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Insular dwarfism

From Wikipedia, the free encyclopedia

Skeletons of Malta's extinct Palaeoloxodon falconeri, the smallest known species of elephant. Adult males measured about one meter in shoulder height and weighed about 305 kg. Females were smaller.
Skeletons of Malta's extinct Palaeoloxodon falconeri, the smallest known species of elephant. Adult males measured about one meter in shoulder height and weighed about 305 kg. 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[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 dinosaurs, like Europasaurus, and modern animals such as elephants and their relatives. 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.

Possible causes

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]

Dwarfism vs. 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]


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.[8][9] 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.[10]


Example Species Range Time frame Continental relative

A. atacis Ibero-Armorican Island Late Cretaceous / Maastrichtian
Tapuiasaurus NT.jpg

Europasaurus skull.JPG

E. holgeri Lower Saxony Late Jurassic / Middle Kimmeridgian
Giraffatitan scale.png

Magyarosaurus- human size.JPG

M. dacus Hateg Island Late Cretaceous / Maastrichtian
Rapetosaurus BW.jpg


L. astibiae Ibero-Armorican Island Late Cretaceous
Paludititan nalatzsensis.jpg

P. nalatzensis Hateg Island Late Cretaceous / Maastrichtian
Epachtosaurus sciuttoi.jpg



Example Species Range Time frame Continental relative
Langenburg theropod size.png

Langenberg Quarry
torvosaur (blue)
Unnamed Lower Saxony Late Jurassic / Middle Kimmeridgian
Torvosaurus gurneyi.png

Struthiosaurus austriacus.jpg

S. austriacus

S. transylvanicus

S. languedocensis
Ibero-Armorican, Australoalpine, and Hateg islands Late Cretaceous
Edmontonia Scale.svg

Telmatosaurus Scale.svg

T. transsylvanicus Hateg Island Late Cretaceous
Hadrosaurus Scale.svg

Tethyshadros Scale.svg

T. insularis Trieste province Late Cretaceous
Thecodontosaurus Scale.svg

T. antiquus Southern England Late Triassic / Rhaetian
Human-plateosaurus size comparison.svg

Iguanodontian Sizes.svg

Zalmoxes[9] (purple)
Z. robustus

Z. shqiperorum
Hateg Island Late Cretaceous
Perot Museum Tenontosaurus.jpg


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).


Example Binomial name Native range Status Continental relative Insular / mainland
length or mass ratio
Apteribis sp. (5212794163).jpg

Hawaiian flightless ibises
Apteribis glenos Molokai Extinct (Late Quaternary)
White Ibis in Florida.jpg

American ibises
Apteribis brevis Maui
Cozumel curassow[13] Crax rubra griscomi Cozumel Unknown
Crax rubra (Great Curassow) - female.jpg

Great curassow
Baudin emus.jpg

Kangaroo Island emu[14]
Dromaius novaehollandiae baudinianus Kangaroo Island, South Australia Extinct (c. AD 1827)
Emu RWD1.jpg

Emu size.png

King Island emu[15] (black)
Dromaius novaehollandiae minor King Island, Tasmania Extinct (AD 1822) LR ≈ 0.48 [b]
Naturalis Biodiversity Center - RMNH.AVES.128765 2 - Toxostoma guttatum (Ridgway, 1885) - Mimidae - bird skin specimen.jpeg

Cozumel thrasher[13]
Toxostoma gluttatum Cozumel Critically endangered
Curve-billed Thrasher.jpg

Other thrashers


Example Binomial name Native range Status Continental relative Insular / mainland
length or mass ratio
20150510-IMG 0786.jpg

Madagascar dwarf chameleon
Brookesia minima Nosy Be island, Madagascar Endangered
Brookesia species male female (Journal.pone.0031314.g010).png

Madagascar leaf chameleons
Brookesia micra on a match head.jpg

Nosy Hara chameleon[16]
Brookesia micra Nosy Hara island, Madagascar Vulnerable
Roxby Island tiger snake[5] Notechis scutatus Roxby Island, South Australia Unknown
Tiger snake 2.jpg

Tiger snake
Dwarf Burmese python Python bivittatus progschai Java, Bali, Sumbawa and Sulawesi, Indonesia Unknown
Burmese python (6887388927).jpg

