Giant tortoise

Giant tortoises are any of several species of various large land tortoises, which include a number of extinct species, as well as a few living species formerly common on the islands of the western Indian Ocean and on the Galápagos Islands.[1]

An Aldabra giant tortoise, an example of a giant tortoise.

HistoryEdit

As of February 2021, giant tortoises are found on two remote groups of tropical islands: Aldabra Atoll and Fregate Island in the Seychelles and the Galápagos Islands in Ecuador. These tortoises can weigh as much as 417 kg (919 lb) and can grow to be 1.3 m (4 ft 3 in) long. Giant tortoises originally made their way to islands from the mainland via oceanic dispersal. Tortoises are aided in such dispersal by their ability to float with their heads up and to survive for up to six months without food or fresh water.[2] Giant tortoises were once all placed in a single genus (often referred to as Testudo or Geochelone), but more recent studies have shown that giant tortoises represent several distinct lineages that are not closely related to one another.[2] These lineages appear to have developed large size independently and, as a result, giant tortoises are polyphyletic. For example, the Aldabra Atoll (Aldabrachelys) and Mascarenes (Cylindraspis) giant tortoises are related to Malagasy tortoises (Asterochelys) while the Galapagos giant tortoises are related to South American mainland tortoises, particularly the Chaco tortoise (Chelonoidis chiliensis). Giant tortoises are classified into several distinct genera, including Aldabrachelys, Centrochelys (in part, often excluding the extant African spurred tortoise (Centrochelys sulcata)), Chelonoidis (in part), †Cylindraspis,Hesperotestudo,Megalochelys andTitanochelon.

The phenomenon of animal species evolving in cache to unusually large size on islands (in comparison to continental relatives) is known as island gigantism or insular gigantism. This may occur due to factors such as relaxed predation pressure, competitive release, or as an adaptation to increased environmental fluctuations on islands.[3][4] However, giant tortoises are no longer considered to be classic examples of island gigantism, as similarly massive tortoises are now known to have once been widespread. Giant tortoises were formerly common (prior to the Quaternary extinctions) across the Cenozoic faunas of Eurasia, Africa and the Americas.[5][6]

Giant tortoises are notably absent from Australia and the South Pacific. However, extinct giant horned turtles (Meiolaniidae) likely filled a similar niche, with Late Pleistocene-Holocene meiolaniid species being known from Australia, New Caledonia, Lord Howe Island, Vanuatu, and the Fijian Archipelago. The identity of the Vanuatu meiolaniid has been controversial, however, with some studies concluding the remains actually belong to a giant tortoise, which are otherwise unknown from this region.[7] Older (Early Miocene) meiolaniids are also known from the St. Bathans fauna in New Zealand.

Table of extant and recently extinct island giant tortoises across the world. Taxonomy of extinct species follows Rhodin et al. (2015)[8] and taxonomy of extant species follows Rhodin et al. (2017) unless otherwise noted
Island Archipelago Region Genus Taxa
Eleuthera Lucayan Archipelago/Bahamas Caribbean Chelonoidis Chelonoidis alburyorum[9]
Andros Island Lucayan Archipelago/Bahamas Caribbean Chelonoidis Chelonoidis alburyorum[9]
Nassau Lucayan Archipelago/Bahamas Caribbean Chelonoidis Chelonoidis alburyorum[9]
Mayaguana Lucayan Archipelago/Bahamas Caribbean Chelonoidis Chelonoidis alburyorum[9]
Crooked Island Lucayan Archipelago/ Bahamas Caribbean Chelonoidis Chelonoidis alburyorum[9]
Gran Abaco Lucayan Archipelago/ Bahamas Caribbean Chelonoidis Chelonoidis alburyorum[10]
Cabo Verde Cape Verde Archipelago Atlantic Ocean Centrochelys Centrochelys atlantica
Bermuda Bermuda Atlantic Ocean Hesperotestudo Hesperotestudo bermudae[11]
Tenerife Canary Islands Atlantic Ocean Centrochelys Centrochelys burchardi
Cuba Greater Antilles Caribbean Chelonoidis Chelonoidis cubensis[12]
Malta Malta Mediterranean Centrochelys[a] Centrochelys robusta
Gran Canaria Canary Islands Atlantic Ocean Centrochelys Centrochelys vulcanica
Sombrero Island Lesser Antilles Caribbean Chelonoidis Chelonoidis sombrerensis
Mona Island Greater Antilles Caribbean Chelonoidis Chelonoidis monensis
Navassa Island Greater Antilles Caribbean Chelonoidis Chelonoidis sp. (extinct)
Cosmoledo Seychelles Indian Ocean Aldabrachelys Aldabrachelys sp. (extinct)[14]
Denis Seychelles Indian Ocean Aldabrachelys Aldabrachelys sp. (extinct)[14]
Assumption Seychelles Indian Ocean Aldabrachelys Aldabrachelys sp. (extinct)[14]
Astove Seychelles Indian Ocean Aldabrachelys Aldabrachelys sp. (extinct)[14]
Comoros Islands Comoros Islands Indian Ocean Aldabrachelys Aldabrachelys sp. (extinct)[14]
Grand Turk Lucayan Archipelago/Bahamas Caribbean Chelonoidis Chelonoidis alburyorum
Middle Caicos Lucayan Archipelago/Bahamas Caribbean Chelonoidis Chelonoidis alburyorum
Réunion Mascarene Islands Indian Ocean Cylindraspis Cylindraspis indica (Réunion giant tortoise)
Rodrigues Mascarene Islands Indian Ocean Cylindraspis Cylindraspis peltastes (Rodrigues domed giant tortoise)

