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On the island [[Tilos]], the species ''Palaeoloxodon tiliensis'' has been described from remains found in Charkadio cave.<ref>G. Theodorou, N. Symeonidis, E. Stathopoulou ''Elephas tiliensis'' n. sp. from Tilos island (Dodecanese, Greece) Hell. J. Geosci., 42 (2007), pp. 19-32</ref> This species was medium-sized at around {{Convert|1.9|m|ft}} tall. Remains of the species are suggested to date to Late Pleistocene. Radiocarbon dating done in the 1970s suggested that the species survived until around 3,500 years ago, which would make the latest surviving ''Palaeoloxodon'' species and the youngest elephant in Europe, but these dates are tentative and await corroboration by other research.<ref name=":0" /> |
On the island [[Tilos]], the species ''Palaeoloxodon tiliensis'' has been described from remains found in Charkadio cave.<ref>G. Theodorou, N. Symeonidis, E. Stathopoulou ''Elephas tiliensis'' n. sp. from Tilos island (Dodecanese, Greece) Hell. J. Geosci., 42 (2007), pp. 19-32</ref> This species was medium-sized at around {{Convert|1.9|m|ft}} tall. Remains of the species are suggested to date to Late Pleistocene. Radiocarbon dating done in the 1970s suggested that the species survived until around 3,500 years ago, which would make the latest surviving ''Palaeoloxodon'' species and the youngest elephant in Europe, but these dates are tentative and await corroboration by other research.<ref name=":0" /> |
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On the island of [[Astypalaia]], a single tusk of a dwarf elephant of unknown age was excavated in the late 1990s. Due to the isolated status of the island it very likely represents an endemic species. Though the size of the animal is difficult to constrain precisely, it was probably similar in size to ''P. tiliensis''.<ref name=":0" /> |
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⚫ | On [[Kasos]], which during the Pleistocene was connected with the islands of [[Karpathos]] and [[Saria Island|Saria]], a single dwarf ''Palaeoloxodon'' molar has been found |
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⚫ | On [[Kasos]], which during the Pleistocene was connected with the islands of [[Karpathos]] and [[Saria Island|Saria]], a single dwarf ''Palaeoloxodon'' molar has been found. Due to the tooth closely resembling those of the species ''P. creutzburgi'' from Crete (which is adjacent to Kasos) in size and shape, it has been referred to as ''P.'' [[Species affinis|aff.]] ''creutzburgi.''<ref name=":4" /><ref name=":0" /> |
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===Cyprus=== |
===Cyprus=== |
Revision as of 19:49, 25 May 2024
Dwarf elephants are prehistoric members of the order Proboscidea which, through the process of allopatric speciation on islands, evolved much smaller body sizes (around 1–2.3 metres (3 ft 3 in – 7 ft 7 in) shoulder height) in comparison with their immediate ancestors. Dwarf elephants are an example of insular dwarfism, the phenomenon whereby large terrestrial vertebrates (usually mammals) that colonize islands evolve dwarf forms, a phenomenon attributed to adaptation to resource-poor environments and lack of predation and competition.
Fossil remains of dwarf elephants have been found on the Mediterranean islands of Cyprus, Malta, Crete, Sicily, Sardinia, the Cyclades Islands and the Dodecanese Islands, which are mostly members of the genus Palaeoloxodon, descending from the large 4 metres (13 ft) tall mainland European species Palaeoloxodon antiquus, though two species represent dwarf mammoths. Dwarf species of elephants and Stegodon have been found on the islands of Indonesia and the Philippines. The Channel Islands of California once supported a dwarf species descended from Columbian mammoths,[1] while the woolly mammoths that existed on Wrangel Island north of Siberia were once considered dwarfs, but are not anymore.
