Biology:Insular dwarfism

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Short description: Form of phyletic dwarfism occurring on islands
For other uses, see Dwarf (disambiguation).
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.

Insular dwarfism, a form of phyletic dwarfism,[1] is the process and condition of large animals evolving or having a reduced body size[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 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"). 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.

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.Cite error: Closing </ref> missing for <ref> tag || L. astibiae || Ibero-Armorican Island || Late Cretaceous |- | 120px
Paludititan || P. nalatzensis || Hațeg Island || Late Cretaceous / Maastrichtian || 120px
Epachthosaurus |}

Other

Example Species Range Time frame Continental relative
120px
Langenberg Quarry
torvosaur (blue)		 Unnamed Lower Saxony Late Jurassic / Middle Kimmeridgian 120px
Torvosaurus
120x120px
Struthiosaurus[9]		 S. austriacus Ibero-Armorican, Australoalpine, and Hațeg Islands Late Cretaceous 120px
Edmontonia
S. transylvanicus
S. languedocensis
120px
Telmatosaurus		 T. transsylvanicus Hațeg Island 120px
Hadrosaurids
120px
Thecodontosaurus[10]		 T. antiquus Southern England Late Triassic / Rhaetian 120px
Plateosaurs
120px
Zalmoxes[10] (purple)		 Z. robustus Hațeg Island Late Cretaceous 120px
Tenontosaurus
Z. shqiperorum

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

Example Binomial name Native range Status Continental relative Insular / mainland
length or mass ratio
120px
Hawaiian flightless ibises		 Apteribis glenos Molokai Extinct (Late Quaternary) 120px
American ibises
Apteribis brevis Maui
Cozumel curassow[11] Crax rubra griscomi Cozumel Unknown 120px
Great curassow
120px
Kangaroo Island emu[12]		 Dromaius novaehollandiae baudinianus Kangaroo Island, South Australia Extinct (c. AD 1827) 120px
Emu
120px
King Island emu[13] (black)		 Dromaius novaehollandiae minor King Island, Tasmania Extinct (AD 1822) LR ≈ 0.48[lower-alpha 2]
Dwarf yellow eyed penguin[14] Megadyptes antipodes richdalei Chatham Islands, New Zealand Extinct (after 1300 AD) 120px
Yellow-eyed penguin
120px
Cozumel thrasher[11]		 Toxostoma gluttatum Cozumel Critically endangered 120px
Other thrashers

Squamates

Example Binomial name Native range Status Continental relative Insular / mainland
length or mass ratio
120px
Madagascar dwarf chameleon		 Brookesia minima Nosy Be island, Madagascar Endangered 120px
Madagascar leaf chameleons
120px
Nosy Hara chameleon[15]		 Brookesia micra Nosy Hara island, Madagascar Vulnerable
Roxby Island tiger snake[5] Notechis scutatus Roxby Island, South Australia Unknown 120px
Tiger snake
Dwarf Burmese python Python bivittatus progschai Java, Bali, Sumbawa and Sulawesi, Indonesia 120px
Burmese python		 LR ≈ 0.44[lower-alpha 3]
Tanahjampea reticulated python[18] Python reticulatus jampeanus Tanahjampea, between Sulawesi and Flores 120px
Reticulated python		 LR ≈ 0.41, males
LR ≈ 0.49, females[lower-alpha 4]

Mammals

Pilosans

Example Binomial name Native range Status Continental relative
120px
Pygmy three-toed sloth		 Bradypus pygmaeus Isla Escudo de Veraguas, Panama Critically endangered 120px
Brown-throated sloth
120px
Acratocnus		 A. antillensis Cuba, Hispaniola and Puerto Rico Extinct (c. 3000 BC) 120px
Continental ground sloths
A. odontrigonus
A. ye
Imagocnus I. zazae Cuba Extinct (Early Miocene)
120px
Megalocnus		 M. rodens Cuba and Hispaniola Extinct (c. 2700 BC)
M. zile
120px
Neocnus		 Neocnus spp. Extinct (c. 3000 BC)

