# Insular dwarfism

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Form of phyletic dwarfism occurring on islands

For other uses, see [Dwarf (disambiguation)](/source/Dwarf_(disambiguation)).

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

**Insular dwarfism**, a form of [phyletic dwarfism](/source/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](/source/Dwarfing). This process has occurred many times throughout evolutionary history, with examples including various species of [dwarf elephants](/source/Dwarf_elephant) that evolved during the [Pleistocene](/source/Pleistocene) epoch, as well as more ancient examples, such as the dinosaurs *[Europasaurus](/source/Europasaurus)* and *[Magyarosaurus](/source/Magyarosaurus)*. This process, and other "[island genetics](/source/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](/source/Caves), desert [oases](/source/Oases), isolated valleys and isolated mountains ("[sky islands](/source/Sky_island)").[*[citation needed](https://en.wikipedia.org/wiki/Wikipedia:Citation_needed)*] Insular dwarfism is one aspect of the more general ["island effect" or "Foster's rule"](/source/Foster's_rule), which posits that when mainland animals colonize islands, small species tend to evolve larger bodies ([island gigantism](/source/Island_gigantism)), and large species tend to evolve smaller bodies. This is itself one aspect of [island syndrome](/source/Island_syndrome), which describes the differences in [morphology](/source/Morphology_(biology)), [ecology](/source/Ecology), [physiology](/source/Physiology) and [behaviour](/source/Ethology) of insular species compared to their continental counterparts.

## 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](/source/Gestation_period) and [generation times](/source/Generation_time).[3]

In the tropics, small size should make [thermoregulation](/source/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](/source/Tiger_snake), 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](/source/Island_gigantism). An excellent example is the [dodo](/source/Dodo), the ancestors of which were normal-sized [pigeons](/source/Pigeon). There are also several species of [giant rats](/source/Papagomys), one still extant, that coexisted with both *[Homo floresiensis](/source/Homo_floresiensis)* and the dwarf [stegodonts](/source/Stegodont) 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](/source/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](/source/Regression_line) for [endothermic](/source/Endotherm) top predators, [ectothermic](/source/Ectotherm) 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](/source/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](/source/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](/source/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](/source/Ha%C8%9Beg_Island) (now the Haţeg or Hatzeg basin in [Romania](/source/Romania)) during the [Mesozoic](/source/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

Example Species Range Time frame Continental relative Ampelosaurus A. atacis Ibero-Armorican Island Late Cretaceous / Maastrichtian Nemegtosaurids Europasaurus E. holgeri Lower Saxony Late Jurassic / Middle Kimmeridgian Brachiosaurs Magyarosaurus M. dacus Hațeg Island Late Cretaceous / Maastrichtian Rapetosaurus Lirainosaurus[12] L. astibiae Ibero-Armorican Island Late Cretaceous Paludititan P. nalatzensis Hațeg Island Late Cretaceous / Maastrichtian Epachthosaurus

#### Other

Example Species Range Time frame Continental relative Langenberg Quarry torvosaur (blue) Unnamed Lower Saxony Late Jurassic / Middle Kimmeridgian Torvosaurus Struthiosaurus[13] S. austriacus Ibero-Armorican, Australoalpine, and Hațeg Islands Late Cretaceous Edmontonia S. transylvanicus S. languedocensis Telmatosaurus T. transsylvanicus Hațeg Island Hadrosaurids Thecodontosaurus[10] T. antiquus Southern England Late Triassic / Rhaetian Plateosaurs Zalmoxes[10] (purple) Z. robustus Hațeg Island Late Cretaceous Tenontosaurus Z. shqiperorum

In addition, the genus *[Balaur](/source/Balaur_(dinosaur))* was initially described as a *[Velociraptor](/source/Velociraptor)*-sized [dromaeosaurid](/source/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](/source/Jeholornis)* (thus actually an example of [insular gigantism](/source/Insular_gigantism)).