Burmese python
LR ≈ 0.44 [c]
Tanahjampea reticulated python[19] Python reticulatus jampeanus Tanahjampea, between Sulawesi and Flores Unknown
Python reticulatus сетчатый питон-2.jpg

Reticulated python
LR ≈ 0.41, males
LR ≈ 0.49, females [d]



Example Binomial name Native range Status Continental relative
Bradypus pygmaeus.jpg

Pygmy three-toed sloth
Bradypus pygmaeus Isla Escudo de Veraguas, Panama Critically endangered
Bradypus variegatus, the Brown-throated Three-toed Sloth (12687597105).jpg

Brown-throated sloth

A. antillensis

A. odontrigonus

A. ye
Cuba, Hispaniola and Puerto Rico Extinct (c. 3000 BC)
Megalonyx size.svg

Continental ground sloths
Imagocnus I. zazae Cuba Extinct (Early Miocene)

M. rodens

M. zile
Cuba and Hispaniola Extinct (c. 2700 BC)
Synocnus comes.jpg

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

Cretan dwarf mammoth
Mammuthus creticus Crete Extinct
Mammuthus Scale.svg

M. exilis skeletal.png

Channel Islands mammoth
Mammuthus exilis Santa Rosae island Extinct (Late Pleistocene)
M. columbi skeletals.png

Columbian mammoth
Sardinian mammoth Mammuthus lamarmorai Sardinia Extinct (Late Pleistocene)
Steppe mammoth size 2.jpg

Steppe mammoth
Saint Paul Island woolly mammoth[22][23] Mammuthus primigenius Saint Paul Island, Alaska Extinct (c. 3750 BC)
M. primigenius.png

Woolly mammoth
Elephas skeleton.JPG

Siculo-Maltese elephants
Palaeoloxodon antiquus leonardi

P. mnaidriensis

P. melitensis

P. falconeri
Sicily and Malta Extinct

Straight-tusked elephant
Cretan elephants Palaeoloxodon chaniensis

P. creutzburgi
Crete Extinct
Elephas cypriotes Tusk and Molar.jpg

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[24] Sinomastodon bumiajuensis Bumiayu Island (now part of Java) Extinct (Early Pleistocene)

のんほいパーク - アケボノゾウ.jpg

Japanese stegodont[25]
Stegodon aurorae Japan and Taiwan[26] Extinct (Early Pleistocene)
Stegodon skeletal.png

Chinese Stegodon
Greater Flores dwarf stegodont[3] Stegodon florensis Flores Extinct (Late Pleistocene)
Stegodon’s ivory displayed at Philippine National Museum.jpg

Sundaland Stegodon
Javan dwarf stegodonts Stegodon hypsilophus[24]

S. semedoensis[27]

S. sp.[24]
Java Extinct (Quaternary)
Mindanao pygmy stegodont[28] Stegodon mindanensis Mindanao and Sulawesi Extinct (Middle Pleistocene)
Sulawesi dwarf stegodont[24] Stegodon sompoensis Sulawesi Extinct
Lesser Flores dwarf stegodont[3] Stegodon sondaari Flores Extinct (Middle Pleistocene)
Sumba dwarf stegodont[29] Stegodon sumbaensis Sumba, Indonesia Extinct (Middle Pleistocene)
Timor dwarf stegodont[24] Stegodon timorensis Timor Extinct
Dwarf stegolophodont[30] Stegolophodon pseudolatidens Japan Extinct (Miocene)
Stegolophodon latidens.JPG



Example Binomial name Native range Status Continental relative
Nosy Hara dwarf lemur[31] Cheirogaleus sp. Nosy Hara island off Madagascar Unknown

Dwarf lemurs
Specimen LB1.jpg

Flores Man[32]
Homo floresiensis Flores Extinct (Late Pleistocene)
Homme de Tautavel 01-08.jpg

Homo erectus

Callao Man
Homo luzonensis[33][34] Luzon, Philippines Extinct (Late Pleistocene)
Modern pygmies of Flores[35] Homo sapiens Flores Extant
Australia Aboriginal Culture 001 (5444690703).jpg

Homo sapiens
Early Palau modern humans (disputed)[36] Homo sapiens Palau Extinct (?)
Great Andamanese - two men - 1875.jpg

Homo sapiens Andaman Islands Extant
Macaca majori.JPG

Sardinian macaque[39]
Macaca majori Sardinia Extinct (Pleistocene)
Barbary macaques of Gibraltar in search of food.jpg

