Jump to content

Whale shark

This is a good article. Click here for more information.
Page semi-protected
From Wikipedia, the free encyclopedia
(Redirected from Whale sharks)

Whale shark
Temporal range: 28.4–0 Ma[1]
Whale shark in the Andaman Sea around the Similan Islands
The size of various whale shark individuals with a human for scale
CITES Appendix II (CITES)[3]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Order: Orectolobiformes
Family: Rhincodontidae
Genus: Rhincodon
A. Smith, 1829[7][6]
Species:
R. typus
Binomial name
Rhincodon typus
Range of whale shark[needs update]
Synonyms
  • Micristodus punctatus Gill, 1865
  • Rhineodon Denison, 1937
  • Rhiniodon typus A. Smith, 1828[4]
  • Rhinodon pentalineatus Kishinouye, 1901
  • Rhinodon typicus Müller & Henle, 1839

The whale shark (Rhincodon typus) is a slow-moving, filter-feeding carpet shark and the largest known extant fish species. The largest confirmed individual had a length of 18.8 m (61.7 ft).[8] The whale shark holds many records for size in the animal kingdom, most notably being by far the most massive living non-cetacean animal. It is the sole member of the genus Rhincodon and the only extant member of the family Rhincodontidae, which belongs to the subclass Elasmobranchii in the class Chondrichthyes. Before 1984 it was classified as Rhiniodon into Rhinodontidae.

Whale sharks inhabit the open waters of all tropical oceans. They are rarely found in water below 21 °C (70 °F).[2] Whale sharks' lifespans are estimated to be between 80 and 130 years, based on studies of their vertebral growth bands and the growth rates of free-swimming sharks.[9][10][11] Whale sharks have very large mouths and are filter feeders, which is a feeding mode that occurs in only two other sharks, the megamouth shark and the basking shark. They feed almost exclusively on plankton and small fishes, and do not pose any threat to humans.

The species was distinguished in April 1828 after the harpooning of a 4.6 m (15 ft) specimen in Table Bay, South Africa. Andrew Smith, a military doctor associated with British troops stationed in Cape Town, described it the following year.[12] The name "whale shark" refers to the animal's appearance and large size; it is a fish, not a mammal, and (like all sharks) is not closely related to whales.[13]

Description

Whale sharks possess a broad, flattened head with a large mouth and two small eyes located at the front corners.[14][15] Unlike many other sharks, whale shark mouths are located at the front of the head rather than on the underside of the head.[16] A 12.1 m (39.7 ft) whale shark was reported to have a mouth 1.55 m (5.1 ft) across.[17] Whale shark mouths can contain over 300 rows of tiny teeth and 20 filter pads which it uses to filter feed.[18] The spiracles are located just behind the eyes. Whale sharks have five large pairs of gills. Their skin is dark grey with a white belly marked with an arrangement of pale grey or white spots and stripes that is unique to each individual. The skin can be up to 15 cm (5.9 in) thick and is very hard and rough to the touch. The whale shark has three prominent ridges along its sides, which start above and behind the head and end at the caudal peduncle.[15] The shark has two dorsal fins set relatively far back on the body, a pair of pectoral fins, a pair of pelvic fins and a single medial anal fin. The caudal fin has a larger upper lobe than the lower lobe (heterocercal).

Whale sharks have been found to possess dermal denticles on the surface of their eyeballs that are structured differently from their body denticles. The dermal denticles, as well as the whale shark's ability to retract its eyes deep into their sockets, serve to protect the eyes from damage.[19][20]

Evidence suggests that whale sharks can recover from major injuries and may be able to regenerate small sections of their fins. Their spot markings have also been shown to reform over a previously wounded area.[21]

The complete and annotated genome of the whale shark was published in 2017.[22]

Rhodopsin, the light-sensing pigment in the rod cells of the retina, is normally sensitive to green and used to see in dim light, but in the whale shark (and the bottom-dwelling cloudy catshark) two amino acid substitutions make the pigment more sensitive to blue light instead, the light that dominates the deep ocean. One of these mutations also makes rhodopsin vulnerable to higher temperatures. In humans, a similar mutation leads to congenital stationary night blindness, as the human body temperature makes the pigment decay.[23][24] This pigment becomes unstable in shallow water, where the temperature is higher and the full spectrum of light is present. To protect from this instability, the whale shark deactivates the pigment when in shallow water (as otherwise the pigment would hinder full color vision). In the colder environment at 2,000 meters below the surface where the shark dives, it is activated again.[25] The mutations thus allow the shark to see well at both ends of its great vertical range.[26][27] The eyes have also lost all cone opsins except LWS.[28]

Size

The whale shark is the largest non-cetacean animal in the world. However, the maximum size and growth patterns of the species are not well understood.[8][29]

Limited evidence, mostly from males, suggests that sexual maturity occurs around 8 to 9 meters (26 to 30 ft) in length, with the possibility of females sexually maturing at a similar size or larger.[14][30][31][32]

Various studies have aimed to estimate the growth and longevity of whale sharks, either by analysing evidence from vertebral growth rings or measurements taken from re-sighted sharks over several years. This information is used to model growth curves, which can predict asymptotic length. The growth curves produced from these studies have estimated asymptotic lengths ranging from 15 to 21.9 meters (49 to 72 ft).[11][33][34][29][9]

A 2020 study looked at the growth of whale shark individuals over a 10-year period around the Ningaloo Reef and concluded the species exhibits sexual dimorphism with regard to size, with females growing larger than males. The study found that males on average reach 8 to 9 meters (26 to 30 ft) in length. The same study had less female data but estimated an average length of around 14.5 m (48 ft). However, this value dropped to 12.1 m (40 ft) if data from aquarium whale sharks was included. The authors noted that these estimates represent average asymptotic size and are not the maximum sizes possible. Additionally, they acknowledged the potential for regional size variation.[29]

Most previous growth studies have had data predominately from males and none have data from sharks over ~10 m (33 ft). Not all previous studies created separate growth curves for males and females, instead combining data from both sexes. Those studies that made sex-specific growth curves have estimated large asymptotic length estimates for males, with lengths of 18 m (59 ft) or more. However, mostly immature males were available in these studies, with few adults to constrain the upper portion of the growth curves.[29]