Cylindraspis vosmaeri (Rodrigues saddle-backed giant tortoise)

Maurititus Mascarene Islands Indian Ocean Cylindraspis Cylindraspis inepta (Mauritius saddle-backed giant tortoise)

Cylindraspis triserrata (Mauritius domed giant tortoise)

Curaçao Lesser Antilles Caribbean Chelonoidis Chelonoidis sp.
Ibiza Balearic Islands Mediterranean Titanochelon Titanochelon sp.
Central granitic islands (Cerf,

Cousine, Frégate, Mahé,

Praslin, Round, and Silhouette)[b]

Seychelles Indian Ocean Aldabrachelys Aldabrachelys gigantea arnoldi (Arnold's giant tortoise or Seychelles saddle-backed giant tortoise)

Aldabrachelys gigantea daudinii (Daudin's giant tortoise)

Aldabrachelys gigantea hololissa (Seychelles giant tortoise or Seychelles domed giant tortoise)

Glorioso Islands Glorioso Islands Indian Ocean Aldabrachelys Aldabrachelys sp.
Aldabra Atoll Seychelles Indian Ocean Aldabrachelys Aldabrachelys gigantea gigantea (Aldabra giant tortoise)
Hispaniola Greater Antilles Caribbean Chelonoidis Chelonoidis marcanoi[12]
Madagascar Madagascar Indian Ocean Aldabrachelys Aldabrachelys abrupta (abrupt giant tortoise)[16]

Aldabrachelys grandidieri (Grandidier's giant tortoise)

San Cristóbal Island Galápagos Islands Pacific Ocean Chelonoidis Chelonoidis chathamensis (San Cristobal giant tortoise)
Isabela Island Galápagos Islands Pacific Ocean Chelonoidis Chelonoidis becki (Volcán Wolf giant tortoise)

Chelonoidis vicina (southern Isabela giant tortoise)

Santiago Island Galápagos Islands Pacific Ocean Chelonoidis Chelonoidis darwini (Santiago giant tortoise)
Santa Cruz Island Galápagos Islands Pacific Ocean Chelonoidis Chelonoidis donfaustoi (eastern Santa Cruz giant tortoise)

Chelonoidis porteri (western Santa Cruz giant tortoise)

