Insular dwarfism

The skeleton of a dwarf elephant from the island of Crete.

Insular dwarfism, a form of phyletic dwarfism, is the process and condition of the reduction in size of large animals over a number of generations 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.

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.

In the tropics, small size should make thermoregulation easier.

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.

Among carnivores, the main factor is thought to be the size and availability of prey resources, and competition is believed to be less important. 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.

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. 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.

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. 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). 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).

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.

Famous examples

Homo floresiensis skull cast

Among the most famous examples of insular dwarfism are:

• Dinosaurs from Mesozoic islands such as Haţeg Island, now in Romania:
  • Europasaurus, a brachiosaur
  • Magyarosaurus, a titanosaur
  • Zalmoxes, an iguanodontid related to Rhabdodon
  • Balaur, an eumaniraptoran
  • Telmatosaurus, a hadrosaur
  • Struthiosaurus, a nodosaurid (also present outside Romania)
• Dwarf pilosans of the Caribbean:
  • Ground sloths in the recent natural history of Cuba, Hispaniola, and Puerto Rico, such as Megalocnus.
  • Pygmy three-toed sloth of Isla Escudo de Veraguas, Bradypus pygmaeus.
• Reduced island proboscideans of the late Pleistocene and early Holocene:
  • Channel Islands mammoth, Mammuthus exilis, which lived on the prehistoric island of Santa Rosae off the coast of California
  • Dwarfed population of woolly mammoth, Mammuthus primigenius, in Saint Paul Island off Alaska (the mammoths of Wrangel Island, north of Siberia, are no longer considered dwarfs)
  • Sardinian mammoth, Mammuthus lamarmorae
  • Cretan mammoth, Mammuthus creticus
  • Siculo-Maltese elephants: Palaeoloxodon antiquus leonardi, P. mnaidriensis, P. melitensis, P. falconeri
  • Cretan elephants: Palaeoloxodon chaniensis, P. creutzburgi
  • Cyprus dwarf elephant, Palaeoloxodon cypriotes
  • Naxos dwarf elephant, Palaeoloxodon sp.
  • Rhodes and Tilos dwarf elephant, Palaeoloxodon tiliensis
  • Dwarf stegodonts from the recent natural history of the Philippines, Flores, Sulawesi, Sumba and Timor, such as Stegodon sondaarii, S. florensis and S.sompoensis
• Dwarf hominins:
  • A recently confirmed separate species of hominid called Homo floresiensis, from fossils found on Flores Island in Indonesia.
  • Small-bodied humans from Palau, Micronesia, similar in size to the Flores hominins (disputed).

Additional examples

Channel Island fox

Lowland anoa

Kangaroo Island (left) and King Island emus

Brookesia micra

Carnivorans

• Canids
  • Channel Island fox in California, Urocyon littoralis
  • Cozumel fox, Urocyon sp.
  • Honshū wolf in Japan, Canis lupus hodophilax †
  • Sardinian dhole, Cynotherium sardous †
• Felids
  • Bali tiger, Panthera tigris balica †
  • Javan tiger, Panthera tigris sondaica †
  • Zanzibar leopard, Panthera pardus adersi (probably extinct)
• Procyonids
  • Cozumel raccoon, Procyon pygmaeus

Ungulates

• Bovids
  • Cebu tamaraw, Bubalus cebuensis †
  • Lowland anoa, Bubalus depressicornis
  • Tamaraw, Bubalus mindorensis
  • Mountain anoa, Bubalus quarlesi
  • Balearic Islands cave goat, Myotragus balearicus †
  • Nesogoral † from Sardinia
• Cervids
  • Key deer of the Florida Keys, Odocoileus virginianus clavium
  • Extinct red deer populations in Jersey, Cervus elaphus
  • Corsican red deer, Cervus elaphus corsicanus
  • Svalbard reindeer, Rangifer tarandus platyrhynchus
  • Philippine sambar, Rusa mariana
  • Several species of Candiacervus † from Crete
  • Cervus astylodon †, Ryukyu Islands of Japan
  • Hoplitomeryx †, former Gargano Island in South Italy
• Hippopotamids
  • Cretan dwarf hippopotamus, Hippopotamus creutzburgi †
  • Maltese hippopotamus, Hippopotamus melitensis †
  • Cyprus dwarf hippopotamus, Hippopotamus minor †
  • Sicilian hippopotamus, Hippopotamus pentlandi †
  • Several species of Malagasy hippopotamus, Hippopotamus sp. †

Birds

• King Island emu, Dromaius novaehollandiae ater †
• Kangaroo Island emu, Dromaius baudinianus †

Lizards

• Madagascar dwarf chamaeleon, Brookesia minima
• Brookesia micra, a less than 3.0 cm long chameleon from the islet of Nosy Hara off the northern tip of Madagascar, one of the "smallest amniote vertebrates in the world"

Snakes

• Tiger snake of Roxby Island (South Australia), Notechis scutatus
• Reticulated python of Tanahjampea island, Python reticulatus jampeanus, between Sulawesi and Flores in Indonesia

See also

• Dwarfing
• Foster's rule
• Franz Nopcsa
• Island gigantism
• Island tameness
• Pleistocene extinctions

External links

• Strange world of island species October 31, 2004 The Observer