Island gigantism
Island gigantism, or insular gigantism, is a biological phenomenon in which the size of an animal species isolated on an island increases dramatically in comparison to its mainland relatives. Island gigantism is one aspect of the more general "island effect" or "Foster's rule", which posits that when mainland animals colonize islands, small species tend to evolve larger bodies, and large species tend to evolve smaller bodies. Following the arrival of humans and associated introduced predators, many giant as well as other island endemics have become extinct. A similar size increase, as well as increased woodiness, has been observed in some insular plants.
Possible causes
Large mammalian carnivores are often absent on islands because of insufficient range or difficulties in over-water dispersal. In their absence, the ecological niches for large predators may be occupied by birds, reptiles or smaller carnivorans, which can then grow to larger-than-normal size. For example, on prehistoric Gargano Island in the Miocene-Pliocene Mediterranean, on islands in the Caribbean like Cuba, and on Madagascar and New Zealand, some or all apex predators were birds like eagles, falcons and owls, including some of the largest known examples of these groups. However, birds and reptiles generally make less efficient large predators than advanced carnivorans.Since small size usually makes it easier for herbivores to escape or hide from predators, the decreased predation pressure on islands can allow them to grow larger. Small herbivores may also benefit from the absence of competition from missing types of large herbivores.
Benefits of large size that have been suggested for island tortoises include decreased vulnerability to scarcity of food and/or water, through ability to survive for longer intervals without them, or ability to travel longer distances to obtain them. Periods of such scarcity may be a greater threat on oceanic islands than on the mainland.
Thus, island gigantism is usually an evolutionary trend resulting from the removal of constraints on the size of small animals related to predation and/or competition. Such constraints can operate differently depending on the size of the animal, however; for example, while small herbivores may escape predation by hiding, large herbivores may deter predators by intimidation. As a result, the complementary phenomenon of island dwarfism can also result from the removal of constraints related to predation and/or competition on the size of large herbivores. In contrast, insular dwarfism among predators more commonly results from the imposition of constraints associated with the limited prey resources available on islands. As opposed to island dwarfism, island gigantism is found in most major vertebrate groups and in invertebrates.
Territorialism may favor the evolution of island gigantism. A study on Anaho Island in Nevada determined that reptile species that were territorial tended to be larger on the island compared to the mainland, particularly in the smaller species. In territorial species, larger size makes individuals better able to compete to defend their territory. This gives additional impetus to evolution toward larger size in an insular population.
A further means of establishing island gigantism may be a founder effect operative when larger members of a mainland population are superior in their ability to colonize islands.
Island size plays a role in determining the extent of gigantism. Smaller islands generally accelerate the rate of evolution of changes in organism size, and organisms there evolve greater extremes in size.
Examples
Examples of island gigantism include:Mammals
Many rodents grow larger on islands, whereas carnivorans, proboscideans and artiodactyls usually become smaller.[Eulipotyphla]ns
[Rodent]s
[Lagomorph]s
[Primates]
[Carnivora]ns
Birds
Stem birds
[Ratites]
[Waterfowl]
[Pangalliformes]
[Gruiformes]
[Pigeons]
Birds of prey">Accipitriformes">Birds of prey
[Parrot]s
[Owl]s
[Caprimulgiformes]
[Passeriform]s
| Example | Binomial name | Native range | Current status | Continental relative |
Chatham raven | Corvus moriorum | Chatham Islands | Extinct | New Zealand raven |
Long-legged bunting | Emberiza alcoveri | Tenerife | Extinct | Cabanis's bunting |
| Giant nukupu'u | Hemignathus vorpalis | Hawaii | Extinct | Finches |
| Tasmanian superb fairywren | Malurus cyaneus cyaneus | Tasmania | Least Concern | Superb fairywren |
| Kangaroo Island superb fairywren | Malurus cyaneus ashbyi | Kangaroo Island | Least Concern | Superb fairywren |
| Stout-legged wren | Pachyplichas yaldwyni | South Island of New Zealand | Extinct | Other passeriforms |
Capricorn silvereye | Zosterops lateralis chlorocephalus | Capricorn and Bunker Group of the Australian Great Barrier Reef | Unknown | Silvereye |
Reptiles
[Pterosaur]s
[Iguanid]s
[Gecko]s
[Skink]s
Wall lizards">Lacertidae">Wall lizards
[Snake]s
Dubious examples
- The Komodo dragon of Flores and nearby islands, the largest extant lizard, and a similar giant monitor lizard from Timor have been regarded as examples of giant insular carnivores. Since islands tend to offer limited food and territory, their mammalian carnivores are usually smaller than continental ones. These cases involve ectothermic carnivores on islands too small to support much mammalian competition. However, these lizards are not as large as their extinct Australian relative megalania, and it has been proposed based on fossil evidence that the ancestors of these varanids first evolved their large size in Australia and then dispersed to Indonesia. If this is true, rather than being insular giants they would be viewed as examples of phyletic gigantism. Supporting this interpretation is evidence for a lizard in Pliocene India, Varanus sivalensis, comparable in size to komodoensis. Nevertheless, given that Australia is often described as the world's largest island and that the related megalania, the largest terrestrial lizard known in the fossil record, was restricted to Australia, the perception of the largest Australasian/Indonesian lizards as insular giants may still have some validity.
- Giant tortoises in the Galápagos Islands and the Seychelles, the largest extant tortoises, as well as extinct tortoises of the Mascarenes and Canary Islands, are often considered examples of island gigantism. However, during the Pleistocene, comparably sized or larger tortoises were present in Australia, southern Asia, Madagascar, North America and South America, and on a number of other, more accessible islands. In the late Pliocene they were also present in Africa.. The present situation of large tortoises being found only on remote islands appears to reflect that these islands were discovered by humans recently and have not been heavily populated, making their tortoises less subject to overexploitation.
Amphibians">Lissamphibia">Amphibians
[Arthropod]s
[Gastropod]s
Flora
In addition to size increase, island grass plants may also exhibit "insular woodiness". The most notable examples are the megaherbs of New Zealand's subantarctic islands. Increased leaf and seed size was also reported in some island species regardless of growth form.| Example | Binomial name | Native range | Current status | Continental relative |
Campbell Island carrot | Anisotome latifolia | Campbell and Auckland Islands | Unknown | Apiaceae |
Ross lily | Bulbinella rossii | Campbell and Auckland Islands | Unknown | New Zealand Maori lily |
Chatham Islands korokio | Corokia macrocarpa | Chatham Islands | Unknown | New Zealand korokio |
Black-eyed daisy | Damnamenia vernicosa | Auckland and Campbell Islands | Unknown | Astereae |
Cucumber tree | Dendrosicyos socotranus | Socotra | Vulnerable | Gourds |
Coco de mer | Lodoicea maldivica | Seychelles | Endangered | Borassoid palms |
| Pleurophyllum criniferum | Antipodes, Auckland and Campbell Islands | Unknown | Cineraria | |
Silver-leaf daisy | Pleurophyllum hookeri | Macquarie Island, Auckland and Campbell Islands | Unknown | Cineraria |
Campbell Island daisy | Pleurophyllum speciosum | Campbell and Auckland Islands | Unknown | Cineraria |
Chatham Islands nikau | Rhopalostylis aff. sapida | Chatham Islands | Unknown | Nikau |
Macquarie Island cabbage | Stilbocarpa polaris | Macquarie Island and New Zealand subantarctic islands | Vulnerable | Araliaceae |