The largest total length for the species is uncertain due to a lack of detailed documentation of the largest reported individuals. Whale sharks as large as 18 to 20 meters (59 to 66 ft) in length have been reported in scientific literature. However, most whale sharks observed are smaller.[8]

Large whale sharks are difficult to measure accurately, both on the land and in the water. When on land, the total length measurement can be affected by how the tail is positioned, either angled as it would be in life or stretched as far as possible. Historically, techniques such as comparisons to objects of known size and knotted ropes have been used for in-water measurements, but these techniques may be inaccurate.[31] Various forms of photogrammetry have been used to improve the accuracy of in-water measurements, including underwater and aerial techniques.[31][35][36]

Reports of large whale sharks

Since the 1800s, there have been accounts of very large whale sharks. Some of these are as follows:

In 1868, the Irish natural scientist Edward Perceval Wright obtained several small whale shark specimens in the Seychelles. Wright was informed of one whale shark that was measured as exceeding 45 ft (14 m). Wright claimed to have observed specimens over 50 ft (15 m) and was told of specimens upwards of 70 ft (21 m).[37]

Hugh M. Smith described a huge animal caught in a bamboo fish trap in Thailand in 1919. The shark was too heavy to pull ashore, and no measurements were taken. Smith learned through independent sources that it was at least 10 wa (a Thai unit of length measuring between a person's outstretched arms). Smith noted that one wa could be interpreted as either 2 m (6.6 ft) or the approximate average of 1.7 to 1.8 m (5.6–5.9 ft), based on the local fishermen.[38] Later sources have stated this whale shark as approximately 18 m (59 ft), but the accuracy of the estimate has been questioned.[14][8]

In 1934, a ship named the Maunganui came across a whale shark in the southern Pacific Ocean and rammed it. The shark became stuck on the prow of the ship, supposedly with 15 ft (4.6 m) on one side and 40 ft (12.2 m) on the other, suggesting a total length of about 55 ft (17 m).[39][40]

Scott A. Eckert & Brent S. Stewart reported on satellite tracking of whale sharks from 1994 to 1996. Out of the 15 individuals tracked, two females were reported as measuring 15 m (49 ft) and 18 m (59 ft).[41] A 20.75 m (68.1 ft) long whale shark was reported as being stranded along the Ratnagiri coast in 1995.[42][43] A female individual with a standard length of 15 m (49.2 ft) and an estimated total length at 18.8 m (61.7 ft) was reported from the Arabian Sea in 2001.[44] In a 2015 study reviewing the size of marine megafauna, McClain and colleagues considered this female as being the most reliable and accurately measured.[8]

On 7 February 2012, a large whale shark was found floating 150 kilometres (93 mi) off the coast of Karachi, Pakistan. The length of the specimen was said to be between 11 and 12 m (36 and 39 ft), with a weight of around 15,000 kg (33,000 lb).[45]

  • Jaws
    Jaws
  • Teeth
    Teeth
  • Eye
    Eye
  • Close up showing eyeball denticles
    Close up showing eyeball denticles
  • Top of head
    Top of head

Distribution and habitat

The whale shark inhabits all tropical and warm-temperate seas. The fish is primarily pelagic, and can be found in both coastal and oceanic habitats.[46] Tracking devices have shown that the whale shark displays dynamic patterns of habitat utilization, likely in response to availability of prey. Whale sharks observed off the northeast Yucatan Peninsula tend to engage in inshore surface swimming between sunrise and mid-afternoon, followed by regular vertical oscillations in oceanic waters during the afternoon and overnight. About 95% of the oscillating period was spent in epipelagic depths (<200 metres (660 ft)), but whale sharks also took regular deep dives (>500 metres (1,600 ft)), often descending in brief "stutter steps", perhaps for foraging. The deepest recorded dive was 1,928 metres (6,325 ft), making the whale shark the deepest diving fish to be recorded. Whale sharks were also observed to remain continuously at depths of greater than 50 metres (160 ft) for three days or more.[47][48][49]

The whale shark is migratory[10] and has two distinct subpopulations: an Atlantic subpopulation, from Maine and the Azores to Cape Agulhas, South Africa, and an Indo-Pacific subpopulation which holds 75% of the entire whale shark population. It usually roams between 30°N and 35°S where water temperatures are higher than 21 °C (70 °F) but have been spotted as far north as the Bay of Fundy, Canada and the Sea of Okhotsk north of Japan and as far south as Victoria, Australia.[2]

Seasonal feeding aggregations occur at several coastal sites such as the Persian Gulf and Gulf of Oman, Ningaloo Reef in Western Australia, Darwin Island in the Galápagos, Quintana Roo in Mexico, Mafia Island of Pwani Region in Tanzania, Inhambane province in Mozambique, the Philippines, around Mahe in the Seychelles,[46] the Gujarat[46] and Kerala coasts of India,[50][51] Taiwan, southern China[46] and Qatar.[52]

In 2011, more than 400 whale sharks gathered off the Yucatan Coast. It was one of the largest gatherings of whale sharks recorded.[53] Aggregations in that area are among the most reliable seasonal gatherings known for whale sharks, with large numbers occurring in most years between May and September. Associated ecotourism has grown rapidly to unsustainable levels.[54]

Growth and reproduction

Growth, longevity, and reproduction of the whale shark are poorly understood.[11][30][34]

Vertebral growth bands have been used to estimate the age, growth, and longevity of whale sharks. However, there was uncertainty as to whether vertebrae growth bands are formed annually or biannually.[33][9][11] A 2020 study compared the ratio of Carbon-14 isotopes found in growth bands of whale shark vertebrae to nuclear testing events in the 1950–60s, finding that growth bands are laid down annually. The study found an age of 50 years for a 10 m (33 ft) female and 35 years for a 9.9 m (32 ft) male.[34] Various studies looking at vertebrae growth bands and measuring whale sharks in the wild have estimated their lifespans from ~80 years and up to ~130 years.[9][10][11]

Evidence suggests that males grow faster than females in the earlier stages of life but ultimately reach a smaller maximum size.[29] Whale sharks exhibit late sexual maturity.[34] One study looking at free-swimming whale sharks estimated the age at maturity in males at ~25 years.[11]

Pupping of whale sharks has not been observed, but mating has been witnessed twice in St Helena.[55] Mating in this species was filmed for the first time in whale sharks off Ningaloo Reef via airplane in Australia in 2019, when a larger male unsuccessfully attempted to mate with a smaller, immature female.[56]

