Insular dwarfism

The skeleton of a dwarf elephant from the island of Crete

Insular dwarfism, a form of phyletic dwarfism,[1] is the process and condition of the reduction in size of large animals over a number of generations[lower-alpha 1] when their population's range is limited to a small environment, primarily islands. This natural process is distinct from the intentional creation of dwarf breeds, called dwarfing. This process has occurred many times throughout evolutionary history, with examples including dinosaurs, like Europasaurus, and modern animals such as elephants and their relatives. This process, and other "island genetics" artifacts, can occur not only on traditional islands, but also in other situations where an ecosystem is isolated from external resources and breeding. This can include caves, desert oases, isolated valleys and isolated mountains ("sky islands"). Insular dwarfism is one aspect of the more general "island rule", which posits that when mainland animals colonize islands, small species tend to evolve larger bodies, and large species tend to evolve smaller bodies.

Possible causes of insular dwarfism

There are several proposed explanations for the mechanism which produces such dwarfism.[3][4]

One is a selective process where only smaller animals trapped on the island survive, as food periodically declines to a borderline level. The smaller animals need fewer resources and smaller territories, and so are more likely to get past the break-point where population decline allows food sources to replenish enough for the survivors to flourish. Smaller size is also advantageous from a reproductive standpoint, as it entails shorter gestation periods and generation times.[3]

In the tropics, small size should make thermoregulation easier.[3]

Among herbivores, large size confers advantages in coping with both competitors and predators, so a reduction or absence of either would facilitate dwarfing; competition appears to be the more important factor.[4]

Among carnivores, the main factor is thought to be the size and availability of prey resources, and competition is believed to be less important.[4] In tiger snakes, insular dwarfism occurs on islands where available prey is restricted to smaller sizes than are normally taken by mainland snakes. Since prey size preference in snakes is generally proportional to body size, small snakes may be better adapted to take small prey.[5]

Dwarfism versus gigantism

The inverse process, wherein small animals breeding on isolated islands lacking the predators of large land masses may become much larger than normal, is called island gigantism. An excellent example is the dodo, the ancestors of which were normal-sized pigeons. There are also several species of giant rats, one still extant, that coexisted with both Homo floresiensis and the dwarf stegodons on Flores.

The process of insular dwarfing can occur relatively rapidly by evolutionary standards. This is in contrast to increases in maximum body size, which are much more gradual. When normalized to generation length, the maximum rate of body mass decrease during insular dwarfing was found to be over 30 times greater than the maximum rate of body mass increase for a ten-fold change in mammals.[6] The disparity is thought to reflect the fact that pedomorphism offers a relatively easy route to evolve smaller adult body size; on the other hand, the evolution of larger maximum body size is likely to be interrupted by the emergence of a series of constraints that must be overcome by evolutionary innovations before the process can continue.[6]

Factors influencing the extent of dwarfing

For both herbivores and carnivores, island size, the degree of island isolation and the size of the ancestral continental species appear not to be of major direct importance to the degree of dwarfing.[4] However, when considering only the body masses of recent top herbivores and carnivores, and including data from both continental and island land masses, the body masses of the largest species in a land mass were found to scale to the size of the land mass, with slopes of about 0.5 log(body mass/kg) per log(land area/km2).[7] There were separate regression lines for endothermic top predators, ectothermic top predators, endothermic top herbivores and (on the basis of limited data) ectothermic top herbivores, such that food intake was 7 to 24-fold higher for top herbivores than for top predators, and about the same for endotherms and ectotherms of the same trophic level (this leads to ectotherms being 5 to 16 times heavier than corresponding endotherms).[7]

Insular dwarfism in dinosaurs

Europasaurus skull

Recognition that insular dwarfism could apply to dinosaurs arose through the work of Ferenc Nopcsa, a Hungarian-born aristocrat, adventurer, scholar, and paleontologist. Nopcsa studied Transylvanian dinosaurs intensively, noticing that they were smaller than their cousins elsewhere in the world. For example, he unearthed six-meter-long sauropods, a group of dinosaurs which elsewhere commonly grew to 30 meters or more, which he named Magyarosaurus. Nopcsa deduced that the area where the remains were found was an island, Hațeg Island (now the Haţeg or Hatzeg basin in Romania) during the Mesozoic era. Nopcsa's proposal of dinosaur dwarfism on Hațeg Island is today widely accepted after further research confirmed that the remains found are not from juveniles.[8]

Famous examples

Homo floresiensis skull cast

Among the most famous examples of insular dwarfism are:

Additional examples

Channel Island fox
Lowland anoa
Kangaroo Island (left) and King Island emus
Brookesia micra

Carnivorans

Ungulates

Birds

Lizards

Snakes

See also

Wikinews has related news: Bones of "small-bodied humans" found in cave

Notes

  1. ↑ An example of noninsular phyletic dwarfism is the evolution of the dwarfed marmosets and tamarins among New World monkeys, culminating in the appearance of the smallest example, Cebuella pygmaea.[2]