Proboscideans

Example Binomial name Native range Status Continental relative
Sulawesi dwarf elephant Elephas celebensis Sulawesi Extinct (Early Pleistocene) 120px
Asian elephant
Cabarruyan dwarf elephant Elephas beyeri Luzon Extinct
Cretan dwarf mammoth Mammuthus creticus Crete 120px
Mammuthus
120px
Channel Islands mammoth		 Mammuthus exilis Santa Rosae island Extinct (Late Pleistocene) 120px
Columbian mammoth
120px

Sardinian mammoth

Mammuthus lamarmorai Sardinia 120px
Steppe mammoth
Saint Paul Island woolly mammoth[21][22] Mammuthus primigenius Saint Paul Island, Alaska Extinct (c. 3750 BC) 120px
Woolly mammoth
120px
Siculo-Maltese elephants		 Palaeoloxodon antiquus leonardi Sicily and Malta Extinct 150px
Straight-tusked elephant
(left)
P. mnaidriensis
P. melitensis
P. falconeri
Cretan elephants Palaeoloxodon chaniensis
 Crete
P. creutzburgi
120px
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[23] Sinomastodon bumiajuensis Bumiayu Island (now part of Java) Extinct (Early Pleistocene) 120px
Sinomastodon
120px
Japanese stegodont[24][25]		 Stegodon miensis Japan (Also Taiwan for S. aurorae)[26] 120px
Chinese Stegodon
Stegodon protoaurorae
Stegodon aurorae
Javan dwarf stegodonts Stegodon hypsilophus[23] Java Extinct (Quaternary)
S. semedoensis[27]
S. sp.[23]
Mindanao pygmy stegodont[28] Stegodon mindanensis Mindanao and Sulawesi Extinct (Middle Pleistocene)
Sulawesi dwarf stegodont[23] Stegodon sompoensis Sulawesi Extinct
Lesser Flores dwarf stegodont[3] Stegodon sondaari Flores Extinct (Middle Pleistocene)
Sumba dwarf stegodont[29] Stegodon sumbaensis Sumba, Indonesia
Timor dwarf stegodont[23] Stegodon timorensis Timor Extinct
Dwarf stegolophodont[30] Stegolophodon pseudolatidens Japan Extinct (Miocene) 120px
Stegolophodon

Primates

Example Binomial name Native range Status Continental relative
Nosy Hara dwarf lemur[31] Cheirogaleus sp. Nosy Hara island off Madagascar Unknown 120px
Dwarf lemurs
120px
Flores Man[32]		 Homo floresiensis Flores Extinct (Late Pleistocene) 100px
Homo erectus
120px
Callao Man		 Homo luzonensis[33][34] Luzon, Philippines
Modern pygmies of Flores[35] Homo sapiens Flores Extant other members of Homo sapiens
Early Palau modern humans (disputed)[36] Palau Extinct (?)
Andamanese[37] Andaman Islands Extant
120px
Sardinian macaque[38]		 Macaca majori Sardinia Extinct (Pleistocene) 120px
Barbary macaque
120px
Zanzibar red colobus		 Piliocolobus kirkii Unguja Endangered 120px
Udzungwa red colobus

