View source for Biology:Dwarf elephant

{{short description|Prehistoric elephant species}}
[[Image:Elephas skeleton.JPG|right|thumb|''Palaeoloxodon falconeri'' skeleton cast]]
'''Dwarf elephants''' are prehistoric members of the order [[Biology:Proboscidea|Proboscidea]] which, through the process of [[Biology:Allopatric speciation|allopatric speciation]] on islands, evolved much smaller body sizes (around {{convert|1|-|2.3|m}}) in comparison with their immediate ancestors. Dwarf elephants are an example of [[Biology:Insular dwarfism|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. Some modern populations of [[Biology:Asian elephant|Asian elephant]]s have also undergone size reduction on islands to a lesser degree, resulting in populations of pygmy elephants.

[[Biology:Fossil|Fossil]] remains of dwarf elephants have been found on the Mediterranean islands of [[Place:Cyprus|Cyprus]], Malta (at Għar Dalam), Crete (in Chania  at Vamos, Stylos and in a now-underwater cave on the coast), [[Place:Sicily|Sicily]], Sardinia, the Cyclades Islands and the Dodecanese Islands, which are mostly members of the genus ''[[Biology:Palaeoloxodon|Palaeoloxodon]]'', descending from the large {{Convert|4|m|ft}} tall mainland European species ''Palaeoloxodon antiquus''. Dwarf species of elephants and ''[[Biology:Stegodon|Stegodon]]'' have been found on the islands of Indonesia and the Philippines. The Channel Islands of California once supported [[Biology:Mammuthus exilis|a dwarf species]] descended from [[Biology:Columbian mammoth|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 [[Biology:Woolly mammoth|woolly mammoth]]s that existed on Wrangel Island north of Siberia were once considered dwarfs, but are not anymore.

