Insular dwarfism

Form of phyletic dwarfism occurring on islands From Wikipedia, the free encyclopedia

Insular dwarfism, a form of phyletic dwarfism,[1] is the process and condition of large animals evolving or having a reduced body size[a] 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 various species of dwarf elephants that evolved during the Pleistocene epoch, as well as more ancient examples, such as the dinosaurs Europasaurus and Magyarosaurus. This process, and other "island genetics" artifacts, can occur not only on 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").[citation needed] Insular dwarfism 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 (island gigantism), and large species tend to evolve smaller bodies. This is itself one aspect of island syndrome, which describes the differences in morphology, ecology, physiology and behaviour of insular species compared to their continental counterparts.

Skeletons of the extinct Palaeoloxodon falconeri, native to Sicily and Malta, it is one of the smallest known species of dwarf elephant. Adult males measured about one meter (3.3 ft) in shoulder height and weighed about 250 kg (550 lb). Females were smaller.

Possible causes

Structure of insular dwarfism web

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]

Differences of dwarfism and 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 stegodonts 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]

It has been suggested that for dwarf elephants, competition was an important factor in body size, with islands with competing herbivores having significantly larger dwarf elephants than those where competing herbivores were absent.[8]

Examples

Non-avian dinosaurs

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. 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.[9][10] 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.[11]

Sauropods

More information Example, Species ...
Example Species Range Time frame Continental relative

Ampelosaurus
A. atacisIbero-Armorican IslandLate Cretaceous / Maastrichtian
Nemegtosaurids

Europasaurus
E. holgeriLower SaxonyLate Jurassic / Middle Kimmeridgian
Brachiosaurs

Magyarosaurus
M. dacusHațeg IslandLate Cretaceous / Maastrichtian
Rapetosaurus

Lirainosaurus[12]
L. astibiaeIbero-Armorican IslandLate Cretaceous

Paludititan
P. nalatzensisHațeg IslandLate Cretaceous / Maastrichtian
Epachthosaurus
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Other

More information Example, Species ...
Example Species Range Time frame Continental relative

Langenberg Quarry
torvosaur (blue)
UnnamedLower SaxonyLate Jurassic / Middle Kimmeridgian
Torvosaurus

Struthiosaurus[13]
S. austriacusIbero-Armorican, Australoalpine, and Hațeg IslandsLate Cretaceous
Edmontonia
S. transylvanicus
S. languedocensis

Telmatosaurus
T. transsylvanicusHațeg Island
Hadrosaurids

Thecodontosaurus[10]
T. antiquusSouthern EnglandLate Triassic / Rhaetian
Plateosaurs

Zalmoxes[10] (purple)
Z. robustusHațeg IslandLate Cretaceous
Tenontosaurus
Z. shqiperorum
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In addition, the genus Balaur was initially described as a Velociraptor-sized dromaeosaurid (and in consequence a dubious example of insular dwarfism), but has been since reclassified as a secondarily flightless stem bird, closer to modern birds than Jeholornis (thus actually an example of insular gigantism).

Birds

More information Example, Binomial name ...
Example Binomial name Native range Status Continental relative Insular / mainland
length or mass ratio

Hawaiian flightless ibises
Apteribis glenosMolokaiExtinct (Late Quaternary)
American ibises
Apteribis brevisMaui
Cozumel curassow[14]Crax rubra griscomiCozumelUnknown
Great curassow

Kangaroo Island emu[15]
Dromaius novaehollandiae baudinianusKangaroo Island, South AustraliaExtinct (c. AD 1827)
Emu

King Island emu[16] (black)
Dromaius novaehollandiae minorKing Island, TasmaniaExtinct (AD 1822)LR ≈ 0.48[b]
Dwarf yellow eyed penguin[17]Megadyptes antipodes richdaleiChatham Islands, New ZealandExtinct (after 1300 AD)
Yellow-eyed penguin

Cozumel thrasher[14]
Toxostoma gluttatumCozumelCritically endangered
Other thrashers
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Squamates