References

  1. ↑ Prothero, D. R.; Sereno, P. C. (Winter 1982). "Allometry and Paleoecology of Medial Miocene Dwarf Rhinoceroses from the Texas Gulf Coastal Plain". Paleobiology. 8 (1): 16–30. JSTOR 2400564.
  2. ↑ Perelman, P.; et al. (2011). "A Molecular Phylogeny of Living Primates". PLOS Genetics. 7 (3): 1–17. doi:10.1371/journal.pgen.1001342. PMC 3060065Freely accessible. PMID 21436896.
  3. 1 2 3 4 Van Den Bergh, G. D.; Rokhus Due Awe; Morwood, M. J.; Sutikna, T.; Jatmiko; Wahyu Saptomo, E. (May 2008). "The youngest Stegodon remains in Southeast Asia from the Late Pleistocene archaeological site Liang Bua, Flores, Indonesia". Quaternary International. 182 (1): 16–48. doi:10.1016/j.quaint.2007.02.001. Retrieved 2011-11-27.
  4. 1 2 3 4 Raia, P.; Meiri, S. (August 2006). "The island rule in large mammals: paleontology meets ecology". Evolution. 60 (8): 1731–1742. doi:10.1111/j.0014-3820.2006.tb00516.x. Retrieved 2011-11-27.
  5. 1 2 Keogh, J. S.; Scott, I. A. W.; Hayes, C. (January 2005). "Rapid and repeated origin of insular gigantism and dwarfism in Australian tiger snakes". Evolution. 59 (1): 226–233. doi:10.1111/j.0014-3820.2005.tb00909.x.
  6. 1 2 Evans, A. R.; et al. (2012-01-30). "The maximum rate of mammal evolution". PNAS. 109. doi:10.1073/pnas.1120774109. Retrieved 2011-02-11.
  7. 1 2 Burness, G. P.; Diamond, J.; Flannery, T. (2001-12-04). "Dinosaurs, dragons, and dwarfs: The evolution of maximal body size". Proceedings of the National Academy of Sciences. 98 (25): 14518–14523. doi:10.1073/pnas.251548698. ISSN 0027-8424. JSTOR 3057309. PMC 64714Freely accessible. PMID 11724953. Retrieved 2012-01-28.
  8. ↑ Dyke, G. (2011-09-20). "The Dinosaur Baron of Transylvania". Scientific American. 305 (4): 80–83. doi:10.1038/scientificamerican1011-80. PMID 22106812.
  9. ↑ "Dwarf dinosaur island really did exist, scientists claim". Telegraph Media Group. 2010-02-22. Retrieved 2010-02-26.
  10. ↑ Benton, M. J.; Csiki, Z.; Grigorescu, D.; Redelstorff, R.; Sander, P. M.; Stein, K.; Weishampel, D. B. (2010-01-28). "Dinosaurs and the island rule: The dwarfed dinosaurs from Haţeg Island" (PDF). Palaeogeography, Palaeoclimatology, Palaeoecology. Elsevier. 293 (3–4): 438–454. doi:10.1016/j.palaeo.2010.01.026. Retrieved 2010-02-26.
  11. ↑ Össi, A.; Prondvai, E.; Butler, R.; Weishampel, D. B. (2012-09-21). "Phylogeny, Histology and Inferred Body Size Evolution in a New Rhabdodontid Dinosaur from the Late Cretaceous of Hungary". PLOS ONE. 7 (9): e44318. doi:10.1371/journal.pone.0044318. PMC 3448614Freely accessible. PMID 23028518.
  12. ↑ Extinct dwarf elephants from the Mediterranean islands
  13. ↑ Schirber, Michael. Surviving Extinction: Where Woolly Mammoths Endured. Live Science. Imaginova Corporation. Retrieved 2007-07-20.
  14. ↑ Tikhonov, Alexei; Larry Agenbroad; Sergey Vartanyan (2003). Comparative analysis of the mammoth populations on Wrangel Island and the Channel Islands. DEINSEA 9: 415–420. ISSN 0923-9308
  15. ↑ North American Extinctions v. World
  16. ↑ Scientist to study Hobbit morphing, abc.net.au
  17. ↑ "Ancient Small People on Palau Not Dwarfs, Study Says". National Geographic News. August 27, 2008.
  18. ↑ Lister, A. M. (1989-11-30). "Rapid dwarfing of red deer on Jersey in the Last Interglacial". Nature. 342 (6249): 539–542. doi:10.1038/342539a0. PMID 2685610. Retrieved 2011-02-28.
  19. ↑ Heupink, T. H.; Huynen, L.; Lambert, D. M. (2011). "Ancient DNA Suggests Dwarf and 'Giant' Emu Are Conspecific". PLoS ONE. 6 (4): e18728. doi:10.1371/journal.pone.0018728. PMC 3073985Freely accessible. PMID 21494561.
  20. ↑ Parker S (1984) The extinct Kangaroo Island Emu, a hitherto-unrecognised species. Bulletin of the British Ornithologists' Club 104: 19–22.
  21. ↑ Glaw, F.; Köhler, J.; Townsend, T. M.; Vences, M. (2012-02-14). "Rivaling the World's Smallest Reptiles: Discovery of Miniaturized and Microendemic New Species of Leaf Chameleons (Brookesia) from Northern Madagascar". PLoS ONE. 7 (2): e31314. doi:10.1371/journal.pone.0031314. PMC 3279364Freely accessible. PMID 22348069. Retrieved 2012-02-17.
  22. ↑ Auliya, M.; Mausfeld, P.; Schmitz, A.; Böhme, W. (2002-04-09). "Review of the reticulated python (Python reticulatus Schneider, 1801) with the description of new subspecies from Indonesia". Naturwissenschaften. 89 (5): 201–213. doi:10.1007/s00114-002-0320-4. Retrieved 2012-04-08.

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