The Mediterranean islands
Dwarf elephants first inhabited the Mediterranean islands during the Pleistocene, including all the major islands with the apparent exception of Corsica and the Balearics. Mediterranean dwarf elephants have generally been considered as members of the genus Palaeoloxodon, derived from the continental straight-tusked elephant, Palaeoloxodon antiquus (Falconer & Cautley, 1847), Syn.: Elephas antiquus. An exception is the dwarf Middle-Late Pleistocene Sardinian mammoth, Mammuthus lamarmorai (Major, 1883), the first endemic elephant of the Mediterranean islands recognized as belonging to the mammoth line. Mammuthus creticus from the Early Pleistocene of Crete, formerly considered a member of Palaeoloxodon, is now also considered to be a mammoth, and approaches the size of the smallest dwarf elephants.[3]
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. As the Ice Age came to an end, sea levels rose, stranding elephants on the island. The island of Sicily appears to have been colonised by proboscideans in at least three separate waves of colonisation. These endemic dwarf elephants were taxonomically different on each island or group of very close islands, like the Cyclades archipelago.
There are many uncertainties about the time of colonisation, the phylogenetic relationships and the taxonomic status of dwarf elephants on the Mediterranean islands. Extinction of the insular dwarf elephants has not been correlated with the arrival of humans to the islands. Furthermore, it has been suggested by the palaeontologist Othenio Abel in 1914,[4] that the finding of skeletons of such elephants sparked the idea that they belonged to giant one-eyed monsters, because the center nasal opening was thought to be the socket of a single eye, and thus perhaps were, for example, the origin of the one-eyed Cyclopes of Greek mythology.
Italy and Malta
Island | Taxon | Author |
---|---|---|
Sardinia | Mammuthus lamarmorai | (Major, 1883) |
Malta | Palaeoloxodon falconeri | (Busk, 1869) |
Palaeoloxodon mnaidriensis | (Adams, 1874) | |
Sicily | Palaeoloxodon falconeri | (Busk, 1869) |
Palaeoloxodon cf. mnaidriensis | (Adams, 1874) |
Sicily and Malta were inhabited by two successive waves of dwarf elephants derived from P. antiquus, which first arrived on the islands at least 500,000 years ago. The first of these species is P. falconeri , which is one of the smallest dwarf elephant species at around 1 metre (3.3 ft) tall, and was strongly modified from its ancestor in numerous aspects, which lived in a depauperate fauna with no other large mammal species.[6][7] Later, around 200,000 years ago, this species was replaced by a second colonisation by P. antiquus, which gave rise to the larger (though still considerably dwarfed) 2 metres (6.6 ft) tall species P. mnaidriensis, which on Sicily lived alongside a number other large mammal species, including herbivores and carnivores.[7][8][9] The youngest records of this species on Sicily date to around 20,000 years ago, close to the time of arrival of modern humans on Sicily.[10]
The dwarf mammoth species Mammuthus lamarmorai colonised Sardinia sometime after 450,000 years ago. It is suggested to have survived into the Last Glacial Period, until at least 60-30,000 years ago.[11]
Greece
Crete
Island | Taxon | Author |
---|---|---|
Crete | Mammuthus creticus | (Bate, 1907) |
Palaeoloxodon creutzburgi | (Kuss, 1965) |
Mammuthus creticus is the earlier species known from the Early Pleistocene. It is the smallest mammoth[12] and is among the smallest dwarf elephants known, with a shoulder height of about 1 metre (3.3 ft) and a weight of about 180 kilograms (400 lb).[13] Palaeoloxodon creutzburgi from the Middle Pleistocene and Late Pleistocene is significantly larger, with an estimated body mass comparable to living Asian elephant, around 40% the size of its mainland ancestor.[14]