==The Mediterranean islands==
{{Location map many | Mediterranean
  | relief = yes
  | default_width = 500
  | caption = Mediterranean islands with remains of dwarf proboscideans<br />[[Image:Red pog.svg|10px]] genus ''Palaeoloxodon''<br />10px genus ''Mammuthus''<br />10px both genera present in Crete<br />[[Image:Black pog.svg|10px]] Proboscidean remains from the Cyclades that can't be confidently assigned to a single proboscidean genus or to a dwarf species.<ref name="Athanassiou2019">{{cite journal | doi = 10.1016/j.quascirev.2019.06.028| title = Pleistocene insular Proboscidea of the Eastern Mediterranean: A review and update| journal = Quaternary Science Reviews| volume = 218| pages = 306–321 | year = 2019| last1 = Athanassiou | first1 = A.| last2 = Van der Geer | first2 = A. A. E.| last3 = Lyras | first3 = G. A.| s2cid = 199107354}}</ref>
| label1 = Sardinia
  | pos1 = left
  | mark1 = Blue pog.svg
  | lat1_deg = 40.09
  | lon1_deg = 9.05
| label2 = Favignana
  | pos2 = left
  | lat2_deg = 37.91
  | lon2_deg = 12.32
| label3 = Sicily
  | pos3 = right
  | lat3_deg = 37.60
  | lon3_deg = 14.08
| label4 = Malta
  | pos4 = bottom
  | lat4_deg = 35.89
  | lon4_deg = 14.42
| label5 = Crete
  | pos5 = bottom
  | mark5 = Red-blue pog.svg
  | lat5_deg = 35.20
  | lon5_deg = 24.83
| label6 = Kythnos
  | pos6 = none
  | mark6 = Black pog.svg
  | mark6size = 6
  | lat6_deg = 37.39
  | lon6_deg = 24.42
| label7 = Serifos
  | pos7 = none
  | mark7 = Black pog.svg
  | mark7size = 6
  | lat7_deg = 37.16
  | lon7_deg = 24.48
| label8 = Milos
  | pos8 = none
  | mark8 = Black pog.svg
  | mark8size = 6
  | lat8_deg = 36.69
  | lon8_deg = 24.45
| label9 = Delos
  | pos9 = none
  | mark9size = 6
  | lat9_deg = 37.39
  | lon9_deg = 25.27
| label10 = Paros
  | pos10 = none
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  | lat10_deg = 37.05
  | lon10_deg = 25.18
| label11 = Naxos
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  | lon11_deg = 25.47
| label12 = Astypalaia
  | pos12 = none
  | lat12_deg = 36.58
  | lon12_deg = 26.38
| label13 = Kasos
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  | lat13_deg = 35.39
  | lon13_deg = 26.92
| label14 = Tilos
  | pos14 = none
  | lat14_deg = 36.43
  | lon14_deg = 27.37
| label15 = Rhodes
  | pos15 = right
  | lat15_deg = 36.20
  | lon15_deg = 27.97
| label16 = Cyprus
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  | lat16_deg = 35.03
  | lon16_deg = 33.23
}}
[[File:Palaeoloxodon falconeri Size Comparison.svg|left|thumb|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 [[Earth:Pleistocene|Pleistocene]], including all the major islands with the apparent exception of [[Place:Corsica|Corsica]] and the Balearics. Mediterranean dwarf elephants have generally been considered as members of the genus ''[[Biology:Palaeoloxodon|Palaeoloxodon]]'', derived from the continental [[Biology:Straight-tusked elephant|straight-tusked elephant]], ''Palaeoloxodon antiquus'' (Falconer & Cautley, 1847), [[Biology:Synonym (taxonomy)|Syn.]]: ''Elephas antiquus''. An exception is the dwarf Middle-Late Pleistocene Sardinian [[Biology:Mammoth|mammoth]], ''[[Biology:Mammuthus lamarmorai|Mammuthus lamarmorai]]'' (Major, 1883), the first endemic elephant of the Mediterranean islands recognized as belonging to the mammoth line. ''[[Biology:Mammuthus creticus|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.<ref>{{Cite journal |last1=Herridge |first1=Victoria L. |last2=Lister |first2=Adrian M. |date=2012-05-09 |title=Extreme insular dwarfism evolved in a mammoth |url=http://dx.doi.org/10.1098/rspb.2012.0671 |journal=Proceedings of the Royal Society B: Biological Sciences |volume=279 |issue=1741 |pages=3193–3200 |doi=10.1098/rspb.2012.0671 |pmid=22572206 |pmc=3385739 |issn=0962-8452}}</ref>

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,<ref>Abel's surmise is noted by Adrienne Mayor in ''The First Fossil Hunters: Paleontology in Greek and Roman Times'' (Princeton University Press) 2000. [See illus. ed., 2001: {{ISBN|0691089779}}]</ref> 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 [[Biology:Cyclopes|Cyclopes]] of [[Religion:Greek mythology|Greek mythology]].

===Italy and Malta===
[[Image:0472 - Siracusa - Museo archeologico - Elephas Falconeri Foto Giovanni Dall'Orto - 22-May-2008.jpg|thumb|''Palaeoloxodon falconeri'']]
{| class="wikitable"
|-
! Island !! Taxon !! Author
|-
| Sardinia || ''[[Biology:Mammuthus lamarmorai|Mammuthus lamarmorai]]'' || ({{Smallcaps|Major}}, 1883)
|-
| rowspan="2" style="background: lightblue" | Malta || ''[[Biology:Palaeoloxodon falconeri|Palaeoloxodon falconeri]]'' || ({{Smallcaps|Busk}}, 1869)
|-
| ''[[Biology:Palaeoloxodon mnaidriensis|Palaeoloxodon mnaidriensis]]'' || ({{Smallcaps|Adams}}, 1874)
|-
| rowspan="2" style="background: lightblue" | [[Place:Sicily|Sicily]] || ''[[Biology:Palaeoloxodon falconeri|Palaeoloxodon falconeri]]'' || ({{Smallcaps|Busk}}, 1869)
|-
| ''[[Biology:Palaeoloxodon mnaidriensis|Palaeoloxodon cf. mnaidriensis]]'' || ({{Smallcaps|Adams}}, 1874)
|}