Fernandina Island Galápagos Islands Pacific Ocean Chelonoidis Chelonoidis phantastica (Fernandina giant tortoise)
Pinta Island Galápagos Islands Pacific Ocean Chelonoidis Chelonoidis abingdonii (Pinta giant tortoise)
Floreana Island Galápagos Islands Pacific Ocean Chelonoidis Chelonoidis nigra (Floreana giant tortoise)
Pinzón Island Galápagos Islands Pacific Ocean Chelonoidis Chelonoidis duncanensis (Pinzón giant tortoise)
Española Island Galápagos Islands Pacific Ocean Chelonoidis Chelonoidis hoodensis (Española giant tortoise)
Santa Fe Island Galápagos Islands Pacific Ocean Chelonoidis Chelonoidis sp. (Santa Fe giant tortoise)
Rábida Island Galápagos Islands Pacific Ocean Chelonoidis Chelonoidis wallacei (Rábida giant tortoise)

Although often considered examples of island gigantism, prior to the arrival of Homo sapiens giant tortoises also occurred in non-island locales, as well as on a number of other, more accessible islands. During the Pleistocene, and mostly during the last 50,000 years, tortoises of the mainland of southern Asia (†Megalochelys atlas),[5] North America (†Hesperotestudo spp.)[5] and South America (Chelonoidis spp.),[6] Indonesia,[5] Madagascar (†Aldabrachelys)[5] and even the island of Malta[5]all became extinct. Giant tortoises (†Titanochelon) also inhabited mainland Europe until the Early Pleistocene (2.0 Mya).[13] The giant tortoises formerly of Africa died out somewhat earlier, during the Late Pliocene.[17] While the timing of the disappearances of various extinct giant tortoise species seems to correlate with the arrival of humans, direct evidence for human involvement in these extinctions is usually lacking; however, such evidence has been obtained in the case of the distantly-related giant meiolaniid turtle Meiolania damelipi in Vanuatu.[18][19] One interesting relic is the shell of an extinct giant tortoise found in a submerged sinkhole in Florida with a wooden spear piercing through it, carbon dated to 12,000 years ago.[20][better source needed] Today, only one of the species of the Indian Ocean survives in the wild; the Aldabra giant tortoise (two more are claimed to exist in captive or re-released populations, but some[vague] genetic studies have cast doubt on the validity of these as separate species)[citation needed] and 10 extant species in the Galápagos.

Life expectancyEdit

Giant tortoises are among the world's longest-living animals, with an average lifespan of 100 years or more.[21] The Madagascar radiated tortoise Tu'i Malila was 188 at her death in Tonga in 1965.[citation needed] Harriet (initially thought to be one of the three Galápagos tortoises brought back to England from Charles Darwin's Beagle voyage, but later shown to be from an island not even visited by Darwin) was reported by the Australia Zoo to be 176 years old when she died in 2006.[22] Also, on 23 March 2006, an Aldabra giant tortoise named Adwaita died at the Alipore Zoological Gardens in Kolkata. He was brought to the zoo in the 1870s from the estate of Lord Clive and is thought to have been around 255 years old when he died.[23][better source needed] Around the time of its discovery, they were caught for food in such large numbers that they became virtually extinct by 1900.[citation needed] Giant tortoises are now protected by strict conservation laws and are categorized as threatened species.

Aldabra giant tortoiseEdit

The Aldabra giant tortoise lives on the remote Aldabra Atoll, one of the Seychelles group of islands in the Indian Ocean.

Distribution and habitatEdit

 
An Aldabra giant tortoise cooling down in a freshwater pond on Curieuse, Seychelles

Aldabra giant tortoises have large dome-shaped shells in order to protect their delicate bodies that lie beneath their shells. They also have long necks in order to eat leaves from the higher branches of trees. The males, although not much bigger than the females, weigh nearly 100 kg (220 lbs) more. They move slowly and have small, thick legs and round, almost flat feet that assist them in walking on sand.

The Aldabra giant tortoise mainly inhabits grasslands and swamps on Aldabra Atoll's islands, which form a part of the Seychelles island chain in the Indian Ocean. In the past, they shared the islands with multiple other giant tortoise species, but many of them were hunted to extinction in the 1700s and 1800s.[citation needed] Despite the fact that they are usually found in regions of dense low-lying vegetation, they have been known to wander into areas with more sparse vegetation and rocks when food is scarce. They can also be seen resting in shaded areas or shallow pools of water in order to cool themselves on hot days.[23][better source needed] Aldabra giant tortoises tend to spend their lives grazing, but will cover surprising distances in search of food and have also been observed on bare rock and thin soil. They can drink from very shallow pools through their nostrils; the former genus Dipsochelys refers to this adaptation.[24][full citation needed]

