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'''Dwarf elephants''' are prehistoric members of the order [[Proboscidea]] which, through the process of [[allopatric speciation]] on islands, evolved much smaller body sizes (around {{convert|1|-|2.3|m}} 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.
'''Dwarf elephants''' are prehistoric members of the order [[Proboscidea]] which, through the process of [[allopatric speciation]] on islands, evolved much smaller body sizes (around {{convert|1|-|2.3|m}} 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|Cyclades Islands]] and the [[Dodecanese Islands]], which are mostly members of the genus ''[[Palaeoloxodon]]'', descending from the large {{Convert|4|m|ft}} tall mainland European species ''[[Palaeoloxodon antiquus]].'' Dwarf species of elephants and ''[[Stegodon]]'' have been found on the islands of Indonesia and the Philippines. The [[Channel Islands (California)|Channel Islands of California]] once supported [[Mammuthus exilis|a dwarf species]] descended from [[Columbian mammoth]]s,<ref>{{cite web | url = https://www.nps.gov/chis/learn/historyculture/pygmymammoth.htm | title = The Pygmy Mammoth (U.S. National Park Service) }}</ref> while the [[woolly mammoth]]s that existed on [[Wrangel Island]] north of [[Siberia]] were once considered dwarfs, but are not anymore.
[[Fossil]] remains of dwarf elephants have been found on the [[Mediterranean]] islands of [[Cyprus]], [[Malta]], [[Crete]]. [[Sicily]], [[Sardinia]], the [[Cyclades|Cyclades Islands]] and the [[Dodecanese Islands]], which are mostly members of the genus ''[[Palaeoloxodon]]'', descending from the large {{Convert|4|m|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 (California)|Channel Islands of California]] once supported [[Mammuthus exilis|a dwarf species]] descended from [[Columbian mammoth]]s,<ref>{{cite web | url = https://www.nps.gov/chis/learn/historyculture/pygmymammoth.htm | title = The Pygmy Mammoth (U.S. National Park Service) }}</ref> while the [[woolly mammoth]]s that existed on [[Wrangel Island]] north of [[Siberia]] were once considered dwarfs, but are not anymore.