Cyclades
Island | Taxon | Author |
---|---|---|
Delos | Palaeoloxodon sp. | Vaufrey, 1929 |
Naxos | Palaeoloxodon lomolinoi | Van der Geer et al., 2014[15] |
Paros | Elephantidae indet. | Georgalas, 1929 |
Kythnos | Elephantidae indet. | Honea, 1975 |
Milos | Elephantidae indet. | Papp, 1953 |
Serifos | Elephantidae indet. | Papp, 1953 |
Remains of paleoloxodontine elephants have been reported from several Cyclades islands. The Delos elephant's size is comparable to Palaeoloxodon mnaidriensis.[15] The Naxos elephant has been described as Palaeoloxodon lomolinoi and it was smaller than the Delos elephant with a calculated body mass of only 10% of that of its mainland ancestor Palaeoloxodon antiquus.[15]
Dodecanese
Island | Taxon | Author |
---|---|---|
Astypalaia | Palaeoloxodon sp. | Athanassiou et al., 2019[2] |
Kasos | Palaeoloxodon aff. creutzburgi | Sen et al., 2014[16] |
Rhodes | Palaeoloxodon sp. | Symeonides et al., 1974 |
Tilos | Palaeoloxodon tiliensis | (Theodorou et al. 2007)[17] |
On the island of Rhodes, bones of an unnamed endemic dwarf elephant have been discovered in cave deposits on the east coast. This elephant was similar in size to Palaeoloxodon mnaidriensis, around 20% the size of its mainland ancestor. The remains, though temporally poorly constrained, are suggested Late Pleistocene age.[14]
On the island Tilos, the species Palaeoloxodon tiliensis has been described from remains found in Charkadio cave.[18] This species was medium-sized at around 1.9 metres (6.2 ft) tall. Remains of the species are suggested to date to Late Pleistocene. Radiocarbon dating done in the 1970s suggested that the species survived until around 3,500 years ago, which would make the latest surviving Palaeoloxodon species and the youngest elephant in Europe, but these dates are tentative and await corroboration by other research.[14]
On the island of Astypalaia, a single tusk of a dwarf elephant of unknown age was excavated in the late 1990s. Due to the isolated status of the island it very likely represents an endemic species. Though the size of the animal is difficult to constrain precisely, it was probably similar in size to P. tiliensis.[14]
On Kasos, which during the Pleistocene was connected with the islands of Karpathos and Saria, a single dwarf Palaeoloxodon molar has been found. Due to the tooth closely resembling those of the species P. creutzburgi from Crete (which is adjacent to Kasos) in size and shape, it has been referred to as P. aff. creutzburgi.[16][14]
Cyprus
Island | Taxon | Author |
---|---|---|
Cyprus | Palaeoloxodon cypriotes | (Bate, 1903)[19] |
Palaeoloxodon xylophagou | Athanassiou et al., 2015[20] |
The Cyprus dwarf elephant (Palaeoloxodon cypriotes) survived at least until 12,000 years ago, around the time of arrival of modern humans to Cyprus (who may have hunted it), making it one of the latest surviving dwarf elephants. It is also one of the smallest dwarf elephant species, comparable in size to P. falconeri, with an estimated shoulder height of 1 metre (3.3 ft). The species likely evolved from the earlier larger (though still strongly dwarfed) Palaeoloxodon xylophagou known from fossils dating to around 200,000 years ago.[21]
Remains of the species were first discovered and recorded by Dorothea Bate in a cave in the Kyrenia hills of northern Cyprus in 1902 and reported in 1903.[19][22]
The Channel Islands of California
The Columbian mammoth (Mammuthus columbi) produced a separate, isolated population at the end of the Pleistocene on the Channel Islands of California, 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, the pygmy mammoth (Mammuthus exilis).[23] Channel Islands mammoths ranged from 150–190 cm (59–75 in) in shoulder height.