Carnivorans

Example Binomial name Native range Status Continental relative Insular / mainland
length or mass ratio
120px
Sicilian wolf		 Canis lupus cristaldii Sicily Extinct (AD 1970) 120px
Gray wolf
120px
Japanese wolf		 Canis lupus hodophilax Japan (excluding Hokkaido) Extinct (AD 1905)
120px
Sardinian dhole
(forward)		 Cynotherium sardous Corsica and Sardinia Extinct (c. 8300 BC) 120px
Xenocyon
Trinil dog Mececyon trinilensis Java Extinct (Pleistocene)
Cozumel Island coati[11] Nasua narica nelsoni Cozumel Critically endangered 120px
Yucatan white-nosed coati
120px
Zanzibar leopard		 Panthera pardus pardus Unguja Critically endangered or Extinct 120px
African leopard
120px
Bali tiger		 Panthera tigris sondaica Bali Extinct (c. AD 1940) 120px
Sumatran tiger
120px
Javan tiger		 Java Extinct (c. AD 1975)
120px
Cozumel raccoon		 Procyon pygmaeus Cozumel Critically endangered 120px
Common raccoon
120px
Island fox		 Urocyon littoralis Six of the Channel Islands of California Near Threatened 120px
Gray fox		 LR ≈ 0.84[lower-alpha 5]
LR ≈ 0.75[lower-alpha 6]
Cozumel fox Urocyon sp. Cozumel Critically endangered or Extinct

Non-ruminant ungulates

Example Binomial name Native range Status Continental relative
120px
Eumaiochoerus		 Eumaiochoerus etruscus Baccinello, Montebamboli Extinct (Miocene) 120px
Microstonyx
120px
Malagasy dwarf hippopotamuses		 Hippopotamus laloumena Madagascar Extinct (c. AD 1000) 120px
Common hippopotamus
H. lemerlei
H. madagascariensis
Bumiayu dwarf hippopotamus[23] Hexaprotodon simplex Bumiayu Island (now Java) Extinct (Early Pleistocene) 120px
Asian hippopotamuses
120px
Cretan dwarf hippopotamus		 Hippopotamus creutzburgi Crete Extinct (Middle Pleistocene) 120px
Hippopotamus antiquus
120px
Maltese dwarf hippopotamus		 Hippopotamus melitensis Malta Extinct (Pleistocene) 120px
Common hippopotamus (H. amphibius)
120px
Sicilian dwarf hippopotamus		 Hippopotamus pentlandi Sicily
120px
Cyprus dwarf hippopotamus		 Hippopotamus minor Cyprus Extinct (c. 8000 BC) Unclear, either H. amphibius or H. antiquus.
Cozumel collared peccary[11] Pecari tajacu nanus Cozumel Unknown 120px
Collared peccary

Bovids

Example Binomial name Native range Status Continental relative
Sicilian bison[24] Bison priscus siciliae Sicily Extinct (Late Pleistocene) 120px
Steppe bison
Sicilian aurochs[41] Bos primigenius siciliae[24] 120px
Eurasian aurochs
Cebu tamaraw Bubalus cebuensis Cebu, Philippines Extinct 120px
Wild water buffalo
120px
Lowland anoa		 Bubalus depressicornis Sulawesi and Buton, Indonesia Endangered
Bubalus grovesi Bubalus grovesi Sulawesi, Indonesia Extinct
120px
Tamaraw		 Bubalus mindorensis Mindoro, Philippines Critically endangered
120px
Mountain anoa		 Bubalus quarlesi Sulawesi and Buton, Indonesia Endangered
120px
Balearic Islands cave goat		 Myotragus balearicus Majorca and Menorca Extinct (after 3000 BC) Gallogoral
Nesogoral[42] Nesogoral spp. Sardinia Extinct
Dahlak Kebir gazelle[43] Nanger soemmerringi ssp. Dahlak Kebir island, Eritrea Vulnerable 120px
Soemmerring's gazelle
120px
Tyrrhenotragus		 Tyrrhenotragus gracillimus Baccinello Extinct Antilopinae sp.