### Birds

Example Binomial name Native range Status Continental relative Insular / mainland length or mass ratio Hawaiian flightless ibises Apteribis glenos Molokai Extinct (Late Quaternary) American ibises Apteribis brevis Maui Cozumel curassow[14] Crax rubra griscomi Cozumel Unknown Great curassow Kangaroo Island emu[15] Dromaius novaehollandiae baudinianus Kangaroo Island, South Australia Extinct (c. AD 1827) Emu King Island emu[16] (black) Dromaius novaehollandiae minor King Island, Tasmania Extinct (AD 1822) LR ≈ 0.48[b] Dwarf yellow eyed penguin[17] Megadyptes antipodes richdalei Chatham Islands, New Zealand Extinct (after 1300 AD) Yellow-eyed penguin Cozumel thrasher[14] Toxostoma gluttatum Cozumel Critically endangered Other thrashers

### [Squamates](/source/Squamates)

Example Binomial name Native range Status Continental relative Insular / mainland length or mass ratio Madagascar dwarf chameleon Brookesia minima Nosy Be island, Madagascar Endangered Madagascar leaf chameleons Nosy Hara chameleon[18] Brookesia micra Nosy Hara island, Madagascar Vulnerable Roxby Island tiger snake[5] Notechis scutatus Roxby Island, South Australia Unknown Tiger snake Dwarf Burmese python Python bivittatus progschai Java, Bali, Sumbawa and Sulawesi, Indonesia Burmese python LR ≈ 0.44[c] Tanahjampea reticulated python[21] Python reticulatus jampeanus Tanahjampea, between Sulawesi and Flores Reticulated python LR ≈ 0.41, males LR ≈ 0.49, females[d]

### Mammals

#### [Pilosans](/source/Pilosans_of_the_Caribbean)

Example Binomial name Native range Status Continental relative Pygmy three-toed sloth Bradypus pygmaeus Isla Escudo de Veraguas, Panama Critically endangered Brown-throated sloth Acratocnus A. antillensis Cuba, Hispaniola and Puerto Rico Extinct (c. 3000 BC) Continental ground sloths A. odontrigonus A. ye Imagocnus I. zazae Cuba Extinct (Early Miocene) Megalocnus M. rodens Cuba and Hispaniola Extinct (c. 2700 BC) M. zile Neocnus Neocnus spp. Extinct (c. 3000 BC)

#### [Proboscideans](/source/Proboscidean)

Main article: [Dwarf elephant](/source/Dwarf_elephant)

Example Binomial name Native range Status Continental relative Sulawesi dwarf elephant Elephas celebensis Sulawesi Extinct (Early Pleistocene) Asian elephant Cabarruyan dwarf elephant Elephas beyeri Luzon Extinct Cretan dwarf mammoth Mammuthus creticus Crete Mammuthus Channel Islands mammoth Mammuthus exilis Santa Rosae island Extinct (Late Pleistocene) Columbian mammoth Sardinian mammoth Mammuthus lamarmorai Sardinia Steppe mammoth Taiwan mammoth[24] Mammuthus trogontherii taiwanicus Taiwan Extinct Saint Paul Island woolly mammoth[25][26] Mammuthus primigenius Saint Paul Island, Alaska Extinct (c. 3750 BC) Woolly mammoth Siculo-Maltese elephants Palaeoloxodon antiquus leonardi Sicily and Malta Extinct Straight-tusked elephant (left) P. mnaidriensis P. melitensis P. falconeri Cretan elephants Palaeoloxodon chaniensis Crete P. creutzburgi Cyprus dwarf elephant Palaeoloxodon cypriotes Cyprus Extinct (c. 9000 BC) Naxos dwarf elephant Palaeoloxodon sp. Naxos Extinct Tilos dwarf elephant Palaeoloxodon tiliensis Tilos Rhodes dwarf elephant Palaeoloxodon sp. Rhodes Bumiayu dwarf sinomastodont[27] Sinomastodon bumiajuensis Bumiayu Island (now part of Java) Extinct (Early Pleistocene) Sinomastodon Japanese stegodont[28][29] Stegodon miensis Japan (Also Taiwan for S. aurorae)[30] Chinese Stegodon Stegodon protoaurorae Stegodon aurorae Greater Flores dwarf stegodont[3] Stegodon florensis Flores Extinct (Late Pleistocene) Sundaland Stegodon Javan dwarf stegodonts Stegodon hypsilophus[27] Java Extinct (Quaternary) S. semedoensis[31] S. sp.[27] Mindanao pygmy stegodont[32] Stegodon mindanensis Mindanao and Sulawesi Extinct (Middle Pleistocene) Sulawesi dwarf stegodont[27] Stegodon sompoensis Sulawesi Extinct Lesser Flores dwarf stegodont[3] Stegodon sondaari Flores Extinct (Middle Pleistocene) Sumba dwarf stegodont[33] Stegodon sumbaensis Sumba, Indonesia Timor dwarf stegodont[27] Stegodon timorensis Timor Extinct Dwarf stegolophodont[34] Stegolophodon pseudolatidens Japan Extinct (Miocene) Stegolophodon