The capture of a ~10.6 m (35 ft) female in July 1996 that was pregnant with ~300 pups indicated that whale sharks are ovoviviparous.[10][57][58] The eggs remain in the body and the females give birth to live young which are 40 to 60 cm (16 to 24 in) long. Evidence indicates the pups are not all born at once, but rather the female retains sperm from one mating and produces a steady stream of pups over a prolonged period.[59]

On 7 March 2009, marine scientists in the Philippines discovered what is believed to be the smallest living specimen of the whale shark. The young shark, measuring only 38 cm (15 in), was found with its tail tied to a stake at a beach in Pilar, Sorsogon, Philippines, and was released into the wild. Based on this discovery, some scientists no longer believe this area is just a feeding ground; this site may be a birthing ground, as well. Both young whale sharks and pregnant females have been seen in the waters of St Helena in the South Atlantic Ocean, where numerous whale sharks can be spotted during the summer.[60][61]

In a report from Rappler last August 2019, whale sharks were sighted during WWF Philippines' photo identification activities in the first half of the year. There were a total 168 sightings – 64 of them "re-sightings" or reappearances of previously recorded whale sharks. WWF noted that "very young whale shark juveniles" were identified among the 168 individuals spotted in the first half of 2019. Their presence suggests that the Ticao Pass may be a pupping ground for whale sharks, further increasing the ecological significance of the area.[62]

Large adult females, often pregnant, are a seasonal presence around the Galapagos Islands, which may have reproductive significance.[63][64] One study between 2011 and 2013 found that 91.5% of the whale sharks observed around Darwin Island were adult females.[64]

Diet

Whale shark filtering plankton in Maldives
A whale shark in the Philippines with remoras
Whale shark with open mouth feeding in La Paz, Baja California Sur, Mexico

The whale shark is a filter feeder – one of only three known filter-feeding shark species (along with the basking shark and the megamouth shark). It feeds on plankton including copepods, krill, fish eggs, Christmas Island red crab larvae[65] and small nektonic life, such as small squid or fish. It also feeds on clouds of eggs during mass spawning of fish and corals.[66] The many rows of vestigial teeth play no role in feeding. Feeding occurs either by ram filtration, in which the animal opens its mouth and swims forward, pushing water and food into the mouth, or by active suction feeding, in which the animal opens and closes its mouth, sucking in volumes of water that are then expelled through the gills. In both cases, the filter pads serve to separate food from water. These unique, black sieve-like structures are presumed to be modified gill rakers. Food separation in whale sharks is by cross-flow filtration, in which the water travels nearly parallel to the filter pad surface, not perpendicularly through it, before passing to the outside, while denser food particles continue to the back of the throat.[67] This is an extremely efficient filtration method that minimizes fouling of the filter pad surface. Whale sharks have been observed "coughing", presumably to clear a build-up of particles from the filter pads. Whale sharks migrate to feed and possibly to breed.[10][68][69]

The whale shark is an active feeder, targeting concentrations of plankton or fish. It is able to ram filter feed or can gulp in a stationary position. This is in contrast to the passive feeding basking shark, which does not pump water. Instead, it swims to force water across its gills.[10][68]

A juvenile whale shark is estimated to eat 21 kg (46 pounds) of plankton per day.[70]

The BBC program Planet Earth filmed a whale shark feeding on a school of small fish. The same documentary showed footage of a whale shark timing its arrival to coincide with the mass spawning of fish shoals and feeding on the resultant clouds of eggs and sperm.[66]

Whale sharks are known to prey on a range of planktonic and small nektonic organisms that are spatiotemporally patchy. These include krill, crab larvae, jellyfish, sardines, anchovies, mackerels, small tunas, and squid. In ram filter feeding, the fish swims forward at constant speed with its mouth fully open, straining prey particles from the water by forward propulsion. This is also called ‘passive feeding’, which usually occurs when prey is present at low density.[71]

Due their mode of feeding, whale sharks are susceptible to the ingestion of microplastics. As such, the presence of microplastics in whale shark scat was recently confirmed.[72]

Relationship with humans

Behavior toward divers

In Oslob, Philippines, whale sharks are fed shrimp to return every morning for tourists and divers.

Despite its size, the whale shark does not pose any danger to humans. Younger whale sharks are gentle and can play with divers. Underwater photographers such as Fiona Ayerst have photographed them swimming close to humans without any danger.[73] Although whale sharks are docile fish, touching or riding the sharks is strictly forbidden and fineable in most countries,[74][75][76] as it can cause serious harm to the animal.[77][78][79]

Swimming alongside an adult free diver

The shark is seen by divers in many places, including the Bay Islands in Honduras, Thailand, Indonesia (Bone Bolango, Cendrawasih Bay), the Philippines, the Maldives close to Maamigili (South Ari Atoll), the Red Sea, Western Australia (Ningaloo Reef, Christmas Island), Taiwan, Panama (Coiba Island), Belize, Tofo Beach in Mozambique, Sodwana Bay (Greater St. Lucia Wetland Park) in South Africa,[73] the Galapagos Islands, Saint Helena, Isla Mujeres (Caribbean Sea), La Paz, Baja California Sur and Bahía de los Ángeles in Mexico, the Seychelles, West Malaysia, islands off eastern peninsular Malaysia, India, Sri Lanka, Oman, Fujairah, Puerto Rico, and other parts of the Caribbean.[80] Juveniles can be found near the shore in the Gulf of Tadjoura, near Djibouti, in the Horn of Africa.[81]

Conservation status

There is currently no robust estimate of the global whale shark population. The species is considered endangered by the IUCN due to the impacts of fisheries, by-catch losses, and vessel strikes,[82] combined with its long lifespan and late maturation.[2] In June 2018 the New Zealand Department of Conservation classified the whale shark as "Migrant" with the qualifier "Secure Overseas" under the New Zealand Threat Classification System.[83]

It is listed, along with six other species of sharks, under the CMS Memorandum of Understanding on the Conservation of Migratory Sharks.[84] In 1998, the Philippines banned all fishing, selling, importing, and exporting of whale sharks for commercial purposes,[85] followed by India in May 2001[86] and Taiwan in May 2007.[87]