Cervids and relatives

Example Binomial name Native range Status Continental relative
120px
Cretan deer[lower-alpha 7]		 Candiacervus spp. Crete Extinct (Pleistocene) Unknown
150px
Sardinian deer[10]		 Praemegaceros cazioti Sardinia Extinct (c. 5500 BC) Praemegaceros
120px
Ryukyu dwarf deer[46]		 Cervus astylodon Ryukyu Islands Extinct 120px
Sika deer (?)
Cervus praenipponicus (?)
Jersey red deer population[47] Cervus elaphus jerseyensis Jersey Extinct (Pleistocene) 120px
Red deer
120px
Corsican red deer		 Cervus elaphus corsicanus Corsica and Sardinia Near Threatened
Sicilian red deer[24] Cervus siciliae Sicily Extinct (Late Pleistocene)
120px
Hoplitomeryx[lower-alpha 8]		 Hoplitomeryx spp. Gargano Island Extinct (Early Pliocene) 120px
Pecorans
Sicilian fallow deer Dama carburangelensis Sicily Extinct (Late Pleistocene) Fallow deer
120px
Florida Key deer		 Odocoileus virginianus clavium Florida Keys Endangered 120px
Virginia deer
120px
Svalbard reindeer		 Rangifer tarandus platyrhynchus Svalbard Vulnerable 120px
Reindeer
120px
Philippine deer		 Rusa marianna Philippines 120px
Sambar deer

Plants

Possible example Binomial name Native range Status Continental relative
120px
Insular elephant cacti[48][49]		 Pachycereus pringlei Remote islands in the Sea of Cortez (e.g. Santa Cruz, San Pedro Mártir) Not evaluated 120px
Mainland elephant cacti

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[13]
  3. ↑ Based on maximum lengths of 2.5 m for the dwarf form[16] and 5.74 m for the mainland form[17]
  4. ↑ Based on maximum Tanahjampea python total lengths (TL) of 2.10 m for males and 3.35 m for females[18] and maximum southern Sumatra python snout to vent lengths (SVL) of 4.5 m for males and 6.1 m for females[19] 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[20]
  5. ↑ For nearby mainland gray foxes[39]
  6. ↑ For mainland gray foxes in general[40]
  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.[44][45]
  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).[44]