#### [Primates](/source/Primates)

Example Binomial name Native range Status Continental relative Nosy Hara dwarf lemur[35] Cheirogaleus sp. Nosy Hara island off Madagascar Unknown Dwarf lemurs Flores Man[36] Homo floresiensis Flores Extinct (Late Pleistocene) Homo erectus Callao Man Homo luzonensis[37][38] Luzon, Philippines Modern pygmies of Flores[39] Homo sapiens Flores Extant other members of Homo sapiens Early Palau modern humans (disputed)[40] Palau Extinct (?) Andamanese[41] Andaman Islands Extant Sardinian macaque[42] Macaca majori Sardinia Extinct (Pleistocene) Barbary macaque Zanzibar red colobus Piliocolobus kirkii Unguja Endangered Udzungwa red colobus

#### [Carnivorans](/source/Carnivorans)

Example Binomial name Native range Status Continental relative Insular / mainland length or mass ratio Sicilian wolf Canis lupus cristaldii Sicily Extinct (AD 1970) Gray wolf Japanese wolf Canis lupus hodophilax Japan (excluding Hokkaido) Extinct (AD 1905) Sardinian dhole (forward) Cynotherium sardous Corsica and Sardinia Extinct (c. 8300 BC) Xenocyon Trinil dog Mececyon trinilensis Java Extinct (Pleistocene) Cozumel Island coati[14] Nasua narica nelsoni Cozumel Critically endangered Yucatan white-nosed coati Zanzibar leopard Panthera pardus pardus Unguja Critically endangered or Extinct African leopard Taiwan leopard [43] Panthera pardus Taiwan Extinct (Pleistocene) Amur leopard Bali tiger Panthera tigris sondaica Bali Extinct (c. AD 1940) Sumatran tiger Javan tiger Java Extinct (c. AD 1975) Cozumel raccoon Procyon pygmaeus Cozumel Critically endangered Common raccoon Island fox Urocyon littoralis Six of the Channel Islands of California Near Threatened Gray fox LR ≈ 0.84[e] LR ≈ 0.75[f] Cozumel fox Urocyon sp. Cozumel Critically endangered or Extinct

#### Non-ruminant [ungulates](/source/Ungulate)

Example Binomial name Native range Status Continental relative Eumaiochoerus Eumaiochoerus etruscus Baccinello, Montebamboli Extinct (Miocene) Microstonyx Malagasy dwarf hippopotamuses Hippopotamus laloumena Madagascar Extinct (c. AD 1000) Common hippopotamus H. lemerlei H. madagascariensis Bumiayu dwarf hippopotamus[27] Hexaprotodon simplex Bumiayu Island (now Java) Extinct (Early Pleistocene) Asian hippopotamuses Cretan dwarf hippopotamus Hippopotamus creutzburgi Crete Extinct (Middle Pleistocene) Hippopotamus antiquus Maltese dwarf hippopotamus Hippopotamus melitensis Malta Extinct (Pleistocene) Common hippopotamus (H. amphibius) Sicilian dwarf hippopotamus Hippopotamus pentlandi Sicily Cyprus dwarf hippopotamus Hippopotamus minor Cyprus Extinct (c. 8000 BC) Unclear, either H. amphibius or H. antiquus. Cozumel collared peccary[14] Pecari tajacu nanus Cozumel Unknown Collared peccary