{{legend|lightblue|connected during [[Earth: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===
====Crete====
[[Image:Cretanelephant-petermaas.jpg|thumb|right|Skeleton of a [[Biology:Mammuthus creticus|Cretan dwarf mammoth]]]]
{| class="wikitable"
|-
! Island !! Taxon !! Author
|-
| rowspan="2" | Crete || ''[[Biology:Mammuthus creticus|Mammuthus creticus]]'' || ({{Smallcaps|Bate}}, 1907)
|-
| ''[[Biology:Palaeoloxodon creutzburgi|Palaeoloxodon creutzburgi]]'' || ({{Smallcaps|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).<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=0 |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:Biology Letters|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''.

''[[Biology:Mammuthus creticus|Mammuthus creticus]]'' is the earlier species known from the [[Earth:Early Pleistocene|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 -->

====Cyclades====
{| class="wikitable"
|-
! Island !! Taxon !! Author
|-
| style="background: lightgreen" | Delos || ''Palaeoloxodon'' sp. || {{Smallcaps|Vaufrey}}, 1929
|-
| style="background: lightgreen" | Naxos || ''Palaeoloxodon lomolinoi'' || {{Smallcaps|Van der Geer}} et al., 2014<ref name="VanderGeer2014">{{cite journal | doi = 10.1016/j.palaeo.2014.04.003| title = A dwarf elephant and a rock mouse on Naxos (Cyclades, Greece) with a revision of the palaeozoogeography of the Cycladic Islands (Greece) during the Pleistocene| journal = Palaeogeography, Palaeoclimatology, Palaeoecology| volume = 404| pages = 133–144 | year = 2014| last1 = Van der Geer | first1 = A. A. E.| last2 = Lyras | first2 = G. A.| last3 = Van den Hoek Ostende | first3 = L. W.| last4 = De Vos | first4 = J.| last5 = Drinia | first5 = H.| hdl = 10795/3263| url = http://repository.edulll.gr/3263| hdl-access = free}}</ref>
|-
| style="background: lightgreen" | Paros || Elephantidae indet. || {{Smallcaps|Georgalas}}, 1929
|-
| Kythnos || Elephantidae indet. || {{Smallcaps|Honea}}, 1975
|-
| [[Place:Milos|Milos]] || Elephantidae indet. || {{Smallcaps|Papp}}, 1953
|-
| Serifos || Elephantidae indet. || {{Smallcaps|Papp}}, 1953
|}

{{legend|lightgreen|connected during LGM<ref name="VanderGeer2014"/>}}

Remains of paleoloxodontine elephants have been reported from several Cyclades islands. The Delos elephant's size is comparable to ''Palaeoloxodon mnaidriensis''.<ref name="VanderGeer2014"/> 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''.<ref name="VanderGeer2014"/>

====Dodecanese====
[[File:Rekhmire_tomb_elephant.jpg|thumb|Leashed dwarf elephant from Rekhmire's tomb, presented as tribute by foreign peoples along with normal sized tusks and a bear or big cat]]
{| class="wikitable"
|-
! Island !! Taxon !! Author
|-
| Astypalaia || ''Palaeoloxodon'' sp. || {{Smallcaps|Athanassiou}} et al., 2019<ref name="Athanassiou2019"/>
|-
| Kasos || ''Palaeoloxodon'' [[Biology:Species affinis|aff.]] ''creutzburgi'' || {{Smallcaps|Sen}} et al., 2014<ref>{{cite journal | doi = 10.5735/086.051.0204| title = Late Pleistocene Dwarf Elephants from the Aegean Islands of Kassos and Dilos, Greece| journal = Annales Zoologici Fennici| volume = 51| pages = 27–42 | year = 2014| last1 = Sen | first1 = S.| last2 = Barrier | first2 = E.| last3 = Crété | first3 = X.| issue = 1–2| s2cid = 85190966}}</ref>
|-
| Rhodes || ''Palaeoloxodon'' sp. || {{Smallcaps|Symeonides}} et al., 1974
|-
| Tilos || ''Palaeoloxodon tiliensis'' || ({{Smallcaps|Theodorou}} et al. 2007)<ref name=Theodorou2007>{{cite journal |author1=Theodorou, G.E. |author2=Symeonides, N. |author3=Stathopoulou, E. | year = 2007 | title = ''Elephas tiliensis'' n. sp. from Tilos island (Dodecanese, Greece) | journal = Hellenic Journal of Geosciences | volume = 42 | pages = 19–32}}</ref>
|}