Barbary macaque
Red Colobus 7.jpg

Zanzibar red colobus
Piliocolobus kirkii Unguja Endangered
Udzungwa Red Colobus Stevage.JPG

Udzungwa red colobus


Example Binomial name Native range Status Continental relative Insular / mainland
length or mass ratio

Japanese wolf
Canis lupus hodophilax Japan (excluding Hokkaido) Extinct (AD 1905)
Canis lupus Europe.jpg

Gray wolf
Adaptations of the Pleistocene island canid Cynotherium sardous (2006) Fig. 1.png

Sardinian dhole
Cynotherium sardous Corsica and Sardinia Extinct (c. 8300 BC)
Xenocyon lycanoides restoration.jpg

Trinil dog Mececyon trinilensis Java Extinct (Pleistocene)
Cozumel Island coati[13] Nasua narica nelsoni Cozumel Critically endangered
White nosed Coati.jpg

Yucatan white-nosed coati
Zanzibar Leopard 2.JPG

Zanzibar leopard
Panthera pardus pardus Unguja Critically endangered or Extinct
Male leopard samburu 2, crop.jpg

African leopard
Bali tiger zanveld.jpg

Bali tiger
Panthera tigris sondaica Bali Extinct (c. AD 1940)
Panthera tigris sumatrae (Sumatran Tiger) close-up.jpg

Sumatran tiger
Panthera tigris sondaica 01.jpg

Javan tiger
Java Extinct (c. AD 1975)
Cozumel Raccoon2.jpg

Cozumel raccoon
Procyon pygmaeus Cozumel Critically endangered

Common raccoon
Urocyon littoralis pair.jpg

Island fox
Urocyon littoralis Six of the Channel Islands of California Near Threatened
Grey Fox (Urocyon cinereoargenteus).jpg

Gray fox
LR ≈ 0.84 [e]
LR ≈ 0.75 [f]
Cozumel fox Urocyon sp. Cozumel Critically endangered or Extinct

Non-ruminant ungulates

Example Binomial name Native range Status Continental relative
Hippo1 final.jpg

Malagasy dwarf hippopotamuses
Hippopotamus laloumena

H. lemerlei

H. madagascariensis
Madagascar Extinct (c. AD 1000)

Common hippopotamus
Bumiayu dwarf hippopotamus[24] Hexaprotodon simplex Bumiayu Island (now Java) Extinct (Early Pleistocene)
Hexaprotodon sivalensis.jpg

Asian hippopotamuses
Hippopotamus cruetzburgi.JPG

Cretan dwarf hippopotamus
Hippopotamus creutzburgi Crete Extinct (Middle Pleistocene)
Museo di paleologia, scheletro di hippopotamus antiquus, recuperato presso figline valdarno.JPG

European hippopotamus
Hippopotamus amphibius Linn at Ghar Dalam, Malta.png

Maltese dwarf hippopotamus
Hippopotamus melitensis Malta Extinct (Pleistocene)

Cyprus dwarf hippopotamus
Hippopotamus minor Cyprus Extinct (c. 8000 BC)
Hippopotamus pentlandi 3.JPG

Sicilian dwarf hippopotamus
Hippopotamus pentlandi Sicily Extinct (Pleistocene)
Cozumel collared peccary[13] Pecari tajacu nanus Cozumel Unknown
Running Javelina.jpg

Collared peccary
Philippine rhinoceros[42] Rhinoceros philippinensis Luzon Extinct (Middle Pleistocene)
Javan Rhino 1900.jpg

Javan rhinoceros


Example Binomial name Native range Status Continental relative
Sicilian bison[25] Bison priscus siciliae Sicily Extinct (Late Pleistocene)
Prazubr rysunek 600.jpg

Steppe bison
Sicilian aurochs[43] Bos primigenius siciliae[25] Sicily Extinct (Late Pleistocene)
Aurochs reconstruction.jpg

Eurasian aurochs
Cebu tamaraw Bubalus cebuensis Cebu, Philippines Extinct
Indian Water Buffalo Bubalus arnee by Dr Raju Kasambe IMG 0347 (11) (cropped).jpg

Wild water buffalo
Lowland anoa.jpg

Lowland anoa
Bubalus depressicornis Sulawesi and Buton, Indonesia Endangered
Bubalus mindorensis by Gregg Yan 01.jpg