#### [Bovids](/source/Bovid)

Example Binomial name Native range Status Continental relative Sicilian bison[28] Bison priscus siciliae Sicily Extinct (Late Pleistocene) Steppe bison Sicilian aurochs[46] Bos primigenius siciliae[28] Eurasian aurochs Cebu tamaraw Bubalus cebuensis Cebu, Philippines Extinct Wild water buffalo Lowland anoa Bubalus depressicornis Sulawesi and Buton, Indonesia Endangered Bubalus grovesi Bubalus grovesi Sulawesi, Indonesia Extinct Tamaraw Bubalus mindorensis Mindoro, Philippines Critically endangered Mountain anoa Bubalus quarlesi Sulawesi and Buton, Indonesia Endangered Balearic Islands cave goat Myotragus balearicus Mallorca and Menorca Extinct (after 3000 BC) Gallogoral Nesogoral[47] Nesogoral spp. Sardinia Extinct Dahlak Kebir gazelle[48] Nanger soemmerringi ssp. Dahlak Kebir island, Eritrea Vulnerable Soemmerring's gazelle Tyrrhenotragus Tyrrhenotragus gracillimus Baccinello Extinct Antilopinae sp.

#### [Cervids](/source/Cervid) and relatives

Example Binomial name Native range Status Continental relative Cretan deer[g] Candiacervus spp. Crete Extinct (Pleistocene) Unknown Sardinian deer[10] Praemegaceros cazioti Sardinia Extinct (c. 5500 BC) Praemegaceros Ryukyu dwarf deer[51] Cervus astylodon Ryukyu Islands Extinct Sika deer (?) Cervus praenipponicus (?) Jersey red deer population[52] Cervus elaphus jerseyensis Jersey Extinct (Pleistocene) Red deer Corsican red deer Cervus elaphus corsicanus Corsica and Sardinia Near Threatened Sicilian red deer[28] Cervus siciliae Sicily Extinct (Late Pleistocene) Hoplitomeryx[h] Hoplitomeryx spp. Gargano Island Extinct (Early Pliocene) Pecorans Sicilian fallow deer Dama carburangelensis Sicily Extinct (Late Pleistocene) Fallow deer Florida Key deer Odocoileus virginianus clavium Florida Keys Endangered Virginia deer Svalbard reindeer Rangifer tarandus platyrhynchus Svalbard Vulnerable Reindeer Philippine deer Rusa marianna Philippines Sambar deer

### [Plants](/source/Plant)

Possible example Binomial name Native range Status Continental relative Insular elephant cacti[53][54] Pachycereus pringlei Remote islands in the Sea of Cortez (e.g. Santa Cruz, San Pedro Mártir) Not evaluated Mainland elephant cacti

## See also

Wikinews has related news:

- [Bones of "small-bodied humans" found in cave](https://en.wikinews.org/wiki/Bones_of_%22small-bodied_humans%22_found_in_cave)

- [Island gigantism](/source/Island_gigantism)

- [Island syndrome](/source/Island_syndrome)

- [Island tameness](/source/Island_tameness)

- [Pleistocene extinctions](/source/Pleistocene_extinctions)

## Notes

1. **[^](#cite_ref-3)** An example of noninsular phyletic dwarfism is the evolution of the dwarfed [marmosets and tamarins](/source/Callitrichidae) among New World monkeys, culminating in the appearance of the smallest example, *[Cebuella pygmaea](/source/Cebuella_pygmaea)*.[2]

1. **[^](#cite_ref-18)** Based on the heights in Fig. 1 of Heupink *et al.*, 2011[16]

1. **[^](#cite_ref-23)** Based on maximum lengths of 2.5 m for the dwarf form[19] and 5.74 m for the mainland form[20]

1. **[^](#cite_ref-27)** 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]

1. **[^](#cite_ref-49)** For nearby mainland gray foxes[44]

1. **[^](#cite_ref-51)** For mainland gray foxes in general[45]

1. **[^](#cite_ref-57)** 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]

1. **[^](#cite_ref-60)** *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]

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1. **[^](#cite_ref-Telegraph_10-0)** ["Dwarf dinosaur island really did exist, scientists claim"](https://web.archive.org/web/20100225003941/http://www.telegraph.co.uk/science/dinosaurs/7291186/Dwarf-dinosaur-island-really-did-exist-scientists-claim.html). [Telegraph Media Group](/source/Telegraph_Media_Group). 2010-02-22. Archived from [the original](https://www.telegraph.co.uk/science/dinosaurs/7291186/Dwarf-dinosaur-island-really-did-exist-scientists-claim.html) on 2010-02-25. Retrieved 2010-02-26.