Species and subspeciesEdit

Canary Islands giant tortoisesEdit

 
A restoration of the Tenerife giant tortoise (Centrochelys burchardi) in the Museo de la Naturaleza y el Hombre of Santa Cruz de Tenerife

Remains have been found in the Canary Islands of two extinct species of large tortoises: the Tenerife giant tortoise (Centrochelys burchardi) and the Gran Canaria giant tortoise (Centrochelys vulcanica). These species are believed to have died out due to volcanic eruptions.[25][full citation needed]

The Canary Islands tortoises were similar to those currently found in the Galapagos and Seychelles. C. burchardi had a larger shell, with a length of approximately 65 to 94 cm, while G. vulcanica had a 61 cm long shell.[25]

The remains date mostly to the Miocene. It is believed that these species survived until the Late Pleistocene.[25]

Galápagos giant tortoisesEdit

The closest living relative of the Galápagos giant tortoise is the small Chaco tortoise from South America, although it is not a direct ancestor. Scientists believe the first tortoises arrived to the Galápagos 2–3 million years ago by drifting 600 miles from the South American coast on vegetation rafts or on their own.[citation needed] They were already large animals before arriving in the Galápagos. Colonizing the easternmost islands of Española and San Cristóbal first, they then dispersed throughout the archipelago, eventually establishing about 16 separate populations on 10 of the largest of the Galápagos Islands. Currently, there are only 10 species of Galápagos giant tortoises left of the original 16 species. During the 16th and 17th centuries, the Galápagos were frequented by buccaneers preying on Spanish treasure ships. Filling a ship's hold with tortoises was an easy way to stock up on food, a tradition that was continued by whalers in the centuries that followed: "whaling skippers were almost lyrical in their praise of tortoise meat, terming it far more delicious than chicken, pork or beef". They said the meat of the giant tortoise was 'succulent meat and the oil from their bodies as pure as butter, but best of all, the giants could hibernate in a ship's damp for a year or more."[26][better source needed] The tortoises also conveniently held water in their necks that could be used as drinking water.

 
Lonesome George, the last known individual of the Pinta giant tortoise (C. abingdonii)

These buccaneers stocked giant tortoises not only because of their meat and oil[clarification needed] but because of these animals' ability to survive for six months to one year without food or water.[citation needed] Once buccaneers, whalers and fur sealers discovered that they could have fresh meat for their long voyages by storing live giant tortoises in the holds of their ships, massive exploitation of the species began. Tortoises were also exploited for their oil,[citation needed] which was used to light the lamps of Quito. Two centuries of exploitation resulted in the loss of between 100,000 and 200,000 tortoises. Three species have been extinct for some time, and a fourth species lost its last member, Lonesome George, in June 2012.[27] In February 2019, a tortoise species once thought to have been extinct since 1906, the Fernandina giant tortoise, was discovered on its namesake island in the Galápagos.[28] It is estimated that 20,000–25,000 wild tortoises live on the islands today.[27][better source needed]

Distribution and habitatEdit

Galápagos tortoises are herbivorous, feeding primarily on cactus pads, grasses, and native fruit. They drink large quantities of water when available that they can store in their bladders for long periods of time. There are two main types of shell among them, the saddle-backed shell and the domed shell. They both provide special adaption to different environments. The saddle-backed tortoises are the smallest Galápagos tortoises, but present a very long neck and pairs of legs. They live on arid zone and feed on cactus. The domed tortoises are bigger with shorter neck and legs, they are found in the more vegetated islands and feed on grass.[29] They spend an average of 16 hours a day resting. Their activity level is driven by ambient temperature and food availability. In the cool season, they are active at midday, sleeping in in the morning and afternoon. In the hot season, their active period is early morning and late afternoon, while midday finds them resting and trying to keep cool under the shade of a bush or half-submerged in muddy wallows.[citation needed]

