Insular dwarfism

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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 in shoulder height and weighed about 250 kg (550 lb). Females were smaller.

Possible causes

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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 & gigantism

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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

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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

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Non-avian dinosaurs

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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

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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 Hateg Island Late Cretaceous / Maastrichtian  
Rapetosaurus
 
Lirainosaurus[12]
L. astibiae Ibero-Armorican Island Late Cretaceous
 
Paludititan
P. nalatzensis Hateg Island Late Cretaceous / Maastrichtian  
Epachthosaurus

Other

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Example Species Range Time frame Continental relative
 
Langenberg Quarry
torvosaur (blue)
Unnamed Lower Saxony Late Jurassic / Middle Kimmeridgian  
Torvosaurus
 
Struthiosaurus[13]
S. austriacus

S. transylvanicus

S. languedocensis
Ibero-Armorican, Australoalpine, and Hateg islands Late Cretaceous  
Edmontonia
 
Telmatosaurus
T. transsylvanicus Hateg Island Late Cretaceous  
Hadrosaurids
 
Thecodontosaurus[10]
T. antiquus Southern England Late Triassic / Rhaetian  
Plateosaurs
 
Zalmoxes[10] (purple)
Z. robustus

Z. shqiperorum
Hateg Island Late Cretaceous  
Tenontosaurus

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

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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
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 Unknown  
Burmese python
LR ≈ 0.44 [c]
Tanahjampea reticulated python[21] Python reticulatus jampeanus Tanahjampea, between Sulawesi and Flores Unknown  
Reticulated python
LR ≈ 0.41, males
LR ≈ 0.49, females [d]

Mammals

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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

A. odontrigonus

A. ye
Cuba, Hispaniola and Puerto Rico Extinct (c. 3000 BC)  
Continental ground sloths
Imagocnus I. zazae Cuba Extinct (Early Miocene)
 
Megalocnus
M. rodens

M. zile
Cuba and Hispaniola Extinct (c. 2700 BC)
 
Neocnus
Neocnus spp. Cuba and Hispaniola Extinct (c. 3000 BC)
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 Extinct  
Mammuthus
 
Channel Islands mammoth
Mammuthus exilis Santa Rosae island Extinct (Late Pleistocene)  
Columbian mammoth
Sardinian mammoth Mammuthus lamarmorai Sardinia Extinct (Late Pleistocene)  
Steppe mammoth
Saint Paul Island woolly mammoth[24][25] Mammuthus primigenius Saint Paul Island, Alaska Extinct (c. 3750 BC)  
Woolly mammoth
 
Siculo-Maltese elephants
Palaeoloxodon antiquus leonardi

P. mnaidriensis

P. melitensis

P. falconeri
Sicily and Malta Extinct  
Straight-tusked elephant
(left)
Cretan elephants Palaeoloxodon chaniensis

P. creutzburgi
Crete Extinct
 
Cyprus dwarf elephant
Palaeoloxodon cypriotes Cyprus Extinct (c. 9000 BC)
Naxos dwarf elephant Palaeoloxodon sp. Naxos Extinct
Rhodes and Tilos dwarf elephant Palaeoloxodon tiliensis Rhodes and Tilos Extinct
Bumiayu dwarf sinomastodont[26] Sinomastodon bumiajuensis Bumiayu Island (now part of Java) Extinct (Early Pleistocene)  
Sinomastodon
 
Japanese stegodont[27][28]
Stegodon miensis

Stegodon protoaurorae

Stegodon aurorae
Japan (Also Taiwan for S. aurorae)[29] Extinct (Early Pleistocene)  
Chinese Stegodon
Greater Flores dwarf stegodont[3] Stegodon florensis Flores Extinct (Late Pleistocene)  
Sundaland Stegodon
Javan dwarf stegodonts Stegodon hypsilophus[26]

