratory of River Ganga". In Sanghi, R. (ed.). Our National River Ganga: Lifeline of Millions. Switzerland: Springer International Publishing. pp. 103–123. doi:10.1007/978-3-319-00530-0_4. ISBN 978-3-319-00529-4.
- Saikia, P. K. (2013). "Indian Gharial (Gavialis gangeticus): Status, Ecology and Conservation". In Singharavelan, N. (ed.). Rare Animals of India. Sharjah, Oakland, Bussum: Bentham Science Publishers. pp. 42–58. ISBN 978-1608054855.
- Das, I. (2002). "Gavialis gangeticus". A Photographic Guide to Snakes and other Reptiles of India. Sanibel Island, Florida: Ralph Curtis Publishing. p. 122. ISBN 0883590565.
- Goin, C. J.; Goin, O. B.; Zug, G. R. (1978). "Gavialis gangeticus". Introduction to Herpetology (Third ed.). San Francisco: W. H. Freeman. p. 341, 345, Figure 17-1. ISBN 0716700204.
- Joshi, A. R. (2018). "Nepali scientists deploy drones to count endangered crocodiles". Mongabay.
- Tarun Nair and Suyash Katdare (2014). "Mayawati And Other River Monsters – In Search Of Gharials In The Ken River". Sanctuary Asia.
- "Gavialidae". reptilis.net.
- Arkive: Gharial
- Adam Britton: Gavialis gangeticus
- Species Gavialis gangeticus at The Reptile Database
- "Gharials". Fort Worth Zoo.
- "Gharial". National Geographic Society.
- "Mystery of crocs' mass die-off". BBC News. 2008.
- Lenin, J. "The song of the Ganges gharial". www.india-seminar.com.