References

  1. ↑ Prothero, Donald Ross; Sereno, Paul Callistus (Winter 1982). "Allometry and Paleoecology of Medial Miocene Dwarf Rhinoceroses from the Texas Gulf Coastal Plain". Paleobiology 8 (1): 16–30. doi:10.1017/S0094837300004322. Bibcode: 1982Pbio....8...16P. https://www.cambridge.org/core/journals/paleobiology/article/abs/allometry-and-paleoecology-of-medial-miocene-dwarf-rhinoceroses-from-the-texas-gulf-coastal-plain/F52722F0471E07FE01184D92C19E1C61. 
  2. ↑ Perelman, P. (2011). "A Molecular Phylogeny of Living Primates". PLOS Genetics 7 (3): 1–17. doi:10.1371/journal.pgen.1001342. PMID 21436896. 
  3. ↑ 3.0 3.1 3.2 3.3 Cite error: Invalid <ref> tag; no text was provided for refs named Van Den Bergh
  4. ↑ 4.0 4.1 4.2 4.3 Raia, Pasquale; Meiri, Shai (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. PMID 17017072. 
  5. ↑ 5.0 5.1 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. PMID 15792242. 
  6. ↑ 6.0 6.1 Evans, A. R. (2012-01-30). "The maximum rate of mammal evolution". PNAS 109 (11): 4187–4190. doi:10.1073/pnas.1120774109. PMID 22308461. Bibcode: 2012PNAS..109.4187E. 
  7. ↑ 7.0 7.1 "Dinosaurs, dragons, and dwarfs: The evolution of maximal body size". Proceedings of the National Academy of Sciences 98 (25): 14518–14523. 2001-12-04. doi:10.1073/pnas.251548698. ISSN 0027-8424. PMID 11724953. Bibcode: 2001PNAS...9814518B. 
  8. ↑ van der Geer, Alexandra A. E.; van den Bergh, Gerrit D.; Lyras, George A.; Prasetyo, Unggul W.; Due, Rokus Awe; Setiyabudi, Erick; Drinia, Hara (August 2016). "The effect of area and isolation on insular dwarf proboscideans" (in en). Journal of Biogeography 43 (8): 1656–1666. doi:10.1111/jbi.12743. ISSN 0305-0270. Bibcode: 2016JBiog..43.1656V. https://onlinelibrary.wiley.com/doi/10.1111/jbi.12743. 
  9. ↑ Carpenter, K. (2001) The Armored Dinosaurs. Indiana University Press, 526 pages.
  10. ↑ 10.0 10.1 10.2 Cite error: Invalid <ref> tag; no text was provided for refs named Benton2010
  11. ↑ 11.0 11.1 11.2 11.3 Cuarón, A. D.; Martínez-Morales, M. A.; McFadden, K. W.; Valenzuela, D.; Gompper, M. E. (2004). "The status of dwarf carnivores on Cozumel Island, Mexico". Biodiversity and Conservation 13 (2): 317–331. doi:10.1023/b:bioc.0000006501.80472.cc. Bibcode: 2004BiCon..13..317C. 
  12. ↑ Parker S (1984) The extinct Kangaroo Island Emu, a hitherto-unrecognised species. Bulletin of the British Ornithologists' Club 104: 19–22.
  13. ↑ 13.0 13.1 Heupink, T. H.; Huynen, L.; Lambert, D. M. (2011). "Ancient DNA Suggests Dwarf and 'Giant' Emu Are Conspecific". PLoS ONE 6 (4). doi:10.1371/journal.pone.0018728. PMID 21494561. Bibcode: 2011PLoSO...618728H. 
  14. ↑ 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.
  15. ↑ 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). doi:10.1371/journal.pone.0031314. PMID 22348069. Bibcode: 2012PLoSO...731314G. 
  16. ↑ 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 3-930612-85-2. pp. 23–27, 198–201.
  17. ↑ 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". Bulletin of the Chicago Herpetological Society 47 (1): 1–6. http://www.vpi.com/sites/default/files/Barker-et-al_CorrectPythonLengths_2.pdf. Retrieved 2020-03-02. 
  18. ↑ 18.0 18.1 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. PMID 12135085. Bibcode: 2002NW.....89..201A. 
  19. ↑ Shine, R.; Harlow, P.S.; Keogh, J.S.; Boeadi, N.I. (1998). "The influence of sex and body size on food habits of a giant tropical snake, Python reticulatus ". Functional Ecology 12 (2): 248–258. doi:10.1046/j.1365-2435.1998.00179.x. Bibcode: 1998FuEco..12..248S. 
