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Dwarf elephants are pre-historic members of the order Proboscidea, that, through the process of allopatric speciation, evolved to a fraction of the size of their modern relatives. Island dwarfing is a biological phenomenon by which the size of animals isolated on an island shrinks dramatically over generations. Skeletal remains of dwarf elephants have been found on the Mediterranean islands of Cyprus, Malta (at Ghar Dalam), Crete, Sicily, Sardinia, the Cyclades Islands and the Dodecanese Islands. Other islands where dwarf elephants have been found are Sulawesi, Flores, Timor and other islands of the Lesser Sundas, the Channel Islands of California, and Wrangel Island off Siberia.
Mediterranean Islands
Dwarf elephants were once part of the Pleistocene fauna of several Mediterranean islands. Mediterranean dwarf elephants have generally been considered as paleoloxodontine, derived from the continental Straight-tusked Elephant, Elephas (Palaeoloxodon) antiquus Falconer & Cautley, 1847. An exception is the dwarf Sardinian Mammoth, Mammuthus lamarmorae (Major, 1883), the only endemic elephant of the Mediterranean islands belonging to the mammoth line (DNA research published in 2006 shows that Elephas creticus may be from the mammoth line too).
During low sea levels, the Mediterranean islands were colonised again and again, giving rise, sometimes on the same island, to several species (or subspecies) of different body sizes. These endemic dwarf elephants were taxonomically different on each island or group of very close islands, like the Cyclades archipelago.
There is still much uncertainty about the time of colonisation, the phylogenetic relationships, and the taxonomic status of dwarf elephants on the Mediterranean islands.
Sardinia
- Mammuthus lamarmorae (Major, 1883)
Sicily & Malta
- Elephas (Palaeoloxodon) antiquus leonardii Aguirre, 1969
- Elephas (Palaeoloxodon) mnaidriensis (Adams, 1874)
- Elephas (Palaeoloxodon) melitensis Falconer, 1868
- Elephas (Palaeoloxodon) falconeri Busk, 1867
Crete
- Elephas (Palaeoloxodon) creticus Bate, 1907
- Elephas (Palaeoloxodon) creutzburgi (Kuss, 1965)
- Elephas (Palaeoloxodon) chaniensis Symeonides et al., 2001
After DNA research, published in 2006, it has been proposed to rename Elephas (Palaeoloxodon) creticus into Mammuthus creticus (Bate, 1907). Others proposed (in 2002) to rename all the described specimens of larger size under the new subspecies name Elephas antiquus creutzburgi (Kuss, 1965).
Cyprus
- Elephas (Palaeoloxodon) cypriotes Bate, 1903
Cyclades Islands
Remains of paleoloxodontine elephants have been reported from the islands of Delos, Naxos, Kythnos, Serifos and Milos. The Delos elephant is of similar size to a small Elephas antiquus, while the Naxos elephant is of similar size to Elephas melitensis. The remains from Kythnos, Serifos and Milos have not been described.Dodecanese Islands
On the island of Rhodes, bones of an endemic dwarf elephant have been discovered. This elephant was similar in size to Elephas mnaidriensis.Two groups of remains of dwarf elephants have been found on the island of Tilos. They are similar in size to Elephas mnaidriensis and the smaller Elephas falconeri, but according to Theodorou (1983, 1988) the two groups indicate sexual dimorphism. The remains had originally been designated to Palaeoloxodon antiquus falconeri (Busk, 1867). However, this name refers to the dwarf elephants from the island of Malta. As a result, since no migration route between the two islands can be proved, this name should not be used when referring to the elephant remnants from Tilos. Theodorou (1983) has accepted the temporal use of this name until further material can be examined.
The Tilos dwarf elephant is the first dwarf elephant whose DNA sequence has been studied. The results of this research are consistent with previous morphological reports, according to which Palaeoloxodon is more closely related to Elephas than to Loxodonta or Mammuthus.
The Tilos dwarf elephant was the latest paleoloxodontine to survive in Europe. They became extinct just less than 4000 years BP, so this elephant survived well into the Holocene.
Channel Islands of California
The Columbian mammoth (Mammuthus columbii) produced a separate, isolated population at the end of the Pleistocene. One of these isolated groups was formed on the Californian Channel Islands, most likely about 40,000 years ago (although the time of isolation is not fully known). Selective forces on the Channel Islands resulted in smaller animals, forming a new species, Mammuthus exilis. Channel Islands mammoths ranged from 150-180 cm in shoulder height.