In 2010, the Gulf of Mexico oil spill resulted in 4,900,000 barrels (780,000 m3) of oil flowing into an area south of the Mississippi River Delta, where one-third of all whale shark sightings in the northern part of the gulf have occurred in recent years. Sightings confirmed that the whale sharks were unable to avoid the oil slick, which was situated on the surface of the sea where the whale sharks feed for several hours at a time. No dead whale sharks were found.[88]

This species was also added to Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) in 2003 to regulate the international trade of live specimens and its parts.[89]

It was reported in 2014 that hundreds of whale sharks were illegally killed every year in China for their fins, skins, and oil.[90]

In captivity

Aquarium photograph of a whale shark in profile with human-shaped shadows in foreground
A whale shark in the Georgia Aquarium

The whale shark is popular in the few public aquariums that keep it, but its large size means that a very large tank is required and it has specialized feeding requirements.[91] Their large size and iconic status have also fueled an opposition to keeping the species in captivity, especially after the early death of some whale sharks in captivity and certain Chinese aquariums keeping the species in relatively small tanks.[92][93]

Whale sharks at the Okinawa Churaumi Aquarium, Japan

The first attempt at keeping whale sharks in captivity was in 1934 when an individual was kept for about four months in a netted-off natural bay in Izu, Japan.[94] The first attempt of keeping whale sharks in an aquarium was initiated in 1980 by the Okinawa Churaumi Aquarium (then known as Okinawa Ocean Expo Aquarium) in Japan.[91] Since 1980, several have been kept at Okinawa, mostly obtained from incidental catches in coastal nets set by fishers (none after 2009), but two were strandings. Several of these were already weak from the capture/stranding and some were released,[91] but initial captive survival rates were low.[93] After the initial difficulties in maintaining the species had been resolved, some have survived long-term in captivity.[91] The record for a whale shark in captivity is an individual that, as of 2021, has lived for more than 26 years in the Okinawa Churaumi Aquarium from Okinawa Ocean Expo Aquarium.[95][91] Following Okinawa, Osaka Aquarium started keeping whale sharks and most of the basic research on the keeping of the species was made at these two institutions.[96]

Since the mid-1990s, several other aquariums have kept the species in Japan (Kagoshima Aquarium, Kinosaki Marine World, Notojima Aquarium, Oita Marine Palace Aquarium, and Yokohama Hakkeijima Sea Paradise), South Korea (Aqua Planet Jeju), China (Chimelong Ocean Kingdom, Dalian Aquarium, Guangzhou Aquarium in Guangzhou Zoo, Qingdao Polar Ocean World and Yantai Aquarium), Taiwan (National Museum of Marine Biology and Aquarium), India (Thiruvananthapuram Aquarium) and Dubai (Atlantis, The Palm), with some maintaining whale sharks for years and others only for a very short period.[94] The whale shark kept at Dubai's Atlantis, The Palm was rescued from shallow waters in 2008 with extensive abrasions to the fins and after rehabilitation it was released in 2010, having lived 19 months in captivity.[97][98] Marine Life Park in Singapore had planned on keeping whale sharks but scrapped this idea in 2009.[99][100]

Outside Asia, the first and so far only place to keep whale sharks is Georgia Aquarium in Atlanta, United States.[94] This is unusual because of the comparatively long transport time and complex logistics required to bring the sharks to the aquarium, ranging between 28 and 36 hours.[96] Georgia keeps two whale sharks: two males, Taroko and Yushan, who both arrived in 2007.[101] Two earlier males at Georgia Aquarium, Ralph and Norton, both died in 2007.[93] Trixie died in 2020. Alice died in 2021. Georgia's whale sharks were all imported from Taiwan and were taken from the commercial fishing quota for the species, usually used locally for food.[96][102] Taiwan closed this fishery entirely in 2008.[102]

Human culture

Snorkeling with whale shark near Isla Mujeres (Mexico) 30 August 2011

In Madagascar, whale sharks are called marokintana in Malagasy, meaning "many stars", after the appearance of the markings on the shark's back.[103]

In the Philippines, it is called butanding and balilan.[104] The whale shark is featured on the reverse of the Philippine 100-peso bill. By law snorkelers must maintain a distance of 4 ft (1.2 m) from the sharks and there is a fine and possible prison sentence for anyone who touches the animals.[105]

Whale sharks are also known as jinbei-zame in Japan (because the markings resemble patterns typically seen on jinbei); gurano bintang (roughly "star from the East"[106]) in Indonesia; and ca ong (literally "sir fish") in Vietnam.[107]

The whale shark is also featured on the latest 2015–2017 edition of the Maldivian 1000 rufiyaa banknote, along with the green turtle.

See also

For a topical guide, see Outline of sharks.