More information Example, Binomial name ...
Example Binomial name Native range Status Continental relative Insular / mainland
length or mass ratio

Madagascar dwarf chameleon
Brookesia minimaNosy Be island, MadagascarEndangered
Madagascar leaf chameleons

Nosy Hara chameleon[18]
Brookesia micraNosy Hara island, MadagascarVulnerable
Roxby Island tiger snake[5]Notechis scutatusRoxby Island, South AustraliaUnknown
Tiger snake
Dwarf Burmese python Python bivittatus progschaiJava, Bali, Sumbawa and Sulawesi, Indonesia
Burmese python
LR ≈ 0.44[c]
Tanahjampea reticulated python[21] Python reticulatus jampeanusTanahjampea, between Sulawesi and Flores
Reticulated python
LR ≈ 0.41, males
LR ≈ 0.49, females[d]
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Mammals

Pilosans

More information Example, Binomial name ...
Example Binomial name Native range Status Continental relative

Pygmy three-toed sloth
Bradypus pygmaeusIsla Escudo de Veraguas, PanamaCritically endangered
Brown-throated sloth

Acratocnus
A. antillensisCuba, Hispaniola and Puerto RicoExtinct (c. 3000 BC)
Continental ground sloths
A. odontrigonus
A. ye
ImagocnusI. zazaeCubaExtinct (Early Miocene)

Megalocnus
M. rodensCuba and HispaniolaExtinct (c. 2700 BC)
M. zile

Neocnus
Neocnus spp.Extinct (c. 3000 BC)
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Proboscideans

More information Example, Binomial name ...
Example Binomial name Native range Status Continental relative
Sulawesi dwarf elephantElephas celebensisSulawesiExtinct (Early Pleistocene)
Asian elephant
Cabarruyan dwarf elephantElephas beyeriLuzonExtinct
Cretan dwarf mammothMammuthus creticusCrete
Mammuthus

Channel Islands mammoth
Mammuthus exilisSanta Rosae islandExtinct (Late Pleistocene)
Columbian mammoth

Sardinian mammoth

Mammuthus lamarmoraiSardinia
Steppe mammoth
Taiwan mammoth[24]Mammuthus trogontherii taiwanicusTaiwanExtinct
Saint Paul Island woolly mammoth[25][26]Mammuthus primigeniusSaint Paul Island, AlaskaExtinct (c. 3750 BC)
Woolly mammoth

Siculo-Maltese elephants
Palaeoloxodon antiquus leonardiSicily and MaltaExtinct
Straight-tusked elephant
(left)
P. mnaidriensis
P. melitensis
P. falconeri
Cretan elephantsPalaeoloxodon chaniensis
Crete
P. creutzburgi

Cyprus dwarf elephant
Palaeoloxodon cypriotesCyprusExtinct (c. 9000 BC)
Naxos dwarf elephantPalaeoloxodon sp.NaxosExtinct
Tilos dwarf elephantPalaeoloxodon tiliensisTilos
Rhodes dwarf elephant Palaeoloxodon sp. Rhodes
Bumiayu dwarf sinomastodont[27]Sinomastodon bumiajuensisBumiayu Island (now part of Java)Extinct (Early Pleistocene)
Sinomastodon

Japanese stegodont[28][29]
Stegodon miensisJapan (Also Taiwan for S. aurorae)[30]
Chinese Stegodon
Stegodon protoaurorae
Stegodon aurorae
Greater Flores dwarf stegodont[3]Stegodon florensisFloresExtinct (Late Pleistocene)
Sundaland Stegodon
Javan dwarf stegodontsStegodon hypsilophus[27]JavaExtinct (Quaternary)
S. semedoensis[31]
S. sp.[27]
Mindanao pygmy stegodont[32]Stegodon mindanensisMindanao and SulawesiExtinct (Middle Pleistocene)
Sulawesi dwarf stegodont[27]Stegodon sompoensisSulawesiExtinct
Lesser Flores dwarf stegodont[3]Stegodon sondaariFloresExtinct (Middle Pleistocene)
Sumba dwarf stegodont[33]Stegodon sumbaensisSumba, Indonesia
Timor dwarf stegodont[27]Stegodon timorensisTimorExtinct
Dwarf stegolophodont[34]Stegolophodon pseudolatidensJapanExtinct (Miocene)
Stegolophodon
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Primates