Wrangel Island
During the last ice age, woolly mammoths (Mammuthus primigenius) lived on Wrangel Island in the Arctic Ocean. It has been shown that mammoths survived on Wrangel Island until 1700 B.C., the most recent survival of any known mammoth population. Wrangel Island is thought to have become separated from the mainland by 12,000 years BP. Survival of a mammoth population may be explained by local topography and climatic features, which permitted relictual preservation of communities of steppe plants. Because of limited food supply, however, they were much smaller than the typical woolly mammoth. Holocene mammoths from Wrangel Island ranged from 180-230 cm in shoulder height.
Indonesia
On Sulawesi and Flores evidence of a succession of distinct endemic island faunas has been found, including dwarfed elephants, dating until the Middle Pleistocene. Around the early Middle Pleistocene these dwarfed elephants were replaced by new immigrants of larger to intermediate sizes.
Flores
The present understanding of the succession of Stegodon species on Flores is that endemic dwarfs, represented by the Early Pleistocene species Stegodon sondaarii, became extinct around 840,000 years ago. These dwarf forms were then replaced by the medium to large-sized Stegodon florensis, a species closely related to the Stegodon trigonocephalus group found both in Java and in the islands of biogeographical Wallacea, separated by deep water from the Asian and Australian continental shelves.Sulawesi
The dwarfed Stegodon sompoensis lived during the Pleistocene on the island of Sulawesi. They had a shoulder height of only 1.5m.References
- Davies, P., Lister, A.M., 2001. Palaeoloxodon cypriotes, the dwarf elephant of Cyprus: size and scaling comparisons with P. falconeri (Sicily-Malta) and mainland P. antiquus. The World of Elephants, Proceedings of the 1st International Congress (October 16-20 2001, Rome): 479-480. PDF fulltext
- Palombo, M.R. 2001. Endemic elephants of the Mediterranean Islands: knowledge, problems and perspectives. The World of Elephants, Proceedings of the 1st International Congress (October 16-20 2001, Rome): 486-491. PDF fulltext
- Poulakakis, N., Mylonas, M., Lymberakis, P., Fassoulas, C. 2002. Origin and taxonomy of the fossil elephants of the island of Crete (Greece): problems and perspectives. Palaeogeography, Palaeoclimatology, Palaeoecology, 2002, 186(1-2), pp 163-183.
- Poulakakis, N., A. Parmakelis, P. Lymberakis, M. Mylonas, E.Zouros, D. S. Reese, S. Glaberman and A. Caccone. 2006. Ancient DNA forces reconsideration of evolutionary history of Mediterranean pygmy elephantids. Biology Letters, 19 april 2006.
- Poulakakis N, Theodorou GE, Zouros E, Mylonas M. 2002. Molecular phylogeny of the extinct pleistocene dwarf elephant Palaeoloxodon antiquus falconeri from Tilos Island, Dodekanisa, Greece. Journal of Molecular Evolution: 55(3):364-74.
- Symeonides, N.K., Theodorou, G.E., Giannopoulos V.I. 2001. New data on Elephas chaniensis (Vamos cave, Chania, Crete). The World of Elephants, Proceedings of the 1st International Congress (October 16-20 2001, Rome): 510-513. PDF fulltext
- Theodorou, G. 1983. The dwarf elephants of the Charkadio cave on the island of Tilos (Dodekanese, Greece). Phd Thesis Athens University: 321 pp.
- Theodorou, G.E. 1988. Environmental factors affecting the evolution of islands endemics: The Tilos example for Greece. Modern Geology, 13: 183-188.
- Theodorou, G.E., Agiadi, K., 2001. Observations on the microstructure of fossil tusks from the Charkadio cave (Tilos, Dodekanese, Greece). The World of Elephants, Proceedings of the 1st International Congress (October 16-20 2001, Rome): 563-567. PDF fulltext
- Tikhonov, A., Agenbroad, L. & Vartanyan, S., 2003 - Comparative analysis of the mammoth populations on Wrangel Island and the Channel Islands - in: Reumer, J.W.F., De Vos, J. & Mol, D. (eds.) - Advances in Mammoth Research (Proceedings of the Second International Mammoth Conference, Rotterdam, May 16-20 1999) - DEINSEA 9: 415-420 0923-9308 Published 24 May 2003
- Van den Bergh, G.D., De Vos, J., Aziz, F., Morwood, M.J. 2001. Elephantoidea in the Indonesian region: new Stegodon findings from Flores. The World of Elephants, Proceedings of the 1st International Congress (October 16-20 2001, Rome): 623-627. PDF fulltext
- Vartanyan, S.L., Garutt, V.E., Sher, A.V. March 1993. Holocene dwarf mammoths from Wrangel Island in the Siberian Arctic. Nature 362, 337 - 340.
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