==The Mediterranean islands==
==The Mediterranean islands==
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| ''[[Palaeoloxodon mnaidriensis|Palaeoloxodon cf. mnaidriensis]]'' || ({{Smallcaps|Adams}}, 1874)
| ''[[Palaeoloxodon mnaidriensis|Palaeoloxodon cf. mnaidriensis]]'' || ({{Smallcaps|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 ''[[Palaeoloxodon falconeri|P. falconeri]] ,'' which is one of the smallest dwarf elephant species at around {{Convert|1|m|ft}} tall, and was strongly modified from its ancestor in numerous aspects, which lived in a depauperate fauna with no other large mammal species.''<ref name=":02">Bonfiglio, L., Marra, A. C., Masini, F., Pavia, M., & Petruso, D. (2002). [https://iris.unito.it/bitstream/2318/91517/1/18%20-%20Bonfiglio%20et%20al%202002%20Sicily%20A%20Review%20BAR.pdf Pleistocene faunas of Sicily: a review]. In W. H. Waldren, & J. A. Ensenyat (Eds.), ''World islands in prehistory: international insular investigations''. British Archaeological Reports, International Series, 1095, 428–436.</ref>''<ref name=":12">{{Cite journal |last=Scarborough |first=Matthew Edward |date=March 2022 |title=Extreme Body Size Variation in Pleistocene Dwarf Elephants from the Siculo-Maltese Palaeoarchipelago: Disentangling the Causes in Time and Space |journal=Quaternary |language=en |volume=5 |issue=1 |pages=17 |doi=10.3390/quat5010017 |issn=2571-550X |doi-access=free |hdl-access=free |hdl=11427/36354}}</ref> 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) {{Convert|2|m|ft}} tall species ''[[Palaeoloxodon mnaidriensis|P. mnaidriensis]],'' which on Sicily lived alongside a number other large mammal species, including herbivores and carnivores.<ref name=":12" /><ref name=":3">{{Cite journal |last1=Baleka |first1=Sina |last2=Herridge |first2=Victoria L. |last3=Catalano |first3=Giulio |last4=Lister |first4=Adrian M. |last5=Dickinson |first5=Marc R. |last6=Di Patti |first6=Carolina |last7=Barlow |first7=Axel |last8=Penkman |first8=Kirsty E.H. |last9=Hofreiter |first9=Michael |last10=Paijmans |first10=Johanna L.A. |date=August 2021 |title=Estimating the dwarfing rate of an extinct Sicilian elephant |url=https://linkinghub.elsevier.com/retrieve/pii/S0960982221007351 |journal=Current Biology |language=en |volume=31 |issue=16 |pages=3606–3612.e7 |doi=10.1016/j.cub.2021.05.037 |pmid=34146486 |s2cid=235477150 |doi-access=free}}</ref><ref name=":122">{{Cite journal |last1=Bethune |first1=Elehna |last2=Kaiser |first2=Thomas M. |last3=Schulz-Kornas |first3=Ellen |last4=Winkler |first4=Daniela E. |date=November 2019 |title=Multiproxy dietary trait reconstruction in Pleistocene Hippopotamidae from the Mediterranean islands |url=https://zenodo.org/record/3248597 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=533 |pages=109210 |bibcode=2019PPP...533j9210B |doi=10.1016/j.palaeo.2019.05.032 |s2cid=181824675}}</ref> 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.<ref name=":2">{{Cite journal |last1=Palombo |first1=Maria Rita |last2=Antonioli |first2=Fabrizio |last3=Di Patti |first3=Carolina |last4=Valeria |first4=Lo Presti |last5=Scarborough |first5=Matthew E. |date=2021-10-03 |title=Was the dwarfed Palaeoloxodon from Favignana Island the last endemic Pleistocene elephant from the western Mediterranean islands? |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2020.1772251 |journal=Historical Biology |language=en |volume=33 |issue=10 |pages=2116–2134 |doi=10.1080/08912963.2020.1772251 |issn=0891-2963 |s2cid=225710152}}</ref>{{legend|lightblue|connected during [[Last Glacial Maximum|LGM]]<ref name="Palombo2020">{{cite journal | doi = 10.1080/08912963.2020.1772251| title = Was the dwarfed ''Palaeoloxodon'' from Favignana Island the last endemic Pleistocene elephant from the western Mediterranean islands?| journal = Historical Biology| pages = 1–19 | year = 2020| last1 = Palombo | first1 = M. R.| last2 = Antonioli | first2 = F.| last3 = Di Patti | first3 = C.| last4 = Presti | first4 = V. L.| last5 = Scarborough | first5 = M. E.| volume = 33| issue = 10| s2cid = 225710152}}</ref>}}

{{legend|lightblue|connected during [[Last Glacial Maximum|LGM]]<ref name="Palombo2020">{{cite journal | doi = 10.1080/08912963.2020.1772251| title = Was the dwarfed ''Palaeoloxodon'' from Favignana Island the last endemic Pleistocene elephant from the western Mediterranean islands?| journal = Historical Biology| pages = 1–19 | year = 2020| last1 = Palombo | first1 = M. R.| last2 = Antonioli | first2 = F.| last3 = Di Patti | first3 = C.| last4 = Presti | first4 = V. L.| last5 = Scarborough | first5 = M. E.| volume = 33| issue = 10| s2cid = 225710152}}</ref>}}