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1. **[^](#cite_ref-16)** Parker S (1984) The extinct Kangaroo Island Emu, a hitherto-unrecognised species. [Bulletin of the British Ornithologists' Club](/source/Bulletin_of_the_British_Ornithologists'_Club) 104: 19–22.

1. ^ [***a***](#cite_ref-Heupink_17-0) [***b***](#cite_ref-Heupink_17-1) Heupink, T. H.; Huynen, L.; Lambert, D. M. (2011). ["Ancient DNA Suggests Dwarf and 'Giant' Emu Are Conspecific"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3073985). *[PLoS ONE](/source/PLoS_ONE)*. **6** (4) e18728. [Bibcode](/source/Bibcode_(identifier)):[2011PLoSO...618728H](https://ui.adsabs.harvard.edu/abs/2011PLoSO...618728H). [doi](/source/Doi_(identifier)):[10.1371/journal.pone.0018728](https://doi.org/10.1371%2Fjournal.pone.0018728). [PMC](/source/PMC_(identifier)) [3073985](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3073985). [PMID](/source/PMID_(identifier)) [21494561](https://pubmed.ncbi.nlm.nih.gov/21494561).

1. **[^](#cite_ref-19)** Cole, Theresa L., et al. "Mitogenomes uncover extinct penguin taxa and reveal island formation as a key driver of speciation." Molecular biology and evolution 36.4 (2019): 784-797.

1. **[^](#cite_ref-Glaw_20-0)** 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"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3279364). *[PLoS ONE](/source/PLoS_ONE)*. **7** (2) e31314. [Bibcode](/source/Bibcode_(identifier)):[2012PLoSO...731314G](https://ui.adsabs.harvard.edu/abs/2012PLoSO...731314G). [doi](/source/Doi_(identifier)):[10.1371/journal.pone.0031314](https://doi.org/10.1371%2Fjournal.pone.0031314). [PMC](/source/PMC_(identifier)) [3279364](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3279364). [PMID](/source/PMID_(identifier)) [22348069](https://pubmed.ncbi.nlm.nih.gov/22348069).

1. **[^](#cite_ref-LangVogel_21-0)** de Lang R, Vogel G (2005). *The Snakes of Sulawesi: A Field Guide to the Land Snakes of Sulawesi with Identification Keys*. Frankfurt Contributions to Natural History Band 25, Edition Chimaira 2005. [ISBN](/source/ISBN_(identifier)) [3-930612-85-2](https://en.wikipedia.org/wiki/Special:BookSources/3-930612-85-2). pp. 23–27, 198–201.

1. **[^](#cite_ref-Barker2012_22-0)** Barker, D.G.; Barten, S.L.; Ehrsam, J.P.; Daddono, L. (2012). ["The Corrected Lengths of Two Well-known Giant Pythons and the Establishment of a New Maximum Length Record for Burmese Pythons, *Python bivittatus*"](https://ghostarchive.org/archive/20221009/http://www.vpi.com/sites/default/files/Barker-et-al_CorrectPythonLengths_2.pdf) (PDF). *Bulletin of the Chicago Herpetological Society*. **47** (1): 1–6. Archived from [the original](http://www.vpi.com/sites/default/files/Barker-et-al_CorrectPythonLengths_2.pdf) (PDF) on 2022-10-09. Retrieved 2020-03-02.

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1. **[^](#cite_ref-28)** Cheng-Han, Sun; Yi-Yang, Cho (2025-04-23). ["Rediscovering Mammoths in Taiwan: The Type Specimens of Mammuthus armeniacus taiwanicus (Proboscidea, Elephantidae)"](https://bioone.org/journals/mammal-study/volume-50/issue-3/ms2024-0028/Rediscovering-Mammoths-in-Taiwan--The-Type-Specimens-of-Mammuthus/10.3106/ms2024-0028.full). *Mammal Study*. **50** (3): 329–336.

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1. **[^](#cite_ref-Vuure2005_52-0)** van Vuure, Cis (2005). [*Retracing the Aurochs: History, Morphology and Ecology of an Extinct Wild Ox*](https://books.google.com/books?id=cyFFAAAAYAAJ&q=Sicily). Coronet Books Incorporated. [ISBN](/source/ISBN_(identifier)) [978-954-642-235-4](https://en.wikipedia.org/wiki/Special:BookSources/978-954-642-235-4). [OCLC](/source/OCLC_(identifier)) [472741798](https://search.worldcat.org/oclc/472741798).