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

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''.

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''.<ref>{{cite journal | last1=Poulakakis | first1=Nikos | last2=Theodorou | first2=Georgios E. | last3=Zouros | first3=Eleftherios | last4=Mylonas | first4=Moysis | title=Molecular Phylogeny of the Extinct Pleistocene Dwarf Elephant Palaeoloxodon antiquus falconeri from Tilos Island, Dodekanisa, Greece | journal=Journal of Molecular Evolution | publisher=Springer Science and Business Media LLC | volume=55 | issue=3 | date=2002-09-01 | issn=0022-2844 | doi=10.1007/s00239-002-2337-x | pages=364–374 | pmid=12187389| s2cid=21140948 }}</ref> On the other hand, DNA analysis in 2017 conclude that the closest living relative of the genus ''Palaeoloxodon'' is the [[Biology:African forest elephant|African forest elephant]] (''Loxodonta cyclotis'').<ref name=Meyer>{{cite journal |last1=Meyer |first1=Matthias |last2=Palkopoulou |first2=Eleftheria |last3=Baleka |first3=Sina |last4=Stiller |first4=Mathias |last5=Penkman |first5=Kirsty E H |author5-link= Kirsty Penkman |last6=Alt |first6=Kurt W |last7=Ishida |first7=Yasuko |last8=Mania |first8=Dietrich |last9=Mallick |first9=Swapan |last10=Meijer |first10=Tom |last11=Meller |first11=Harald |last12=Nagel |first12=Sarah |last13=Nickel |first13=Birgit |last14=Ostritz |first14=Sven |last15=Rohland |first15=Nadin |last16=Schauer |first16=Karol |last17=Schüler |first17=Tim |last18=Roca |first18=Alfred L |last19=Reich |first19=David |last20=Shapiro |first20=Beth |last21=Hofreiter |first21=Michael |title=Palaeogenomes of Eurasian straight-tusked elephants challenge the current view of elephant evolution |journal=eLife |date=6 June 2017 |volume=6 |page=e25413 |doi=10.7554/eLife.25413 |pmid=28585920 |pmc=5461109 |doi-access=free}}</ref> The paper suggests that the current view of elephant evolution is "in need of substantial revision".<ref name=Meyer/> After the study of new osteological material<ref name=Theodorou2007/> 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.

The Tilos species was the last dwarf elephant to become extinct, with the most recent remains having been dated to just 3040-1840 BCE. The Ancient Egyptian tomb of Rekhmire (built 1470-1445 BCE) contains a painting of a small leashed elephant with long tusks, large ears, and brown skin, that is interpreted by some authors as a live depiction of a dwarf elephant.<ref>Masseti, M. (2008). The most ancient explorations of the Mediterranean. ''Proc. Calif. Acad. Sci.'' 4th Ser, 59(Suppl I), 1-18.</ref>