References

  1. ^ "Rhincodon typus in the Paleobiology Database". Fossilworks. Archived from the original on 27 May 2022. Retrieved 17 December 2021.
  2. ^ a b c d Pierce, S.J.; Norman, B. (2016). "Rhincodon typus". IUCN Red List of Threatened Species. 2016: e.T19488A2365291. doi:10.2305/IUCN.UK.2016-1.RLTS.T19488A2365291.en. Retrieved 19 November 2021.
  3. ^ "Appendices | CITES". cites.org. Retrieved 14 January 2022.
  4. ^ a b Smith, Andrew (5 November 1828). "Descriptions of New or imperfectly known Objects of the Animal Kingdom, found in the South of Africa". The South African Commercial Advertiser. Vol. 3, no. 145 – via Center for Research Libraries Document Delivery System. Reprinted in Penrith (1972).
  5. ^ Penrith, M. J. (1972). "Earliest Description and Name for the Whale Shark". Copeia. 1972 (2): 362. doi:10.2307/1442501. JSTOR 1442501.
  6. ^ a b Melville, R. V. (1981). "Opinion 1278. The Generic Name Rhincodon A. Smith, 1829 (Pisces): Conserved". The Bulletin of Zoological Nomenclature. 41 (4): 215–217.
  7. ^ Smith, Andrew (1829). "Contributions to the Natural History of South Africa, &c". The Zoological Journal. 4: 443–444.
  8. ^ a b c d e McClain CR, Balk MA, Benfield MC, Branch TA, Chen C, Cosgrove J, Dove ADM, Gaskins LC, Helm RR, Hochberg FG, Lee FB, Marshall A, McMurray SE, Schanche C, Stone SN, Thaler AD. 2015. "Sizing ocean giants: patterns of intraspecific size variation in marine megafauna". PeerJ 3:e715 doi:10.7717/peerj.715.
  9. ^ a b c d Hsu, Hua Hsun; Joung, Shoou Jeng; Hueter, Robert E.; Liu, Kwang Ming (2014). "Age and growth of the whale shark (Rhincodon typus) in the north-western Pacific". Marine and Freshwater Research. 65 (12): 1145. doi:10.1071/MF13330. ISSN 1323-1650.
  10. ^ a b c d e f Colman, J. G. Froese, Ranier; Pauly, Daniel (eds.). "Rhincodon typus". FishBase. Retrieved 17 September 2006.
  11. ^ a b c d e f Perry, Cameron T.; Figueiredo, Joana; Vaudo, Jeremy J.; Hancock, James; Rees, Richard; Shivji, Mahmood (2018). "Comparing length-measurement methods and estimating growth parameters of free-swimming whale sharks (Rhincodon typus) near the South Ari Atoll, Maldives". Marine and Freshwater Research. 69 (10): 1487. doi:10.1071/MF17393. ISSN 1323-1650.
  12. ^ Martin, R. Aidan. "Rhincodon or Rhiniodon? A Whale Shark by Any Other Name". ReefQuest Centre for Shark Research.
  13. ^ Brunnschweiler, J. M.; Baensch, H.; Pierce, S. J.; Sims, D. W. (3 February 2009). "Deep-diving behaviour of a whale shark Rhincodon typus during long-distance movement in the western Indian Ocean". Journal of Fish Biology. 74 (3): 706–14. Bibcode:2009JFBio..74..706B. doi:10.1111/j.1095-8649.2008.02155.x. PMID 20735591.
  14. ^ a b c Colman, J. G. (1997). "A review of the biology and ecology of the whale shark". Journal of Fish Biology. 51 (6): 1219–1234. Bibcode:1997JFBio..51.1219C. doi:10.1111/j.1095-8649.1997.tb01138.x. ISSN 1095-8649. PMID 29991171.
  15. ^ a b Norman, Brad (2002). CITES identification manual. Environment Australia. ISBN 0-642-54900-1. OCLC 54364165.
  16. ^ "Whale Sharks, Rhincodon typus". MarineBio.org. Retrieved 17 May 2018.
  17. ^ Kaikini, A. S.; Ramamohana Rao, V.; Dhulkhed, M. H. (1959). "A note on the whale shark Rhincodon typus Smith, stranded off Mangalore". Central Marine Fisheries Research Unit, Mangalore.
  18. ^ Compagno, L. J. V. "Species Fact Sheet, Rhincodon typus". Food and Agriculture Organization of the United Nations. Retrieved 19 September 2006.
  19. ^ Tomita, Taketeru; Murakumo, Kiyomi; Komoto, Shinya; Dove, Alistair; Kino, Masakatsu; Miyamoto, Kei; Toda, Minoru (29 June 2020). "Armored eyes of the whale shark". PLOS ONE. 15 (6): e0235342. Bibcode:2020PLoSO..1535342T. doi:10.1371/journal.pone.0235342. ISSN 1932-6203. PMC 7323965. PMID 32598385.
  20. ^ McGreevy, Nora. "Whale Sharks Have Tiny Teeth on Their Eyeballs". Smithsonian Magazine.
  21. ^ Womersley, Freya; Hancock, James; Perry, Cameron T.; Rowat, David (February 2021). "Wound-healing capabilities of whale sharks (Rhincodon typus) and implications for conservation management". Conservation Physiology. 9 (1): coaa120. doi:10.1093/conphys/coaa120. PMC 7859907. PMID 33569175.
  22. ^ Read, Timothy D.; Petit, Robert A.; Joseph, Sandeep J.; Alam, Md. Tauqeer; Weil, M. Ryan; Ahmad, Maida; Bhimani, Ravila; Vuong, Jocelyn S.; Haase, Chad P. (December 2017). "Draft sequencing and assembly of the genome of the world's largest fish, the whale shark: Rhincodon typus Smith 1828". BMC Genomics. 18 (1): 532. doi:10.1186/s12864-017-3926-9. ISSN 1471-2164. PMC 5513125. PMID 28709399.
  23. ^ "How whale shark rhodopsin evolved to see, in the deep blue sea". ScienceDaily.
  24. ^ "Whale sharks see in dark due to mutation behind blindness in humans". The Japan Times. 9 April 2023.
  25. ^ White, Caleb (3 April 2023). "Whale Sharks Can See in the Dark Due to Genetic Mutation; Light-Sensing Pigment in Retina Works in Dim Environment". Science Times.
  26. ^ Yamaguchi, Kazuaki; Koyanagi, Mitsumasa; Sato, Keiichi; Terakita, Akihisa; Kuraku, Shigehiro (28 March 2023). "Whale shark rhodopsin adapted to deep-sea lifestyle by a substitution associated with human disease". Proceedings of the National Academy of Sciences. 120 (13): e2220728120. Bibcode:2023PNAS..12020728Y. doi:10.1073/pnas.2220728120. ISSN 0027-8424. PMC 10068813. PMID 36943890.