1. **[^](#cite_ref-Geer2011_53-0)** van der Geer, A.; Lyras, G; de Vos, J.; Dermitzakis, M. (14 February 2011). ["Sardinia and Corsica"](https://books.google.com/books?id=JmSsNuwMAxgC&q=Chapter+Nine+Sardinia). [*Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands*](https://books.google.com/books?id=JmSsNuwMAxgC). John Wiley & Sons. [ISBN](/source/ISBN_(identifier)) [978-1-4443-9128-2](https://en.wikipedia.org/wiki/Special:BookSources/978-1-4443-9128-2). [OCLC](/source/OCLC_(identifier)) [894698082](https://search.worldcat.org/oclc/894698082).

1. **[^](#cite_ref-Chiozzi2014_54-0)** Chiozzi, G.; Bardelli, G.; Ricci, M.; De Marchi, G.; Cardini, A. (2014). ["Just another island dwarf? Phenotypic distinctiveness in the poorly known Soemmerring's Gazelle, *Nanger soemmerringii* (Cetartiodactyla: Bovidae), of Dahlak Kebir Island"](https://doi.org/10.1111%2Fbij.12239). *Biological Journal of the Linnean Society*. **111** (3): 603–620. [doi](/source/Doi_(identifier)):[10.1111/bij.12239](https://doi.org/10.1111%2Fbij.12239). [hdl](/source/Hdl_(identifier)):[11380/1061537](https://hdl.handle.net/11380%2F1061537).

1. ^ [***a***](#cite_ref-Mazza2015_55-0) [***b***](#cite_ref-Mazza2015_55-1) Mazza, P.P.A.; Rossi, M.A.; Agostini, S. (2015). "Hoplitomerycidae (Late Miocene, Italy), an Example of Giantism in Insular Ruminants". *Journal of Mammalian Evolution*. **22** (2): 271–277. [doi](/source/Doi_(identifier)):[10.1007/s10914-014-9277-2](https://doi.org/10.1007%2Fs10914-014-9277-2). [S2CID](/source/S2CID_(identifier)) [16437411](https://api.semanticscholar.org/CorpusID:16437411).

1. **[^](#cite_ref-Geer2018_56-0)** van der Geer, A.A.E. (2018). ["Uniformity in variety: Antler morphology and evolution in a predator-free environment"](https://doi.org/10.26879%2F834). *Palaeontologia Electronica* (21.1.9A): 1–31. [doi](/source/Doi_(identifier)):[10.26879/834](https://doi.org/10.26879%2F834).

1. **[^](#cite_ref-Kaifu2015_58-0)** Kaifu, Y.; Fujita, M.; Yoneda, M.; Yamasaki, S. (15 February 2015). ["Pleistocene Seafaring and Colonization of the Ryukyu Islands, Southwestern Japan"](https://books.google.com/books?id=AsgbBgAAQBAJ&q=Cervus+astylodon). In Kaifu, Y.; Izuho, M.; Goebel, T.; Sato, H.; Ono, A. (eds.). [*Emergence and Diversity of Modern Human Behavior in Paleolithic Asia*](https://books.google.com/books?id=AsgbBgAAQBAJ). Texas A&M University Press. [ISBN](/source/ISBN_(identifier)) [978-1-62349-277-9](https://en.wikipedia.org/wiki/Special:BookSources/978-1-62349-277-9). [OCLC](/source/OCLC_(identifier)) [985023261](https://search.worldcat.org/oclc/985023261).

1. **[^](#cite_ref-59)** Lister, A. M. (1989-11-30). "Rapid dwarfing of red deer on Jersey in the Last Interglacial". *[Nature](/source/Nature_(journal))*. **342** (6249): 539–542. [Bibcode](/source/Bibcode_(identifier)):[1989Natur.342..539L](https://ui.adsabs.harvard.edu/abs/1989Natur.342..539L). [doi](/source/Doi_(identifier)):[10.1038/342539a0](https://doi.org/10.1038%2F342539a0). [PMID](/source/PMID_(identifier)) [2685610](https://pubmed.ncbi.nlm.nih.gov/2685610). [S2CID](/source/S2CID_(identifier)) [4343091](https://api.semanticscholar.org/CorpusID:4343091).