===Cyprus===
{| class="wikitable"
|-
! Island !! Taxon !! Author
|-
| rowspan="2"| [[Place:Cyprus|Cyprus]] || ''[[Biology:Cyprus dwarf elephant|Palaeoloxodon cypriotes]]'' || ({{Smallcaps|Bate}}, 1903)<ref name="Bate1903">Bate, D. M. A.: ''Preliminary Note on the Discovery of a Pigmy Elephant in the Pleistocene of Cyprus'' in ''Proceedings of the Royal Society of London'' Vol. 71 (1902 - 1903), pp. 498-500</ref>
|-
| ''Palaeoloxodon xylophagou'' || {{Smallcaps|Athanassiou}} et al., 2015<ref>{{cite journal | doi = 10.1016/j.quaint.2015.05.065| title = Cranial evidence for the presence of a second endemic elephant species on Cyprus| journal = Quaternary International| volume = 379| pages = 47–57 | year = 2015| last1 = Athanassiou | first1 = A.| last2 = Herridge | first2 = V.| last3 = Reese | first3 = D. S.| last4 = Iliopoulos | first4 = G.| last5 = Roussiakis | first5 = S.| last6 = Mitsopoulou | first6 = V.| last7 = Tsiolakis | first7 = E.| last8 = Theodorou | first8 = G.}}</ref>
|}

The Cyprus dwarf elephant survived at least until 11,000 BCE.<!-- typo? --> Its estimated body weight was only {{convert|200|kg|abbr=on}}, only 2% of its {{convert|10000|kg|abbr=on}} ancestor. Molars of this dwarf elephant are reduced to approximately 40% the size of mainland [[Biology:Straight-tusked elephant|straight-tusked elephant]]s.

Remains of the species were first discovered and recorded by Dorothea Bate in a cave in the Kyrenia hills of [[Place:Cyprus|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>

==The Channel Islands of California==
[[File:M._exilis_skeletal.png|thumb|Skeletal restoration of ''Mammuthus exilis'']]
The [[Biology:Mammoth|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 [[Biology:Pygmy mammoth|pygmy mammoth]] (''Mammuthus exilis'').<ref>{{cite news |url=http://www.latimes.com/local/lanow/la-me-ln-mammoth-fossil-channel-islands-20160915-snap-story.html |title=Well-preserved mammoth skull unearthed on Channel Islands puzzles scientists |newspaper={{wipe|Los Angeles Times}} |first=Veronica |last=Rocha |date=16 September 2016 |access-date=24 September 2016}}</ref> Channel Islands mammoths ranged from {{convert|150|–|190|cm|abbr=on}} in shoulder height.

==Wrangel Island==
During the last ice age, [[Biology:Woolly mammoth|woolly mammoth]]s (''Mammuthus primigenius'') lived on Wrangel Island in the [[Earth:Arctic Ocean|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 {{convert|180|–|230|cm|abbr=on}} in shoulder height and were for a time considered "dwarf mammoths".<ref name=Nature>{{cite journal |author1=Vartanyan, S.L. |author2=Garutt, V.E. |author3=Sher, A.V. | year = 1993 | title = Holocene dwarf mammoths from Wrangel Island in the Siberian Arctic | journal = Nature | volume = 362 | pages = 337–340 | doi=10.1038/362337a0 | issue=6418 | pmid=29633990|s2cid=4249191 }}</ref> 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".<ref>{{cite journal|last=Tikhonov |first=Alexei |author2=Larry Agenbroad |author3=Sergey Vartanyan |year=2003 |title=Comparative analysis of the mammoth populations on Wrangel Island and the Channel Islands |journal=Deinsea |volume=9 |pages=415–420 |issn=0923-9308 |url=http://www.nmr.nl/nmr/pages/showPage.do?itemid=1896&instanceid=16 |url-status=dead |archive-url=https://web.archive.org/web/20120611113051/http://www.nmr.nl/nmr/pages/showPage.do?itemid=1896&instanceid=16 |archive-date=2012-06-11 }}</ref>

==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 ''[[Biology:Stegodon|Stegodon]]''.