Bubalus mindorensis Mindoro, Philippines Critically endangered
Buablus quarlesi2.jpg

Mountain anoa
Bubalus quarlesi Sulawesi and Buton, Indonesia Endangered
Myotragus balearicus.JPG

Balearic Islands cave goat
Myotragus balearicus Majorca and Menorca Extinct (after 3000 BC) Gallogoral
Nesogoral[44] Nesogoral spp. Sardinia Extinct
Dahlak Kebir gazelle[45] Nanger soemmerringi ssp. Dahlak Kebir island, Eritrea Vulnerable
The book of antelopes (1894) Gazella soemmerringi (white background).png

Soemmerring's gazelle

Cervids and relatives

Example Binomial name Native range Status Continental relative
Candiacervus ropalophorus.jpg

Cretan dwarf megacerines[g]
Candiacervus spp. Crete Extinct (Pleistocene)
Praemegaceros obscurus.JPG

Praemegaceros verticornis[9]
Megaloceros Species.jpg

Sardinian megacerine[9]
(second from left)
Praemegaceros cazioti Sardinia Extinct (c. 5500 BC)
Ryukyu dwarf deer[48] Cervus astylodon Ryukyu Islands Extinct
The deer of all lands (1898) Manchurian sika white background.png

Sika deer (?)

Cervus praenipponicus (?)
Jersey red deer population[49] Cervus elaphus jerseyensis Jersey Extinct (Pleistocene)

Red deer

Corsican red deer
Cervus elaphus corsicanus Corsica and Sardinia Near Threatened
Pleistocene Sicilian deer[25] Cervus siciliae Sicily Extinct (Late Pleistocene)
Hoplitomeryx matthei.jpg

Hoplitomeryx spp. Gargano Island Extinct (Early Pliocene)
Antilocapra americana male (Wyoming, 2012).jpg

Sicilian megacerine[25] Megaloceros carburangelensis Sicily Extinct (Late Pleistocene)
Mégalocéros B&W rogné.png

Irish elk
Key deer male.jpg

Florida Key deer
Odocoileus virginianus clavium Florida Keys Endangered
White-tailed deer.jpg

Virginia deer
Spitsbergen reindeer01.jpg

Svalbard reindeer
Rangifer tarandus platyrhynchus Svalbard Unknown

Rusa marianna by Gregg Yan.jpg

Philippine deer
Rusa marianna Philippines Vulnerable
Sambar (Rusa unicolor cambojensis) (7109798353).jpg

Sambar deer


Possible example Binomial name Native range Status Continental relative
El Tecolote.JPG

Insular elephant cacti[50][51]
Pachycereus pringlei Remote islands in the Sea of Cortez
(e.g. Santa Cruz, San Pedro Mártir)
Not evaluated
Pachycereus pringlei cardon sahueso.JPG

Mainland elephant cacti

See also


  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[15]
  3. ^ Based on maximum lengths of 2.5 m for the dwarf form[17] and 5.74 m for the mainland form[18]
  4. ^ Based on maximum Tanahjampea python total lengths (TL) of 2.10 m for males and 3.35 m for females[19] and maximum southern Sumatra python snout to vent lengths (SVL) of 4.5 m for males and 6.1 m for females[20] 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[21]
  5. ^ For nearby mainland gray foxes[40]
  6. ^ For mainland gray foxes in general[41]
  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.[46][47]
  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).[46]


  1. ^ Prothero, D. R.; Sereno, P. C. (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. JSTOR 2400564.
  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, G. D.; Rokhus Due Awe; Morwood, M. J.; Sutikna, T.; 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, P.; Meiri, S. (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. Retrieved 2011-02-11.
  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. ^ "Dwarf dinosaur island really did exist, scientists claim". Telegraph Media Group. 2010-02-22. Retrieved 2010-02-26.
  9. ^ a b c d e 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.
  10. ^ 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.
  11. ^ 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.
  12. ^ Carpenter, K. (2001) The Armored Dinosaurs. Indiana University Press, 526 pages.
  13. ^ 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. CiteSeerX doi:10.1023/ S2CID 25730672.
  14. ^ Parker S (1984) The extinct Kangaroo Island Emu, a hitherto-unrecognised species. Bulletin of the British Ornithologists' Club 104: 19–22.
  15. ^ 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.
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