S. semedoensis[30]

S. sp.[26]
Java Extinct (Quaternary)
Mindanao pygmy stegodont[31] Stegodon mindanensis Mindanao and Sulawesi Extinct (Middle Pleistocene)
Sulawesi dwarf stegodont[26] Stegodon sompoensis Sulawesi Extinct
Lesser Flores dwarf stegodont[3] Stegodon sondaari Flores Extinct (Middle Pleistocene)
Sumba dwarf stegodont[32] Stegodon sumbaensis Sumba, Indonesia Extinct (Middle Pleistocene)
Timor dwarf stegodont[26] Stegodon timorensis Timor Extinct
Dwarf stegolophodont[33] Stegolophodon pseudolatidens Japan Extinct (Miocene)  
Stegolophodon
Example Binomial name Native range Status Continental relative
Nosy Hara dwarf lemur[34] Cheirogaleus sp. Nosy Hara island off Madagascar Unknown  
Dwarf lemurs
 
Flores Man[35]
Homo floresiensis Flores Extinct (Late Pleistocene)  
Homo erectus
 
Callao Man
Homo luzonensis[36][37] Luzon, Philippines Extinct (Late Pleistocene)
Modern pygmies of Flores[38] Homo sapiens Flores Extant other members of Homo sapiens
Early Palau modern humans (disputed)[39] Homo sapiens Palau Extinct (?)
Andamanese[40] Homo sapiens Andaman Islands Extant
 
Sardinian macaque[41]
Macaca majori Sardinia Extinct (Pleistocene)  
Barbary macaque
 
Zanzibar red colobus
Piliocolobus kirkii Unguja Endangered  
Udzungwa red colobus
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
 
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

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Example Binomial name Native range Status Continental relative
 
Eumaiochoerus
Eumaiochoerus etruscus Baccinello, Montebamboli Extinct (Miocene)  
Microstonyx
 
Malagasy dwarf hippopotamuses
Hippopotamus laloumena

H. lemerlei

H. madagascariensis
Madagascar Extinct (c. AD 1000)  
Common hippopotamus
Bumiayu dwarf hippopotamus[26] 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 Extinct (Pleistocene)
 
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
Example Binomial name Native range Status Continental relative
Sicilian bison[27] Bison priscus siciliae Sicily Extinct (Late Pleistocene)  
Steppe bison
Sicilian aurochs[44] Bos primigenius siciliae[27] Sicily Extinct (Late Pleistocene)  
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 Majorca and Menorca Extinct (after 3000 BC) Gallogoral
Nesogoral[45] Nesogoral spp. Sardinia Extinct
Dahlak Kebir gazelle[46] Nanger soemmerringi ssp. Dahlak Kebir island, Eritrea Vulnerable  
Soemmerring's gazelle
 
Tyrrhenotragus
Tyrrhenotragus gracillimus Baccinello Extinct Antilopinae sp.

Cervids and relatives

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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[49]
Cervus astylodon Ryukyu Islands Extinct  
Sika deer (?)

Cervus praenipponicus (?)
Jersey red deer population[50] Cervus elaphus jerseyensis Jersey Extinct (Pleistocene)  
Red deer
 
Corsican red deer
Cervus elaphus corsicanus Corsica and Sardinia Near Threatened
Sicilian red deer[27] 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 Vulnerable  
Sambar deer
Possible example Binomial name Native range Status Continental relative
 
Insular elephant cacti[51][52]
Pachycereus pringlei Remote islands in the Sea of Cortez
(e.g. Santa Cruz, San Pedro Mártir)
Not evaluated  
Mainland elephant cacti

See also

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Notes

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  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[42]
  6. ^ For mainland gray foxes in general[43]
  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.[47][48]
  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).[47]