===Flores===
[[File:Dwarf stegodon size comparison.svg|thumb|300x300px|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 {{convert|120|cm|ft||abbr=on}} tall at the shoulder and weighed about {{Convert|350-400|kg|lb}}. This species became extinct around 1 million years ago,<ref name="proboscideans">{{Cite journal |last1=Larramendi |first1=A. |year=2016 |title=Shoulder height, body mass and shape of proboscideans |url=https://www.app.pan.pl/archive/published/app61/app001362014.pdf |journal=Acta Palaeontologica Polonica |volume=61 |doi=10.4202/app.00136.2014 |s2cid=2092950}}</ref><ref name="van den Bergh-2022" /><ref name="Geer-2016" /> 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 {{convert|190|cm|ft||abbr=on}} 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 {{convert|130|cm|ft||abbr=on}}, and a body mass of about {{Convert|570|kg|lb}} <ref name="Geer-2016" /><ref>Puspaningrum, Mika; Van Den Bergh, Gerrit; Chivas, Allan; Setiabudi, Erick; Kurniawan, Iwan; Brumm, Adam; and Sutikna, Thomas, "[https://ro.uow.edu.au/smhpapers/2035 Preliminary results of dietary and environmental reconstructions of Early to Middle Pleistocene Stegodons from the So'a Basin of Flores, Indonesia, based on enamel stable isotope records]" (2014). Faculty of Science, Medicine and Health - Papers: part A. 2035.</ref> ''Stegodon florensis'' became extinct about 50,000 years ago, around the time of the arrival of modern humans to Flores.<ref name="van den Bergh-2022">{{Cite journal |last1=van den Bergh |first1=Gerrit D. |last2=Alloway |first2=Brent V. |last3=Storey |first3=Michael |last4=Setiawan |first4=Ruly |last5=Yurnaldi |first5=Dida |last6=Kurniawan |first6=Iwan |last7=Moore |first7=Mark W. |last8=Jatmiko |last9=Brumm |first9=Adam |last10=Flude |first10=Stephanie |last11=Sutikna |first11=Thomas |last12=Setiyabudi |first12=Erick |last13=Prasetyo |first13=Unggul W. |last14=Puspaningrum |first14=Mika R. |last15=Yoga |first15=Ifan |date=October 2022 |title=An integrative geochronological framework for the Pleistocene So'a basin (Flores, Indonesia), and its implications for faunal turnover and hominin arrival |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379122003523 |journal=Quaternary Science Reviews |language=en |volume=294 |pages=107721 |doi=10.1016/j.quascirev.2022.107721|hdl=10072/418777 |s2cid=252290750 |hdl-access=free }}</ref>