More information Example, Binomial name ...
Example Binomial name Native range Status Continental relative
Nosy Hara dwarf lemur[35]Cheirogaleus sp.Nosy Hara island off MadagascarUnknown
Dwarf lemurs

Flores Man[36]
Homo floresiensisFloresExtinct (Late Pleistocene)
Homo erectus

Callao Man
Homo luzonensis[37][38]Luzon, Philippines
Modern pygmies of Flores[39]Homo sapiensFloresExtantother members of Homo sapiens
Early Palau modern humans (disputed)[40]PalauExtinct (?)
Andamanese[41]Andaman IslandsExtant

Sardinian macaque[42]
Macaca majoriSardiniaExtinct (Pleistocene)
Barbary macaque

Zanzibar red colobus
Piliocolobus kirkiiUngujaEndangered
Udzungwa red colobus
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Carnivorans

More information Example, Binomial name ...
Example Binomial name Native range Status Continental relative Insular / mainland
length or mass ratio

Sicilian wolf
Canis lupus cristaldiiSicilyExtinct (AD 1970)
Gray wolf

Japanese wolf
Canis lupus hodophilaxJapan (excluding Hokkaido)Extinct (AD 1905)

Sardinian dhole
(forward)
Cynotherium sardousCorsica and SardiniaExtinct (c. 8300 BC)
Xenocyon
Trinil dogMececyon trinilensisJavaExtinct (Pleistocene)
Cozumel Island coati[14]Nasua narica nelsoniCozumelCritically endangered
Yucatan white-nosed coati

Zanzibar leopard
Panthera pardus pardusUngujaCritically endangered or Extinct
African leopard
Taiwan leopard [43]Panthera pardusTaiwanExtinct (Pleistocene)
Amur leopard

Bali tiger
Panthera tigris sondaicaBaliExtinct (c. AD 1940)
Sumatran tiger

Javan tiger
JavaExtinct (c. AD 1975)

Cozumel raccoon
Procyon pygmaeusCozumelCritically endangered
Common raccoon

Island fox
Urocyon littoralisSix of the Channel Islands of CaliforniaNear Threatened
Gray fox
LR ≈ 0.84[e]
LR ≈ 0.75[f]
Cozumel foxUrocyon sp.CozumelCritically endangered or Extinct
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Non-ruminant ungulates

More information Example, Binomial name ...
Example Binomial name Native range Status Continental relative

Eumaiochoerus
Eumaiochoerus etruscusBaccinello, MontebamboliExtinct (Miocene)
Microstonyx

Malagasy dwarf hippopotamuses
Hippopotamus laloumenaMadagascarExtinct (c. AD 1000)
Common hippopotamus
H. lemerlei
H. madagascariensis
Bumiayu dwarf hippopotamus[27]Hexaprotodon simplexBumiayu Island (now Java)Extinct (Early Pleistocene)
Asian hippopotamuses

Cretan dwarf hippopotamus
Hippopotamus creutzburgiCreteExtinct (Middle Pleistocene)
Hippopotamus antiquus

Maltese dwarf hippopotamus
Hippopotamus melitensisMaltaExtinct (Pleistocene)
Common hippopotamus (H. amphibius)

Sicilian dwarf hippopotamus
Hippopotamus pentlandiSicily

Cyprus dwarf hippopotamus
Hippopotamus minorCyprusExtinct (c. 8000 BC) Unclear, either H. amphibius or H. antiquus.
Cozumel collared peccary[14]Pecari tajacu nanusCozumelUnknown
Collared peccary
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Bovids