===Greece===
===Greece===
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Poulakakis and others proposed in 2002 to rename all the described specimens of larger size than ''Mammuthus creticus'' under the new subspecies name ''Elephas antiquus creutzburgi'' (Kuss, 1965).<ref>{{cite journal|author1=Poulakakis, N. |author2=Mylonas, M. |author3=Lymberakis, P. |author4=and Fassoulas, C. |date=2002-10-01| title = Origin and taxonomy of the fossil elephants of the island of Crete (Greece): problems and perspectives | journal = Palaeogeography, Palaeoclimatology, Palaeoecology | volume = 186 | issue = 1–2| pages = 163–183 | doi = 10.1016/S0031-0182(02)00451-0}}</ref> After DNA research, published in 2006, it has been proposed to rename ''Elephas (Palaeoloxodon) creticus'' into ''Mammuthus creticus'' (Bate, 1907). In a recent study of 2007, it was argued for the groundlessness of the theory by Poulakakis et al. in 2006,<ref name=Poulakakis2006>{{cite journal |author1=Poulakakis N. |author2=Parmakelis A. |author3=Lymberakis P. |author4=Mylonas M. |author5=Zouros E. |author6=Reese D. |author7=Glaberman S. |author8=Caccone A. | year = 2006 | title = Ancient DNA forces reconsideration of evolutionary history of Mediterranean pygmy elephantids | journal = Biol. Lett. | volume = 2 | issue = 3| pages = 451–454 | doi = 10.1098/rsbl.2006.0467 | pmid = 17148428 | pmc = 1686204 }}</ref> showing the weak points of that DNA research.<ref name="BioLetters">{{cite journal |author1=Orlando, L. |author2=Pagés, M. |author3=Calvignac, S. |display-authors=etal |date=2007-02-22 |title=Does the 43bp sequence from an 800000 year old Cretan dwarf elephantid really rewrite the textbook on mammoths? |journal=[[Biology Letters]] |volume=3 |issue=1 |pages=57–59 |doi=10.1098/rsbl.2006.0536 |pmc=2373798 |pmid=17443966}}</ref> However, morphological data is at least equivocal, and may also support placement in ''Mammuthus''.
Poulakakis and others proposed in 2002 to rename all the described specimens of larger size than ''Mammuthus creticus'' under the new subspecies name ''Elephas antiquus creutzburgi'' (Kuss, 1965).<ref>{{cite journal|author1=Poulakakis, N. |author2=Mylonas, M. |author3=Lymberakis, P. |author4=and Fassoulas, C. |date=2002-10-01| title = Origin and taxonomy of the fossil elephants of the island of Crete (Greece): problems and perspectives | journal = Palaeogeography, Palaeoclimatology, Palaeoecology | volume = 186 | issue = 1–2| pages = 163–183 | doi = 10.1016/S0031-0182(02)00451-0}}</ref> After DNA research, published in 2006, it has been proposed to rename ''Elephas (Palaeoloxodon) creticus'' into ''Mammuthus creticus'' (Bate, 1907). In a recent study of 2007, it was argued for the groundlessness of the theory by Poulakakis et al. in 2006,<ref name=Poulakakis2006>{{cite journal |author1=Poulakakis N. |author2=Parmakelis A. |author3=Lymberakis P. |author4=Mylonas M. |author5=Zouros E. |author6=Reese D. |author7=Glaberman S. |author8=Caccone A. | year = 2006 | title = Ancient DNA forces reconsideration of evolutionary history of Mediterranean pygmy elephantids | journal = Biol. Lett. | volume = 2 | issue = 3| pages = 451–454 | doi = 10.1098/rsbl.2006.0467 | pmid = 17148428 | pmc = 1686204 }}</ref> showing the weak points of that DNA research.<ref name="BioLetters">{{cite journal |author1=Orlando, L. |author2=Pagés, M. |author3=Calvignac, S. |display-authors=etal |date=2007-02-22 |title=Does the 43bp sequence from an 800000 year old Cretan dwarf elephantid really rewrite the textbook on mammoths? |journal=[[Biology Letters]] |volume=3 |issue=1 |pages=57–59 |doi=10.1098/rsbl.2006.0536 |pmc=2373798 |pmid=17443966}}</ref> However, morphological data is at least equivocal, and may also support placement in ''Mammuthus''.