  20. ↑ Sheehy, C.M.; Albert, J.S.; Lillywhite, H.B.; Van Damme, R. (2016). "The evolution of tail length in snakes associated with different gravitational environments". Functional Ecology 30 (2): 244–254. doi:10.1111/1365-2435.12472. Bibcode: 2016FuEco..30..244S. ; see Table S1
  21. ↑ Schirber, Michael. Surviving Extinction: Where Woolly Mammoths Endured. Live Science. Imaginova Corporation. Retrieved 2007-07-20.
  22. ↑ The mammoths of Wrangel Island, north of Siberia, are no longer considered dwarfs. See: 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
  23. ↑ 23.0 23.1 23.2 23.3 23.4 23.5 Aziz, F.; van den Bergh, G. D. (September 25, 1995). "A dwarf Stegodon from Sambungmacan (Central Java, Indonesia)" (in en). Proc. Kon. Ned. Akad. V. Wetensch. 98 (3): 229–241. https://www.researchgate.net/publication/275753718. Retrieved 2017-07-31. 
  24. ↑ 24.0 24.1 24.2 24.3 Sondaar, P. Y.; A.A.E. van der Geer (2005). "Evolution and Extinction of Plio-Pleistocene Island Ungulates" (in en). International Journal of the French Quaternary Association 2: 241–256. https://www.researchgate.net/publication/242279220. Retrieved 2017-07-31. 
  25. ↑ Aiba, Hiroaki; Baba, Katsuyoshi; Matsukawa, Masaki (2010-03-10). "A new species of Stegodon (Mammalia, Proboscidea) from the Kazusa Group (lower Pleistocene), Hachioji City, Tokyo, Japan and its evolutionary morphodynamics: STEGODON PROTOAURORAE SP. NOV. AND MORPHODYNAMICS" (in en). Palaeontology 53 (3): 471–490. doi:10.1111/j.1475-4983.2010.00953.x. https://onlinelibrary.wiley.com/doi/10.1111/j.1475-4983.2010.00953.x. 
  26. ↑ van den Bergh, Gert D.; de Vos, John; Sondaar, Paul Y. (25 September 2000). "The Late Quaternary palaeogeography of mammal evolution in the Indonesian Archipelago". Palaeogeography, Palaeoclimatology, Palaeoecology 171 (3–4): 385–408. doi:10.1016/S0031-0182(01)00255-3. http://www.rhinoresourcecenter.com/pdf_files/129/1291330178.pdf. 
  27. ↑ Siswanto, S., & Noerwidi, S. (2014). PROBOSCIDEA FOSSIL FROM SEMEDO SITE: Its Correlation With Biostratigraphy and Human Arrival in Java. Berkala Arkeologi, 34(2).
  28. ↑ Zaim, Y. (20 August 2010). "Geological Evidence for the Earliest Appearance of Hominins in Indonesia". in Fleagle, J. G; Shea, J. J.; Grine, F. E. et al.. Out of Africa I: The First Hominin Colonization of Eurasia. Springer Science & Business Media. p. 106. ISBN 978-90-481-9036-2. OCLC 668096676. https://books.google.com/books?id=CO5zfl460CEC&pg=PA106. 
  29. ↑ Setiyabudi, Erick; Kurniawan, Iwan; Van Den Bergh, Gerrit. "Fossils of Stegodon and Varanus komodoensis Sumba and Flores: a Pleistocene landbridge". Faculty of Science, Medicine and Health. https://ro.uow.edu.au/cgi/viewcontent.cgi?referer=https://en.wikipedia.org/&httpsredir=1&article=3055&context=smhpapers. 
  30. ↑ Saegusa, H. (2008). "Dwarf Stegolophodon from the Miocene of Japan: Passengers on sinking boats". Quaternary International 182 (1): 49–62. doi:10.1016/j.quaint.2007.08.001. Bibcode: 2008QuInt.182...49S. 
  31. ↑ "New group of dwarf lemurs may be world's rarest primate". http://www.bbc.com/earth/story/20150812-tiny-lemur-may-be-worlds-rarest. 
  32. ↑ Scientist to study Hobbit morphing, abc.net.au
  33. ↑ Wade, L. (2019-04-10). "New species of ancient human unearthed in the Philippines". Science 364. doi:10.1126/science.aax6501. 
  34. ↑ Détroit, F.; Mijares, A. S.; Corny, J.; Daver, G.; Zanolli, C.; Dizon, E.; Robles, E.; Grün, R. et al. (2019). "A new species of Homo from the Late Pleistocene of the Philippines". Nature 568 (7751): 181–186. doi:10.1038/s41586-019-1067-9. PMID 30971845. Bibcode: 2019Natur.568..181D. https://hal.archives-ouvertes.fr/hal-02296712/file/Detroit_%26_al_2019_Nature_postprint.pdf. 