Life cycleEdit

Tortoises breed primarily during the hot season from January to May; however, tortoises can be seen mating any month of the year. During the cool season (June to November), female tortoises migrate to nesting zones, which are generally located in low lands of the islands, to lay their eggs. A female can lay from 1–4 nests over a nesting season from June to December. She digs the hole with her hind feet, then lets the eggs drop down into the nest, and finally covers it again with her hind feet. The number of eggs ranges from 2 to 7 for saddle-backed tortoises to sometimes more than 20 to 25 eggs for domed tortoises. The eggs incubate from 110 to 175 days (incubation periods depend on the month the clutch was produced, with eggs laid early in the cool season requiring longer incubation periods than eggs laid at the end of the cool season, when the majority of their incubation will occur at the start of the hot season). After hatching, the young hatchlings remain in the nest for a few weeks before emerging out a small hole adjacent to the nest cap. Usually, the temperature of the nest influences on the sex of the hatchling. Warm temperatures would yield more females, while colder temperatures would yield more males.[citation needed]

SpeciesEdit

Mascarenes giant tortoisesEdit

The Mascarene Islands of Mauritius, Réunion and Rodrigues once harboured five species of giant tortoise, comprising two species occurring on Mauritius, another two on Rodrigues, and one on Réunion. The tortoises were unique to these islands and had gained a number of special adaptations in the absence of ground predators. They differed from any other giant tortoise species because of their modified jaws, reduced scales on the legs and shells averaging just 1mm thick. The shells of the giant tortoises were open-ended; the name Cylindraspis actually means "cylinder-shaped". This was a specific adaptation in response to the lack of predators, where thick, heavily armored shells were no longer necessary.[citation needed]

Around the 16th century, with human arrival and the subsequent introduction of domestic animals, particularly pigs, the tortoises were rapidly[vague][citation needed] hunted to extinction. Unfortunately, the thin shells were of no protection against these new invaders; rats, cats and pigs devoured the eggs and young and thousands were collected alive for provisioning ships. Sometimes they were even hunted for their oil,[citation needed] which was very valuable around that time because it provided a cure for many ailments, including scurvy.

On Mauritius, the giant tortoise disappeared from the main island by the end of the 17th century and the very last tortoises survived until the 1730s on the islets in the north. Around the late 1800s, large number of tortoise bones were discovered in the Mare aux Songes excavations.[citation needed] These resulted in the description of the two species of giant tortoise endemic to Mauritius, the Mauritius saddle-backed (Cylindraspis inepta) and the Mauritius domed (Cylindraspis triserrata).[30]

Today, the only remains from these five species are a number of fossil bones and shells, a few drawings of live animals and one stuffed Rodrigues saddle-backed giant tortoise in France's National Museum of Natural History.[31][better source needed]

SpeciesEdit

FootnotesEdit

  1. ^ Often treated as Centrochelys but Pérez-García et al. (2017) suggests it could pertain to Titanochelon.[13]
  2. ^ Exact geographic range of these subspecies prior to human arrival is unknown, especially as tortoises have been moved between islands, but populations of Aldabrachelys are known to have existed at one time on all of these islands.[15]

See alsoEdit

ReferencesEdit

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  2. ^ a b Le M, Raxworthy CJ, McCord WP, Mertz L (August 2006). "A molecular phylogeny of tortoises (Testudines: Testudinidae) based on mitochondrial and nuclear genes". Molecular Phylogenetics and Evolution. 40 (2): 517–31. doi:10.1016/j.ympev.2006.03.003. PMID 16678445.
  3. ^ Pritchard PC (1996). The Galápagos Tortoises: Nomenclatural and Survival Status. Lunenburg, MA: Chelonian Research Foundation. ISBN 978-0965354004.
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  12. ^ a b Turvey ST, Almonte J, Hansford J, Scofield RP, Brocca JL, Chapman SD (June 2017). "A new species of extinct Late Quaternary giant tortoise from Hispaniola". Zootaxa. 4277 (1): 1–16. doi:10.11646/zootaxa.4277.1.1. PMID 30308657.
  13. ^ a b Pérez-García A, Vlachos E, Arribas A (March 2017). "The last giant continental tortoise of Europe: A survivor in the Spanish Pleistocene site of Fonelas P-1". Palaeogeography, Palaeoclimatology, Palaeoecology. 470: 30–39. doi:10.1016/j.palaeo.2017.01.011.
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External linksEdit