  27. ^ "Mutation behind night blindness in humans helps whale sharks see in the dark". www.science.org. Retrieved 26 March 2023.
  28. ^ Yamaguchi, Kazuaki; Koyanagi, Mitsumasa; Sato, Keiichi; Terakita, Akihisa; Kuraku, Shigehiro (1 October 2021). "Whale shark rhodopsin adapted to its vertically wide-ranging lifestyle": 2021.10.01.462724. doi:10.1101/2021.10.01.462724. S2CID 238261190 – via bioRxiv. {{cite journal}}: Cite journal requires |journal= (help)
  29. ^ a b c d e Meekan, Mark G.; Taylor, Brett M.; Lester, Emily; Ferreira, Luciana C.; Sequeira, Ana M. M.; Dove, Alistair D. M.; Birt, Matthew J.; Aspinall, Alex; Brooks, Kim; Thums, Michele (2020). "Asymptotic Growth of Whale Sharks Suggests Sex-Specific Life-History Strategies". Frontiers in Marine Science. 7. doi:10.3389/fmars.2020.575683. ISSN 2296-7745. S2CID 221712078.
  30. ^ a b Stevens, J. D. (1 March 2007). "Whale shark (Rhincodon typus) biology and ecology: A review of the primary literature". Fisheries Research. Whale Sharks: Science, Conservation and Management. 84 (1): 4–9. Bibcode:2007FishR..84....4S. doi:10.1016/j.fishres.2006.11.008. ISSN 0165-7836.
  31. ^ a b c Rowat, D.; Brooks, K. S. (2012). "A review of the biology, fisheries and conservation of the whale shark Rhincodon typus". Journal of Fish Biology. 80 (5): 1019–1056. Bibcode:2012JFBio..80.1019R. doi:10.1111/j.1095-8649.2012.03252.x. ISSN 1095-8649. PMID 22497372. S2CID 7243391.
  32. ^ Norman, Bradley M.; Stevens, John D. (1 March 2007). "Size and maturity status of the whale shark (Rhincodon typus) at Ningaloo Reef in Western Australia". Fisheries Research. Whale Sharks: Science, Conservation and Management. 84 (1): 81–86. Bibcode:2007FishR..84...81N. doi:10.1016/j.fishres.2006.11.015. ISSN 0165-7836.
  33. ^ a b Wintner, Sabine P. (2000). "Preliminary Study of Vertebral Growth Rings in the Whale Shark, Rhincodon typus, from the East Coast of South Africa". Environmental Biology of Fishes. 59 (4): 441–451. Bibcode:2000EnvBF..59..441W. doi:10.1023/A:1026564707027. S2CID 20461057.
  34. ^ a b c d Ong, Joyce J. L.; Meekan, Mark G.; Hsu, Hua Hsun; Fanning, L. Paul; Campana, Steven E. (6 April 2020). "Annual Bands in Vertebrae Validated by Bomb Radiocarbon Assays Provide Estimates of Age and Growth of Whale Sharks". Frontiers in Marine Science. 7. doi:10.3389/fmars.2020.00188. hdl:20.500.11815/2381.
  35. ^ Rohner, C. A.; Richardson, A. J.; Marshall, A. D.; Weeks, S. J.; Pierce, S. J. (2011). "How large is the world's largest fish? Measuring whale sharks Rhincodon typus with laser photogrammetry". Journal of Fish Biology. 78 (1): 378–385. Bibcode:2011JFBio..78..378R. doi:10.1111/j.1095-8649.2010.02861.x. PMID 21235570. S2CID 6882935.
  36. ^ Whitehead, Darren A.; Ayres, Kathryn A.; Gayford, Joel H.; Ketchum, James T.; Galván-Magana, Felipe; Christiansen, Fredrik (5 July 2022). "Aerial photogrammetry of whale sharks (Rhincodon typus) in the Bay of La Paz, using an unoccupied aerial vehicle". Marine Biology. 169 (7): 94. doi:10.1007/s00227-022-04085-0. ISSN 1432-1793.
  37. ^ Wright, E. Perceval (2011). Six months at the Seychelles: letter to A. Searle Hart, LL. D., S.F.T.C.D. British Library, Historical Print Editions. ISBN 9781241491611. OCLC 835888086.
  38. ^ Smith, H. M. (13 November 1925). "A Whale Shark (Rhineodon) in the Gulf of Siam". Science. 62 (1611): 438. Bibcode:1925Sci....62..438S. doi:10.1126/science.62.1611.438. ISSN 0036-8075. PMID 17732228.
  39. ^ Gudger, E. W. (1938). "Whale Sharks Rammed by Ocean Vessels: How These Sluggish Leviathans Aid in Their Own Destruction". New England Naturalist. 1–15. New England Museum of Natural History: Boston Society of Natural History. OCLC 1759776.
  40. ^ Maniguet, Xavier (1992). The Jaws of Death: Shark as Predator, Man as Prey. HarperCollins Publishers Limited. ISBN 978-0-00-219960-5.
  41. ^ Eckert, Scott A.; Stewart, Brent S. (1 February 2001). "Telemetry and Satellite Tracking of Whale Sharks, Rhincodon Typus, in the Sea of Cortez, Mexico, and the North Pacific Ocean". Environmental Biology of Fishes. 60 (1): 299–308. Bibcode:2001EnvBF..60..299E. doi:10.1023/A:1007674716437. ISSN 1573-5133. S2CID 22173382.
  42. ^ Katkar, B.N. (1996). "Turtles and whale shark landed along ratnagiri coast, maharashtra". Marine Fisheries Information Service. 141: 20.
  43. ^ Venkatesan, V; Ramamurthy, N; Boominathan, N; Gandhi, A (2008). "Stranding of a whale shark, Rhincodon typus (smith) at Pamban, Gulf of Mannar" (PDF). Marine Fisheries Information Service. 198: 19–22.
  44. ^ Borrell, Asunción; Aguilar, Alex; Gazo, Manel; Kumarran, R. P.; Cardona, Luis (2011). "Stable isotope profiles in whale shark (Rhincodon typus) suggest segregation and dissimilarities in the diet depending on sex and size". Environmental Biology of Fishes. 92 (4): 559–567. Bibcode:2011EnvBF..92..559B. doi:10.1007/s10641-011-9879-y. ISSN 0378-1909. S2CID 37683420.
  45. ^ Hasan, Saad (10 February 2012). "Experts to cut up 40.1-foot long whale shark today". The Express Tribune.
  46. ^ a b c d Pierce, S.J.; Norman, Brad (18 March 2016). "Whale Shark". IUCN Red List of Threatened Species. IUCN. Retrieved 28 November 2020.
  47. ^ Pain, Stephanie (31 May 2022). "Call of the deep". Knowable Magazine. Annual Reviews. doi:10.1146/knowable-052622-3. Retrieved 6 June 2022.