''[[Mammuthus creticus]]'' is the earlier species known from the [[Early Pleistocene]]. It is the smallest mammoth<ref name=":1">{{Cite journal |last1=Herridge |first1=V. L. |last2=Lister |first2=A. M. |year=2012 |title=Extreme insular dwarfism evolved in a mammoth |journal=Proceedings of the Royal Society B: Biological Sciences |volume=279 |issue=1741 |pages=3193–3300 |doi=10.1098/rspb.2012.0671 |pmc=3385739 |pmid=22572206}}</ref> and is among the smallest dwarf elephants known, with a shoulder height of about {{Convert|1|m|ft}} and a weight of about {{Convert|180|kg|lb}}.<ref name="proboscideans" /> ''[[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.<ref>{{Cite journal |last1=Athanassiou |first1=Athanassios |last2=van der Geer |first2=Alexandra A.E. |last3=Lyras |first3=George A. |date=August 2019 |title=Pleistocene insular Proboscidea of the Eastern Mediterranean: A review and update |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119300848 |journal=Quaternary Science Reviews |language=en |volume=218 |pages=306–321 |bibcode=2019QSRv..218..306A |doi=10.1016/j.quascirev.2019.06.028 |s2cid=199107354}}</ref><!-- http://rspb.royalsocietypublishing.org/content/early/2012/05/04/rspb.2012.0671 -->
''[[Mammuthus creticus]]'' is the earlier species known from the [[Early Pleistocene]]. It is the smallest mammoth<ref name=":1">{{Cite journal |last1=Herridge |first1=V. L. |last2=Lister |first2=A. M. |year=2012 |title=Extreme insular dwarfism evolved in a mammoth |journal=Proceedings of the Royal Society B: Biological Sciences |volume=279 |issue=1741 |pages=3193–3300 |doi=10.1098/rspb.2012.0671 |pmc=3385739 |pmid=22572206}}</ref> and is among the smallest dwarf elephants known, with a shoulder height of about {{Convert|1|m|ft}} and a weight of about {{Convert|180|kg|lb}}.<ref name="proboscideans" /> ''[[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.<ref name=":0">{{Cite journal |last1=Athanassiou |first1=Athanassios |last2=van der Geer |first2=Alexandra A.E. |last3=Lyras |first3=George A. |date=August 2019 |title=Pleistocene insular Proboscidea of the Eastern Mediterranean: A review and update |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119300848 |journal=Quaternary Science Reviews |language=en |volume=218 |pages=306–321 |bibcode=2019QSRv..218..306A |doi=10.1016/j.quascirev.2019.06.028 |s2cid=199107354}}</ref><!-- http://rspb.royalsocietypublishing.org/content/early/2012/05/04/rspb.2012.0671 -->


====Cyclades====
====Cyclades====
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On the island of [[Rhodes]], bones of an endemic dwarf elephant have been discovered. This elephant was similar in size to ''Palaeoloxodon mnaidriensis''.
On the island of [[Rhodes]], bones of an endemic dwarf elephant have been discovered. This elephant was similar in size to ''Palaeoloxodon mnaidriensis''.<ref name=":0" />