More information Example, Binomial name ...
Example Binomial name Native range Status Continental relative
Sicilian bison[28]Bison priscus siciliaeSicilyExtinct (Late Pleistocene)
Steppe bison
Sicilian aurochs[46]Bos primigenius siciliae[28]
Eurasian aurochs
Cebu tamarawBubalus cebuensisCebu, PhilippinesExtinct
Wild water buffalo

Lowland anoa
Bubalus depressicornisSulawesi and Buton, IndonesiaEndangered
Bubalus grovesiBubalus grovesiSulawesi, IndonesiaExtinct

Tamaraw
Bubalus mindorensisMindoro, PhilippinesCritically endangered

Mountain anoa
Bubalus quarlesiSulawesi and Buton, IndonesiaEndangered

Balearic Islands cave goat
Myotragus balearicusMajorca and MenorcaExtinct (after 3000 BC)Gallogoral
Nesogoral[47]Nesogoral spp.SardiniaExtinct
Dahlak Kebir gazelle[48]Nanger soemmerringi ssp.Dahlak Kebir island, EritreaVulnerable
Soemmerring's gazelle

Tyrrhenotragus
Tyrrhenotragus gracillimusBaccinelloExtinctAntilopinae sp.
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Cervids and relatives

More information Example, Binomial name ...
Example Binomial name Native range Status Continental relative

Cretan deer[g]
Candiacervus spp.CreteExtinct (Pleistocene)Unknown

Sardinian deer[10]
Praemegaceros caziotiSardiniaExtinct (c. 5500 BC) Praemegaceros

Ryukyu dwarf deer[51]
Cervus astylodonRyukyu IslandsExtinct
Sika deer (?)
Cervus praenipponicus (?)
Jersey red deer population[52]Cervus elaphus jerseyensisJerseyExtinct (Pleistocene)
Red deer

Corsican red deer
Cervus elaphus corsicanusCorsica and SardiniaNear Threatened
Sicilian red deer[28]Cervus siciliaeSicilyExtinct (Late Pleistocene)

Hoplitomeryx[h]
Hoplitomeryx spp.Gargano IslandExtinct (Early Pliocene)
Pecorans
Sicilian fallow deerDama carburangelensisSicilyExtinct (Late Pleistocene)Fallow deer

Florida Key deer
Odocoileus virginianus claviumFlorida KeysEndangered
Virginia deer

Svalbard reindeer
Rangifer tarandus platyrhynchusSvalbardVulnerable
Reindeer

Philippine deer
Rusa mariannaPhilippines
Sambar deer
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Plants

More information Possible example, Binomial name ...
Possible example Binomial name Native range Status Continental relative

Insular elephant cacti[53][54]
Pachycereus pringleiRemote islands in the Sea of Cortez (e.g. Santa Cruz, San Pedro Mártir)Not evaluated
Mainland elephant cacti
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See also

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]
  2. Based on the heights in Fig. 1 of Heupink et al., 2011[16]
  3. Based on maximum lengths of 2.5 m for the dwarf form[19] and 5.74 m for the mainland form[20]
  4. Based on maximum Tanahjampea python total lengths (TL) of 2.10 m for males and 3.35 m for females[21] and maximum southern Sumatra python snout to vent lengths (SVL) of 4.5 m for males and 6.1 m for females[22] with SVLs corrected to TLs by multiplying by a factor of 1.127, derived from the average relative tail length (0.113) of African and Indian rock pythons[23]
  5. For nearby mainland gray foxes[44]
  6. For mainland gray foxes in general[45]
  7. Like Hoplitomeryx, Candiacervus appears to be an unusual case in that members of this genus evolved into insular species of a wide range of sizes, not only dwarf forms but also some that might be considered giants.[49][50]
  8. Hoplitomeryx is evidently quite an unusual case, because members of this genus apparently evolved into both dwarf and giant insular forms on the same island(s).[49]

References

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