  48. ^ Public Library of Science (18 November 2015). "Whale sharks in Gulf of Mexico swim near the surface, take deep dives". phys.org. Retrieved 6 June 2022.
  49. ^ Howard, Brian C. (28 June 2016). "Whale Sharks Move in Mysterious Ways: Watch Them Online". National Geographic Society. Archived from the original on 28 June 2016. Retrieved 12 August 2016.
  50. ^ "Drive to conserve whale shark". The Hindu. 30 August 2017 – via www.thehindu.com.
  51. ^ Kaushik, Himanshu (30 August 2014). "Whale sharks found off Gujarat coast no expats, they are Indian". The Times of India. Retrieved 12 May 2016.
  52. ^ Dekker, Stefanie (17 July 2020). "'What a privilege': Swimming with endangered whale sharks in Qatar". Aljazeera. Retrieved 28 July 2020.
  53. ^ de la Parra Venegas, Rafael; Hueter, Robert; Cano, Jaime González; Tyminski, John; Remolina, José Gregorio; Maslanka, Mike; Ormos, Andrea; Weigt, Lee; Carlson, Bruce; Dove, Alistair (29 April 2011). "An Unprecedented Aggregation of Whale Sharks, Rhincodon typus, in Mexican Coastal Waters of the Caribbean Sea". PLOS ONE. 4. 6 (4): e18994. Bibcode:2011PLoSO...618994D. doi:10.1371/journal.pone.0018994. PMC 3084747. PMID 21559508.
  54. ^ Dove, Alistair (27 January 2015), Yucatan Whale Sharks Swimming in Troubled Waters, archived from the original on 7 November 2017
  55. ^ Clingham, Elizabeth; Brown, Judith; Henry, Leeann; Beard, Annalea; Dove, Alistair D (2016). Evidence that St. Helena island is an important multi-use habitat for whale sharks, Rhincodon typus , with the first description of putative mating in this species. doi:10.7287/peerj.preprints.1885v1. OCLC 8162956757.
  56. ^ "Attempted Whale Shark Mating Caught on Camera for the First Time in History". livescience.com. 24 June 2019.
  57. ^ Joung, Shoou-Jeng; et al. (July 1996). "The whale shark, Rhincodon typus, is a livebearer: 300 embryos found in one 'megamamma' supreme". Environ. Biol. Fish. 46 (3): 219–223. Bibcode:1996EnvBF..46..219J. doi:10.1007/BF00004997. S2CID 22250254.
  58. ^ Clark, Eugenie. "Frequently Asked Questions". Sharklady. Archived from the original on 5 March 2001. Retrieved 26 September 2006.
  59. ^ Schmidt, Jennifer V.; Chen, Chien-Chi; Sheikh, Saad I.; Meekan, Mark G.; Norman, Bradley M.; Joung, Shoou-Jeng (4 August 2010). "Paternity analysis in a litter of whale shark embryos". Endangered Species Research. 12 (2): 117–124. doi:10.3354/esr00300.
  60. ^ "Tiny whale shark rescued – World news – World environment". Associated Press via NBC News. 2009.
  61. ^ "St Helena whale sharks cause stir in Atlanta". South Atlantic Media Services. 14 November 2013. Archived from the original on 18 May 2023. Retrieved 12 May 2016.
  62. ^ "'Largest number in years': Over 100 new whale sharks spotted in Donsol". Rappler.com. 30 August 2019. Retrieved 4 March 2020.
  63. ^ Hearn, Alex R.; Green, J.; Román, M. H.; Acuña-Marrero, D.; Espinoza, E.; Klimley, A. P. (October 2016). "Adult female whale sharks make long-distance movements past Darwin Island (Galapagos, Ecuador) in the Eastern Tropical Pacific". Marine Biology. 163 (10). doi:10.1007/s00227-016-2991-y. ISSN 0025-3162.
  64. ^ a b Acuña-Marrero, David; Jiménez, Jesús; Smith, Franz; Doherty, Paul F.; Hearn, Alex; Green, Jonathan R.; Paredes-Jarrín, Jules; Salinas-de-León, Pelayo (31 December 2014). Klimley, A. Peter (ed.). "Whale Shark (Rhincodon typus) Seasonal Presence, Residence Time and Habitat Use at Darwin Island, Galapagos Marine Reserve". PLOS ONE. 9 (12): e115946. doi:10.1371/journal.pone.0115946. ISSN 1932-6203. PMC 4281130. PMID 25551553.
  65. ^ Morelle, Rebecca (17 November 2008). "Shark-cam captures ocean motion". BBC News. Retrieved 30 July 2009.
  66. ^ a b Jurassic Shark (2000) documentary by Jacinth O'Donnell; broadcast on Discovery Channel, 5 August 2006
  67. ^ Motta, Philip J.; et al. (2010). "Feeding anatomy, filter-feeding rate, and diet of whale sharks Rhincodon typus during surface ram filter feeding off the Yucatan Peninsula, Mexico" (PDF). Zoology. 113 (4): 199–212. Bibcode:2010Zool..113..199M. doi:10.1016/j.zool.2009.12.001. PMID 20817493.
  68. ^ a b Martin, R. Aidan. "Elasmo Research". ReefQuest. Retrieved 17 September 2006.
  69. ^ "Whale shark". Ichthyology at the Florida Museum of Natural History. 11 May 2017. Archived from the original on 5 September 2006. Retrieved 17 September 2006.
  70. ^ Schmidt, Jennifer V. (4 December 2010). "Whale Sharks are BIG eaters!". The Shark Research Institute. Archived from the original on 5 May 2015. Retrieved 24 April 2016.
  71. ^ "Rhincodon typus (whale shark)". Animal Diversity Web. Retrieved 1 September 2016.
  72. ^ Yong, Mila Mi Hua; Leistenschneider, Clara; Miranda, Joni Anne; Paler, Maria Kristina; Legaspi, Christine; Germanov, Elitza; Araujo, Gonzalo; Burkhardt-Holm, Patricia; Erni-Cassola, Gabriel (2021). "Microplastics in fecal samples of whale sharks (Rhincodon typus) and from surface water in the Philippines". Microplastics and Nanoplastics. 1 (1): 17. doi:10.1186/s43591-021-00017-9. PMC 8475362. PMID 34939039.
  73. ^ a b Pictures of the Day: Tuesday, Aug. 04, 2009. Time magazine, "A 40-foot whale shark and a brave snorkeler swim off the South African coast."
  74. ^ Russell, Mark 'Crowley' (9 April 2024). "Scuba diver fined for pulling whale shark tail in Thailand". DIVE Magazine. Retrieved 11 July 2024.