References

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  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. Bibcode:1982Pbio....8...16P. doi:10.1017/S0094837300004322. JSTOR 2400564. S2CID 88464305.
  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 3060065. PMID 21436896.
  3. ^ a b c d e Van Den Bergh, Gerrit Dirk; Awe, Rokhus Due; Morwood, Michael John; Sutikna, Thomas; 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. Bibcode:2008QuInt.182...16V. doi:10.1016/j.quaint.2007.02.001.
  4. ^ a b c d 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. S2CID 26853128.
  5. ^ a b 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. S2CID 58524.
  6. ^ a b Evans, A. R.; et al. (2012-01-30). "The maximum rate of mammal evolution". PNAS. 109 (11): 4187–4190. Bibcode:2012PNAS..109.4187E. doi:10.1073/pnas.1120774109. PMC 3306709. PMID 22308461.
  7. ^ a b 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. Bibcode:2001PNAS...9814518B. doi:10.1073/pnas.251548698. ISSN 0027-8424. JSTOR 3057309. PMC 64714. PMID 11724953.
  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". Journal of Biogeography. 43 (8): 1656–1666. doi:10.1111/jbi.12743. ISSN 0305-0270.
  9. ^ "Dwarf dinosaur island really did exist, scientists claim". Telegraph Media Group. 2010-02-22. Archived from the original on 2010-02-25. Retrieved 2010-02-26.
  10. ^ a b c d 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. 293 (3–4): 438–454. Bibcode:2010PPP...293..438B. doi:10.1016/j.palaeo.2010.01.026. Archived from the original (PDF) on 2011-07-10. Retrieved 2017-07-30.
  11. ^ Dyke, G. (2011-09-20). "The Dinosaur Baron of Transylvania". Scientific American. 305 (4): 80–83. Bibcode:2011SciAm.305c..80D. doi:10.1038/scientificamerican1011-80. PMID 22106812.
  12. ^ Company, J. (2010). "Bone histology of the titanosaur Lirainosaurus astibiae (Dinosauria: Sauropoda) from the Latest Cretaceous of Spain". Naturwissenschaften. 98 (1): 67–78. doi:10.1007/s00114-010-0742-3. hdl:10251/148874. PMID 21120450. S2CID 31752413. {{cite journal}}: |last1= has generic name (help)
  13. ^ Carpenter, K. (2001) The Armored Dinosaurs. Indiana University Press, 526 pages.
  14. ^ a b c d 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. Bibcode:2004BiCon..13..317C. CiteSeerX 10.1.1.511.2040. doi:10.1023/b:bioc.0000006501.80472.cc. S2CID 25730672.
  15. ^ Parker S (1984) The extinct Kangaroo Island Emu, a hitherto-unrecognised species. Bulletin of the British Ornithologists' Club 104: 19–22.
  16. ^ a b Heupink, T. H.; Huynen, L.; Lambert, D. M. (2011). "Ancient DNA Suggests Dwarf and 'Giant' Emu Are Conspecific". PLoS ONE. 6 (4): e18728. Bibcode:2011PLoSO...618728H. doi:10.1371/journal.pone.0018728. PMC 3073985. PMID 21494561.
  17. ^ 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.
  18. ^ 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. Bibcode:2012PLoSO...731314G. doi:10.1371/journal.pone.0031314. PMC 3279364. PMID 22348069.
  19. ^ 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.
  20. ^ 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" (PDF). Bulletin of the Chicago Herpetological Society. 47 (1): 1–6. Retrieved 2020-03-02.
  21. ^ a b 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. Bibcode:2002NW.....89..201A. doi:10.1007/s00114-002-0320-4. PMID 12135085. S2CID 4368895.
  22. ^ 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. Bibcode:1998FuEco..12..248S. doi:10.1046/j.1365-2435.1998.00179.x.
  23. ^ 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. Bibcode:2016FuEco..30..244S. doi:10.1111/1365-2435.12472.; see Table S1
  24. ^ Schirber, Michael. Surviving Extinction: Where Woolly Mammoths Endured. Live Science. Imaginova Corporation. Retrieved 2007-07-20.
  25. ^ 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
  26. ^ a b c d e f Aziz, F.; van den Bergh, G. D. (September 25, 1995). "A dwarf Stegodon from Sambungmacan (Central Java, Indonesia)". Proc. Kon. Ned. Akad. V. Wetensch. 98 (3): 229–241. Retrieved 2017-07-31.
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