Two groups of remains of dwarf elephants have been found on the island of [[Tilos]]. They are similar in size to ''Palaeoloxodon mnaidriensis'' and the smaller ''Palaeoloxodon falconeri'', but the two groups indicate sexual dimorphism.<ref name=Theodorou1983>{{cite book | author = Theodorou, G. | year = 1983 | title = The dwarf elephants of the Charkadio cave on the island of Tilos (Dodekanese, Greece) | publisher = Phd Thesis [[Athens University]] | page = 321 pp}}</ref><ref name=Theodorou1988>{{cite journal | author = Theodorou, G.E. | year = 1988 | title = Environmental factors affecting the evolution of islands endemics: The Tilos example for Greece | journal = Modern Geology | volume = 13 | pages = 183–188}}</ref> 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 proven, this name should not be used when referring to the elephant remnants from Tilos. The species has been described as ''Elephas tiliensis'' in 2007<ref name=Theodorou2007/> and it is now assigned to genus ''Palaeoloxodon''.
Two groups of remains of dwarf elephants have been found on the island of [[Tilos]]. They are similar in size to ''Palaeoloxodon mnaidriensis'' and the smaller ''Palaeoloxodon falconeri'', but the two groups indicate sexual dimorphism.<ref name=Theodorou1983>{{cite book | author = Theodorou, G. | year = 1983 | title = The dwarf elephants of the Charkadio cave on the island of Tilos (Dodekanese, Greece) | publisher = Phd Thesis [[Athens University]] | page = 321 pp}}</ref><ref name=Theodorou1988>{{cite journal | author = Theodorou, G.E. | year = 1988 | title = Environmental factors affecting the evolution of islands endemics: The Tilos example for Greece | journal = Modern Geology | volume = 13 | pages = 183–188}}</ref> 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 proven, this name should not be used when referring to the elephant remnants from Tilos. The species has been described as ''Elephas tiliensis'' in 2007<ref name=Theodorou2007/> and it is now assigned to genus ''Palaeoloxodon''.
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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 body weight of only {{convert|200|kg|abbr=on}}, 2% of its {{convert|10000|kg|abbr=on}} mainland ancestor. The species likely evolved from the earlier larger (though still strongly dwarfed) ''Palaeoloxodon xylophagou'' known from fossils dating to around 200,000 years ago.
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 {{Convert|1|m|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.<ref name=":13">{{Cite journal |last1=Athanassiou |first1=Athanassios |last2=Herridge |first2=Victoria |last3=Reese |first3=David S. |last4=Iliopoulos |first4=George |last5=Roussiakis |first5=Socrates |last6=Mitsopoulou |first6=Vassiliki |last7=Tsiolakis |first7=Efthymios |last8=Theodorou |first8=George |date=August 2015 |title=Cranial evidence for the presence of a second endemic elephant species on Cyprus |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618215005856 |journal=Quaternary International |language=en |volume=379 |pages=47–57 |bibcode=2015QuInt.379...47A |doi=10.1016/j.quaint.2015.05.065}}</ref>


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.<ref name="Bate1903"/><ref>Dorothea Bate, ''Cyprus work diary 1901–02'', 3 volumes, Natural History Museum's earth sciences library, palaeontology MSS</ref>
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.<ref name="Bate1903"/><ref>Dorothea Bate, ''Cyprus work diary 1901–02'', 3 volumes, Natural History Museum's earth sciences library, palaeontology MSS</ref>

Revision as of 19:44, 23 April 2024

Palaeoloxodon falconeri skeleton cast

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 elephant is located in Mediterranean
Sardinia
Sardinia
Favignana
Favignana
Sicily
Sicily
Malta
Malta
Crete
Crete
Kythnos
Serifos
Milos
Delos
Paros
Naxos
Astypalaia
Kasos
Kasos
Tilos
Rhodes
Rhodes
Cyprus
Cyprus
class=notpageimage|
Mediterranean islands with remains of dwarf proboscideans
genus Palaeoloxodon
genus Mammuthus
both genera present in Crete
Proboscidean remains from the Cyclades that can't be confidently assigned to a single proboscidean genus or to a dwarf species.[2]
Size comparison of the Sicilian species Palaeoloxodon falconeri, one of the smallest dwarf elephants, compared to a human

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.

Size comparison of dwarf Sardinian mammoth Mammuthus lamarmorai compared to a human

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.[5][6] 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.[6][7][8] 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.[9]

  connected during LGM[10]

Greece

Crete

Island Taxon Author
Crete Mammuthus creticus (Bate, 1907)
Palaeoloxodon creutzburgi (Kuss, 1965)

Poulakakis and others proposed in 2002 to rename all the described specimens of larger size than Mammuthus creticus under the new subspecies name Elephas antiquus creutzburgi (Kuss, 1965).[11] After DNA research, published in 2006, it has been proposed to rename Elephas (Palaeoloxodon) creticus into Mammuthus creticus (Bate, 1907). In a recent study of 2007, it was argued for the groundlessness of the theory by Poulakakis et al. in 2006,[12] showing the weak points of that DNA research.[13] However, morphological data is at least equivocal, and may also support placement in Mammuthus.