===Sulawesi===
During the Late Pliocene-Early Pleistocene on Sulawesi, two species of dwarf proboscidean coinhabited the island, the elephant [[Biology:Stegoloxodon|''Stegoloxodon celebensis'']], and ''Stegodon sompoensis.<ref name="Vos-2007">{{Citation |last1=Vos |first1=John de |title=Patterns in Insular Evolution of Mammals: A Key to Island Palaeogeography |date=2007 |url=http://link.springer.com/10.1007/978-1-4020-6374-9_10 |work=Biogeography, Time, and Place: Distributions, Barriers, and Islands |volume=29 |pages=315–345 |editor-last=Renema |editor-first=Willem |access-date=2023-08-10 |place=Dordrecht |publisher=Springer Netherlands |doi=10.1007/978-1-4020-6374-9_10 |isbn=978-1-4020-6373-2 |last2=Ostende |first2=Lars W. van den Hoek |last3=Bergh |first3=Gert D. van den|series=Topics in Geobiology }}</ref>'' The former was about {{convert|150|cm|ft||abbr=on}} tall,<ref>{{Cite journal |last1=Alink |first1=Gerrit |last2=Adhityatama |first2=Shinatria |last3=Simanjuntak |first3=Truman |date=2017-12-29 |title=The Descriptive Analysis of Palaeolithic Stone Tools from Sulawesi, Collected by the Indonesian-Dutch Expedition in 1970 |url=http://jurnalarkeologi.kemdikbud.go.id/index.php/amerta/article/view/252 |journal=AMERTA |volume=35 |issue=2 |pages=75 |doi=10.24832/amt.v35i2.252|doi-access=free }}</ref> while the latter was around 32% the size of mainland ''Stegodon'' species, with an estimated body mass of about a ton.<ref name="Geer-2016" /> Later in the Pleistocene, these animals were replaced by larger-sized species of ''Stegodon'' and elephants,<ref name="Vos-2007" /><ref>{{Cite journal |last1=Brumm |first1=Adam |last2=Hakim |first2=Budianto |last3=Ramli |first3=Muhammad |last4=Aubert |first4=Maxime |last5=van den Bergh |first5=Gerrit D. |last6=Li |first6=Bo |last7=Burhan |first7=Basran |last8=Saiful |first8=Andi Muhammad |last9=Siagian |first9=Linda |last10=Sardi |first10=Ratno |last11=Jusdi |first11=Andi |last12=Abdullah |last13=Mubarak |first13=Andi Pampang |last14=Moore |first14=Mark W. |last15=Roberts |first15=Richard G. |date=2018-04-11 |editor-last=Hardy |editor-first=Karen |title=A reassessment of the early archaeological record at Leang Burung 2, a Late Pleistocene rock-shelter site on the Indonesian island of Sulawesi |journal=PLOS ONE |language=en |volume=13 |issue=4 |pages=e0193025 |doi=10.1371/journal.pone.0193025 |issn=1932-6203 |pmc=5894965 |pmid=29641524 |doi-access=free }}</ref> 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.<ref name="Geer-2016" />

===Java===
The species ''Stegodon trigonocephalus'' is known from the Early-Middle Pleistocene of Java.<ref name="Puspaningrum-2020">{{Cite journal |last1=Puspaningrum |first1=Mika R. |last2=van den Bergh |first2=Gerrit D. |last3=Chivas |first3=Allan R. |last4=Setiabudi |first4=Erick |last5=Kurniawan |first5=Iwan |date=January 2020 |title=Isotopic reconstruction of Proboscidean habitats and diets on Java since the Early Pleistocene: Implications for adaptation and extinction |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119303178 |journal=Quaternary Science Reviews |language=en |volume=228 |pages=106007 |doi=10.1016/j.quascirev.2019.106007|s2cid=212876762 }}</ref> A population from the Trinil H.K locality, which likely dates to the Middle Pleistocene,<ref>{{Cite journal |last1=L. Hilgen |first1=Sander |last2=Pop |first2=Eduard |last3=Adhityatama |first3=Shinatria |last4=A. Veldkamp |first4=Tom |last5=W.K. Berghuis |first5=Harold |last6=Sutisna |first6=Indra |last7=Yurnaldi |first7=Dida |last8=Dupont-Nivet |first8=Guillaume |last9=Reimann |first9=Tony |last10=Nowaczyk |first10=Norbert |last11=F. Kuiper |first11=Klaudia |last12=Krijgsman |first12=Wout |last13=B. Vonhof |first13=Hubert |last14=Ekowati |first14=Dian Rahayu |last15=Alink |first15=Gerrit |date=February 2023 |title=Revised age and stratigraphy of the classic Homo erectus-bearing succession at Trinil (Java, Indonesia) |journal=Quaternary Science Reviews |language=en |volume=301 |pages=107908 |doi=10.1016/j.quascirev.2022.107908|doi-access=free |hdl=1887/3515249 |hdl-access=free }}</ref> is around 65% the size of mainland ''Stegodon'' species.<ref name="Geer-2016" /> Large individuals are estimated to have reached around {{convert|280|cm|ft||abbr=on}} at the shoulders, with a body mass of around 5 tons.<ref name="proboscideans" /> Other smaller unnamed ''Stegodon'' species are also known from the Early Pleistocene on the island.<ref name="Puspaningrum-2020" /> The extinct dwarf elephant species [[Biology:Stegoloxodon|''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.<ref>{{Cite journal |last1=Markov |first1=Georgi N. |last2=Saegusa |first2=Haruo |date=2008-09-01 |title=On the validity of Stegoloxodon Kretzoi, 1950 (Mammalia: Proboscidea) |url=https://biotaxa.org/Zootaxa/article/view/zootaxa.1861.1.5 |journal=Zootaxa |volume=1861 |issue=1 |pages=55 |doi=10.11646/zootaxa.1861.1.5 |issn=1175-5334}}</ref><ref name="Puspaningrum-2020" />