Wrangel Island
During the last ice age, woolly mammoths (Mammuthus primigenius) lived on Wrangel Island in the Arctic Ocean and survived until 1700 BCE, the most recent survival of any known mammoth population. Wrangel Island is thought to have become separated from the mainland by 12000 BCE. It was assumed that Wrangel Island mammoths ranged from 180–230 cm (71–91 in) in shoulder height and were for a time considered "dwarf mammoths".[24] However this classification has been re-evaluated and since the Second International Mammoth Conference in 1999, these mammoths are no longer considered to be true "dwarf mammoths".[25]
Indonesia and the Philippines
In Indonesia and the Philippines, evidence of a succession of distinct endemic island faunas has been found, including dwarfed elephants and species of Stegodon.
Flores
During the late Early Pleistocene, Flores was inhabited by the dwarf species Stegodon sondaarii, around 15% of the size of mainland Stegodon species, which was around 120 cm (3.9 ft) tall at the shoulder and weighed about 350–400 kilograms (770–880 lb). This species became extinct around 1 million years ago,[13][26][27] being replaced by Stegodon florensis. Stegodon florensis shows a progressive size reduction with time, with the earlier Middle Pleistocene subspecies Stegodon florensis florensis estimated to be around 50% the size of mainland Stegodon species with a shoulder height of around 190 cm (6.2 ft) and a body mass of around 1.7 tons, while the later Stegodon florensis insularis from the Late Pleistocene is estimated to be around 17% the size of mainland Stegodon species, with a shoulder height of around 130 cm (4.3 ft), and a body mass of about 570 kilograms (1,260 lb) [27][28] Stegodon florensis became extinct about 50,000 years ago, around the time of the arrival of modern humans to Flores.[26]
Sulawesi
During the Late Pliocene-Early Pleistocene on Sulawesi, two species of dwarf proboscidean coinhabited the island, the elephant Stegoloxodon celebensis, and Stegodon sompoensis.[29] The former was about 150 cm (4.9 ft) tall,[30] while the latter was around 32% the size of mainland Stegodon species, with an estimated body mass of about a ton.[27] Later in the Pleistocene, these animals were replaced by larger-sized species of Stegodon and elephants,[29][31] with an indeterminate Stegodon species from the Middle Pleistocene of Sulawesi being around 57% the size of mainland species, with an estimated bodymass of about 2 tons.[27]
Java
The species Stegodon trigonocephalus is known from the Early-Middle Pleistocene of Java.[32] A population from the Trinil H.K locality, which likely dates to the Middle Pleistocene,[33] is around 65% the size of mainland Stegodon species.[27] Large individuals are estimated to have reached around 280 cm (9.2 ft) at the shoulders, with a body mass of around 5 tons.[13] Other smaller unnamed Stegodon species are also known from the Early Pleistocene on the island.[32] The extinct dwarf elephant species Stegoloxodon indonesicus is also known from the Early Pleistocene of Java, which is probably closely related to S. celebensis from Sulawesi, but whose relationships to other elephants are obscure.[34][32]
Sumba
The species Stegodon sumbaensis of an uncertain Middle-Late Pleistocene age from Sumba is one the smallest known species, at around 8% of the size of its mainland ancestor, with an estimated body mass of around 250 kilograms (550 lb).[27]
Timor
The species Stegodon timorensis is known from the Middle Pleistocene of Timor. It is a small-sized species, only slightly larger than S. sondaarii,[35] and around 23% the size of mainland species, with an estimated body mass of around 770 kilograms (1,700 lb).[27]
Luzon
On Luzon the dwarf Stegodon luzonensis is known from remains found in the Manila Basin of an uncertain Pleistocene age,[36] as well as remains found near the early Middle Pleistocene Nesorhinus butchery site.[37] It is around 40% the size of mainland Stegodon species, with a body mass of around 1.3 tons.[27] The extinct dwarf elephant Elephas beyeri is also known from the island,[27] which is estimated to have been about 1.2 metres (3.9 ft) in shoulder height.[38]
Mindanao
On the island of Mindanao, the dwarf Stegodon species Stegodon mindanensis was present at some point in the Pleistocene. It has an estimated body mass of around 400 kilograms (880 lb).[39]
See also
References
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