Mammuthus creticus is the earlier species known from the Early Pleistocene. It is the smallest mammoth[14] 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).[15] 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.[16]

Cyclades

Island Taxon Author
Delos Palaeoloxodon sp. Vaufrey, 1929
Naxos Palaeoloxodon lomolinoi Van der Geer et al., 2014[17]
Paros Elephantidae indet. Georgalas, 1929
Kythnos Elephantidae indet. Honea, 1975
Milos Elephantidae indet. Papp, 1953
Serifos Elephantidae indet. Papp, 1953
  connected during LGM[17]

Remains of paleoloxodontine elephants have been reported from several Cyclades islands. The Delos elephant's size is comparable to Palaeoloxodon mnaidriensis.[17] 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.[17]

Dodecanese

Island Taxon Author
Astypalaia Palaeoloxodon sp. Athanassiou et al., 2019[2]
Kasos Palaeoloxodon aff. creutzburgi Sen et al., 2014[18]
Rhodes Palaeoloxodon sp. Symeonides et al., 1974
Tilos Palaeoloxodon tiliensis (Theodorou et al. 2007)[19]

On the island of Rhodes, bones of an endemic dwarf elephant have been discovered. This elephant was similar in size to Palaeoloxodon mnaidriensis.[16]

Two groups of remains of dwarf elephants have been found on the island of Tilos. They are similar in size to Palaeoloxodon mnaidriensis and the smaller Palaeoloxodon falconeri, but the two groups indicate sexual dimorphism.[20][21] 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 proven, this name should not be used when referring to the elephant remnants from Tilos. The species has been described as Elephas tiliensis in 2007[19] and it is now assigned to genus Palaeoloxodon.

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.[22] On the other hand, DNA analysis in 2017 conclude that the closest living relative of the genus Palaeoloxodon is the African forest elephant (Loxodonta cyclotis).[23] The paper suggests that the current view of elephant evolution is "in need of substantial revision".[23] After the study of new osteological material[19] that has been excavated in anatomical connection in the Charkadio Cave on Tilos island the new species name Palaeoloxodon tiliensis has been assigned to the Tilos dwarf elephants.

Cyprus

Island Taxon Author
Cyprus Palaeoloxodon cypriotes (Bate, 1903)[24]
Palaeoloxodon xylophagou Athanassiou et al., 2015[25]

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.[26]

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.[24][27]

The Channel Islands of California

Skeletal restoration of Mammuthus exilis

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).[28] 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".[29] 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".[30]

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

Size comparison of dwarf Flores Stegodon species compared to a human

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,[15][31][32] 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) [32][33] Stegodon florensis became extinct about 50,000 years ago, around the time of the arrival of modern humans to Flores.[31]

Sulawesi

During the Late Pliocene-Early Pleistocene on Sulawesi, two species of dwarf proboscidean coinhabited the island, the elephant Stegoloxodon celebensis, and Stegodon sompoensis.[34] The former was about 150 cm (4.9 ft) tall,[35] while the latter was around 32% the size of mainland Stegodon species, with an estimated body mass of about a ton.[32] Later in the Pleistocene, these animals were replaced by larger-sized species of Stegodon and elephants,[34][36] 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.[32]

Java

The species Stegodon trigonocephalus is known from the Early-Middle Pleistocene of Java.[37] A population from the Trinil H.K locality, which likely dates to the Middle Pleistocene,[38] is around 65% the size of mainland Stegodon species.[32] Large individuals are estimated to have reached around 280 cm (9.2 ft) at the shoulders, with a body mass of around 5 tons.[15] Other smaller unnamed Stegodon species are also known from the Early Pleistocene on the island.[37] 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.[39][37]

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).[32]

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,[40] and around 23% the size of mainland species, with an estimated body mass of around 770 kilograms (1,700 lb).[32]

Luzon

On Luzon the dwarf Stegodon luzonensis is known from remains found in the Manila Basin of an uncertain Pleistocene age,[41] as well as remains found near the early Middle Pleistocene Nesorhinus butchery site.[42] It is around 40% the size of mainland Stegodon species, with a body mass of around 1.3 tons.[32] The extinct dwarf elephant Elephas beyeri is also known from the island,[32] which is estimated to have been about 1.2 metres (3.9 ft) in shoulder height.[43]

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).[44]

See also

References

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