1. **[^](#cite_ref-Wilder2010_61-0)** Wilder, B.T.; Felger, R.S. (30 September 2010). ["Dwarf Giants, Guano, and Isolation: Vegetation and Floristic Diversity of San Pedro Mártir Island, Gulf of California, Mexico"](https://brccapi.sdnhm.org/files/8213/7106/5625/Proceedings42_COLOR1.pdf) (PDF). *Proceedings of the San Diego Society of Natural History*. **42**: 1–24, see pp. 9–13. Retrieved 2020-01-05. (p. 12) The dwarfing of the San Pedro Mártir plants seems to be due to a selection for shorter individuals to survive fierce tropical storms, possible root competition in such a dense forest, and the undefined effect of high levels of nitrogen and phosphorus from the abundant guano that might stunt growth. Genetic studies have not been undertaken...

1. **[^](#cite_ref-Burns2019_62-0)** Burns, K.C. (May 2019). [*Evolution in Isolation: The Search for an Island Syndrome in Plants*](https://books.google.com/books?id=wbOQDwAAQBAJ). [Cambridge University Press](/source/Cambridge_University_Press). pp. 174–177. [doi](/source/Doi_(identifier)):[10.1017/9781108379953](https://doi.org/10.1017%2F9781108379953). [ISBN](/source/ISBN_(identifier)) [978-1-108-37995-3](https://en.wikipedia.org/wiki/Special:BookSources/978-1-108-37995-3). [OCLC](/source/OCLC_(identifier)) [1108160200](https://search.worldcat.org/oclc/1108160200). [S2CID](/source/S2CID_(identifier)) [186536407](https://api.semanticscholar.org/CorpusID:186536407). (pp. 174-175) ... the extent to which its dwarfed stature is genetically determined, and an explanation for why insular dwarfism might be selectively advantageous, awaits additional study.

## External links

- [Strange world of island species October 31, 2004 The Observer](https://www.theguardian.com/life/science/story/0,12996,1340300,00.html)

v t e Biological rules Rules Allen's rule Shorter appendages in colder climates Bateson's rule Extra limbs mirror their neighbours Bergmann's rule Larger bodies in colder climates Cope's rule Bodies get larger over time Deep-sea gigantism Larger bodies in deep-sea animals Dollo's law Loss of complex traits is irreversible Eichler's rule Parasites co-vary with their hosts Emery's rule Insect social parasites are often in same genus as their hosts Fahrenholz's rule Host and parasite phylogenies become congruent Foster's rule (Insular gigantism, Insular dwarfism) Small species get larger, large species smaller, after colonizing islands Gause's law Complete competitors cannot coexist Gloger's rule Lighter coloration in colder, drier climates Haldane's rule Hybrid sexes that are absent, rare, or sterile, are heterogamic Harrison's rule Parasites co-vary in size with their hosts Hamilton's rule Genes increase in frequency when relatedness of recipient to actor times benefit to recipient exceeds reproductive cost to actor Kleiber's law An animals metabolic rate decreases with its size Hennig's progression rule In cladistics, the most primitive species are found in earliest, central, part of group's area Jarman–Bell principle The correlation between the size of an animal and its diet quality; larger animals can consume lower quality diet Jordan's rule Inverse relationship between water temperature and no. of fin rays, vertebrae Lack's principle Birds lay only as many eggs as they can provide food for Rapoport's rule Latitudinal range increases with latitude Rensch's rule Sexual size dimorphism increases with size when males are larger, decreases with size when females are larger Rosa's rule Groups evolve from character variation in primitive species to a fixed character state in advanced ones Schmalhausen's law A population at limit of tolerance in one aspect is vulnerable to small differences in any other aspect Thayer's law The top of an animals coloration is darker than the bottom Thorson's rule Number of eggs of benthic marine invertebrates decreases with latitude Van Valen's law Probability of extinction of a group is constant over time von Baer's laws Embryos start from a common form and develop into increasingly specialised forms Williston's law Parts in an organism become reduced in number and specialized in function Related Countergradient variation Where genetics opposes environment as a factor Gigantothermy Large ectothermic animals more easily maintain constant body temperature

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Adapted from the Wikipedia article [Insular dwarfism](https://en.wikipedia.org/wiki/Insular_dwarfism) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Insular_dwarfism?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.