  75. ^ "Whale Sharks ⋅ Saint Helena Island Info ⋅ About St Helena, in the South Atlantic Ocean". sainthelenaisland.info. Retrieved 11 July 2024.
  76. ^ "Government of Western Australia, Department of Fisheries. Whale Shark" (PDF).
  77. ^ Whitehead, Darren Andrew (2014) Establishing a quantifiable model of whale shark avoidance behaviours to anthropogenic impacts in tourism encounters to inform management actions, University of Hertfordshire.
  78. ^ Reporter, Jess Thomson Science (30 November 2022). "Man filmed riding world's biggest shark in tourism stunt". Newsweek. Retrieved 11 July 2024.
  79. ^ "Friend of the Sea Sustainable Whale Shark Watching Standard" (PDF).
  80. ^ Compagno, Leonard J. V. (26 April 2002). Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date: Bullhead, Mackerel and Carpet Sharks. Vol. 2. Food & Agriculture Organization of the United Nations (FAO). ISBN 978-92-5-104543-5.
  81. ^ Hawes, Craig (2 April 2013) Snorkelling with whale sharks in Djibouti. gulfnews.com
  82. ^ Womersley, Freya C.; et al. (2022). "Global collision-risk hotspots of marine traffic and the world's largest fish, the whale shark". Proceedings of the National Academy of Sciences of the United States of America. 119 (20): e2117440119. Bibcode:2022PNAS..11917440W. doi:10.1073/pnas.2117440119. hdl:10754/676739. PMC 9171791. PMID 35533277.
  83. ^ Duffy, Clinton A. J.; Francis, Malcolm; Dunn, M. R.; Finucci, Brit; Ford, Richard; Hitchmough, Rod; Rolfe, Jeremy (2018). Conservation status of New Zealand chondrichthyans (chimaeras, sharks and rays), 2016 (PDF). Wellington, New Zealand: Department of Conservation. p. 11. ISBN 9781988514628. OCLC 1042901090.
  84. ^ "Memorandum of understanding on the conservation of migratory sharks" (PDF). Convention on migratory species. p. 10. Retrieved 13 February 2012.
  85. ^ Whale Sharks Receive Protection in the Philippines Archived 16 February 2020 at the Wayback Machine. hayop.0catch.com. 27 March 1998
  86. ^ National Regulations on Whale Shark fishing. Department of Sustainability, Environment, Water, Population and Communities.
  87. ^ COA bans fishing for whale sharks. Taipei Times, 27 May 2007, p. 4.
  88. ^ Handwerk, Brian (24 September 2010) Whale Sharks Killed, Displaced by Gulf Oil? National Geographic News.
  89. ^ Whale shark. cites.org
  90. ^ Hilton/Greenpeace, Paul (5 February 2014). "Hundreds of sharks killed in China". ABC News.
  91. ^ a b c d e Matsumoto; Toda; Matsumoto; Ueda; Nakazato; Sato; Uchida (2017). "Notes on Husbandry of Whale Sharks, Rhincodon typus, in Aquaria". In Smith, Mark; Warmolts; Thoney; Hueter; Murray; Ezcurra (eds.). The Elasmobranch Husbandry Manual II. Ohio Biological Survey. pp. 15–22. ISBN 9780867271676. OCLC 1001957014.
  92. ^ "Whale Shark's Death Sparks Debate". wsbtv. 30 November 2007. Archived from the original on 25 March 2010.
  93. ^ a b c Moore, M. (25 October 2010). "Conservationists round on Chinese whale shark aquarium". The Telegraph. Archived from the original on 12 January 2022. Retrieved 8 November 2017.
  94. ^ a b c Mollet, H. (September 2012). "Whale Shark Rhincodon typus Smith, 1828 in Captivity". Archived from the original on 24 March 2014. Retrieved 8 November 2017.
  95. ^ "令和二年度 沖縄美ら海水族館 年報" (PDF). Okinawa Churaumi Aquarium. Retrieved 28 May 2021.
  96. ^ a b c Schreiber, C; Coco, C (2017). "Husbandry of Whale Sharks". In Smith, Mark; Warmolts; Thoney; Hueter; Murray; Ezcurra (eds.). The Elasmobranch Husbandry Manual II. Ohio Biological Survey. pp. 87–98. ISBN 9780867271676. OCLC 1001957014.
  97. ^ "Dubai hotel releases whale shark back into the wild". Associated Press. 20 March 2010. Archived from the original on 7 August 2015.
  98. ^ Bennett; Kaiser; Selvan; Hueter; Tyminski; Lötter (2017). "Rescue, Rehabilitation and Release of a Whale Shark, Rhincodon typus, in the Arabian Gulf". In Smith, Mark; Warmolts; Thoney; Hueter; Murray; Ezcurra (eds.). The Elasmobranch Husbandry Manual II. Ohio Biological Survey. pp. 229–235. ISBN 9780867271676. OCLC 1001957014.
  99. ^ Chua, G. (16 May 2009). "No whale sharks at Sentosa IR". Wild Singapore News. Retrieved 8 November 2017.
  100. ^ "Resorts World considering alternatives to whale shark exhibit". AsianOne Travel. 16 May 2009. Archived from the original on 9 June 2013.
  101. ^ "Aquarium gains two new whale sharks". CNN. 1 June 2007. Archived from the original on 3 June 2007. Retrieved 1 June 2007.
  102. ^ a b Sundquist, T. (18 September 2013). "Transporting the World's Largest Fish: A Whale [Shark] of a Task". Promega Connections. Archived from the original on 8 March 2021. Retrieved 8 November 2017.
  103. ^ Briggs, Helen (17 May 2018). "Madagascar emerges as whale shark hotspot". BBC News. Retrieved 12 September 2018.
  104. ^ Ocean Ambassadors – Sharks. Oneocean.org. Retrieved 23 May 2013.
  105. ^ Cannon, Marisa (21 July 2015). "Swimming with whale sharks in the Philippines". cnn.com. CNN. Retrieved 16 August 2015.
  106. ^ "Rewards from Gurano Bintang Shark Conservation". 5 February 2022.
  107. ^ "Whale Shark". Discovery.com. 5 September 2013. Archived from the original on 4 March 2009. Retrieved 3 November 2011.

Further reading

Wikimedia Commons has media related to Whale sharks.
Wikispecies has information related to Whale sharks.

Retrieved from "https://en.wikipedia.org/w/index.php?title=Whale_shark&oldid=1264471238"