  35. ↑ Tucci, S. (2018-08-03). "Evolutionary history and adaptation of a human pygmy population of Flores Island, Indonesia". Science 361 (6401): 511–516. doi:10.1126/science.aar8486. PMID 30072539. Bibcode: 2018Sci...361..511T. 
  36. ↑ "Ancient Small People on Palau Not Dwarfs, Study Says". National Geographic News. August 27, 2008.
  37. ↑ Mondal, M.; Casals, F.; Xu, T.; Dall'Olio, G. M.; Pybus, M.; Netea, M. G.; Comas, D.; Laayouni, H. et al. (2016). "Genomic analysis of Andamanese provides insights into ancient human migration into Asia and adaptation". Nature Genetics 48 (9): 1066–1070. doi:10.1038/ng.3621. PMID 27455350. http://repositori.upf.edu/bitstream/10230/34401/1/Mondal_NG_Gen.pdf. 
  38. ↑ Rook, L. (2008-12-31). "The first workshop on European fossil primate record (Siena and Grosseto, September 11-13, 2008) with an update on Italian studies in Paleoprimatology". Atti Muss. Stor. Nat. Maremma (22): 129–143. http://museonatura.comune.grosseto.it/fileadmin/templates/pdf/ATTI_DEL_MUSEO/Volume_n.22/Rook_atti_22_129-143.pdf. Retrieved 2019-06-20. 
  39. ↑ Parfit, M.; Groo, M. (22 April 2020). "The uplifting tale of these tiny island foxes, nearly wiped out by disaster". National Geographic. https://www.nationalgeographic.com/animals/2020/04/santa-cruz-island-foxes/. 
  40. ↑ Moore, C.M.; Collins, P.W. (1995). Mammalian Species – Urocyon littoralis. 489. pp. 1–7. http://www.science.smith.edu/departments/Biology/VHAYSSEN/msi/pdf/i0076-3519-489-01-0001.pdf. Retrieved 2011-09-16. 
  41. ↑ van Vuure, Cis (2005). Retracing the Aurochs: History, Morphology and Ecology of an Extinct Wild Ox. Coronet Books Incorporated. ISBN 978-954-642-235-4. OCLC 472741798. https://books.google.com/books?id=cyFFAAAAYAAJ&q=Sicily. 
  42. ↑ van der Geer, A.; Lyras, G; de Vos, J.; Dermitzakis, M. (14 February 2011). "Sardinia and Corsica". Evolution of Island Mammals: Adaptation and Extinction of Placental Mammals on Islands. John Wiley & Sons. ISBN 978-1-4443-9128-2. OCLC 894698082. https://books.google.com/books?id=JmSsNuwMAxgC&q=Chapter+Nine+Sardinia. 
  43. ↑ 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". Biological Journal of the Linnean Society 111 (3): 603–620. doi:10.1111/bij.12239. 
  44. ↑ 44.0 44.1 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:10.1007/s10914-014-9277-2. 
  45. ↑ van der Geer, A.A.E. (2018). "Uniformity in variety: Antler morphology and evolution in a predator-free environment". Palaeontologia Electronica (21.1.9A): 1–31. doi:10.26879/834. 
  46. ↑ Kaifu, Y.; Fujita, M.; Yoneda, M.; Yamasaki, S. (15 February 2015). "Pleistocene Seafaring and Colonization of the Ryukyu Islands, Southwestern Japan". in Kaifu, Y.; Izuho, M.; Goebel, T. et al.. Emergence and Diversity of Modern Human Behavior in Paleolithic Asia. Texas A&M University Press. ISBN 978-1-62349-277-9. OCLC 985023261. https://books.google.com/books?id=AsgbBgAAQBAJ&q=Cervus+astylodon. 
  47. ↑ 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. Bibcode: 1989Natur.342..539L. 
  48. ↑ 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". Proceedings of the San Diego Society of Natural History 42: 1–24; see pp. 9–13. https://brccapi.sdnhm.org/files/8213/7106/5625/Proceedings42_COLOR1.pdf. 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...". 
  49. ↑ Burns, K.C. (May 2019). Evolution in Isolation: The Search for an Island Syndrome in Plants. Cambridge University Press. pp. 174–177. doi:10.1017/9781108379953. ISBN 978-1-108-37995-3. OCLC 1108160200. https://books.google.com/books?id=wbOQDwAAQBAJ. "(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." 



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