=== 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 {{Convert|250|kg|lb}}.<ref name="Geer-2016" />

=== Timor ===
The species ''S. timorensis'' is known from the Middle Pleistocene of Timor. It is a small-sized species, only slightly larger than ''S. sondaarii,''<ref>{{Cite journal |last1=Louys |first1=Julien |last2=Price |first2=Gilbert J. |last3=O’Connor |first3=Sue |date=2016-03-10 |title=Direct dating of Pleistocene stegodon from Timor Island, East Nusa Tenggara |journal=PeerJ |language=en |volume=4 |pages=e1788 |doi=10.7717/peerj.1788 |pmid=26989625 |issn=2167-8359|pmc=4793331 |doi-access=free }}</ref> and around 23% the size of mainland species, with an estimated body mass of around {{Convert|770|kg|lb}}.<ref name="Geer-2016" />

=== Luzon ===
On Luzon the dwarf ''Stegodon luzonensis'' is known from remains found in the Manila Basin of an uncertain Pleistocene age,<ref>von Koenigswald GHR. 1956. Fossil mammals from the Philippines. Quezon City: National Research Council of the Philippines, 1–14</ref> as well as remains found near the early Middle Pleistocene ''[[Biology:Nesorhinus|Nesorhinus]]'' butchery site.<ref>{{Cite journal |last1=Ingicco |first1=T. |last2=van den Bergh |first2=G. D. |last3=Jago-on |first3=C. |last4=Bahain |first4=J.-J. |last5=Chacón |first5=M. G. |last6=Amano |first6=N. |last7=Forestier |first7=H. |last8=King |first8=C. |last9=Manalo |first9=K. |last10=Nomade |first10=S. |last11=Pereira |first11=A. |last12=Reyes |first12=M. C. |last13=Sémah |first13=A.-M. |last14=Shao |first14=Q. |last15=Voinchet |first15=P. |date=May 2018 |title=Earliest known hominin activity in the Philippines by 709 thousand years ago |url=https://www.nature.com/articles/s41586-018-0072-8 |journal=Nature |language=en |volume=557 |issue=7704 |pages=233–237 |doi=10.1038/s41586-018-0072-8 |pmid=29720661 |s2cid=13742336 |issn=0028-0836}}</ref> It is around 40% the size of mainland ''Stegodon'' species, with a body mass of around 1.3 tons.<ref name="Geer-2016" /> The extinct dwarf elephant ''Elephas beyeri'' is also known from the island,<ref name="Geer-2016">{{Cite journal |last1=Geer |first1=Alexandra A. E. |last2=Bergh |first2=Gerrit D. |last3=Lyras |first3=George A. |last4=Prasetyo |first4=Unggul W. |last5=Due |first5=Rokus Awe |last6=Setiyabudi |first6=Erick |last7=Drinia |first7=Hara |date=August 2016 |title=The effect of area and isolation on insular dwarf proboscideans |url=https://onlinelibrary.wiley.com/doi/10.1111/jbi.12743 |journal=Journal of Biogeography |language=en |volume=43 |issue=8 |pages=1656–1666 |doi=10.1111/jbi.12743 |s2cid=87958022 |issn=0305-0270}}</ref> which is estimated to have been about {{Convert|1.2|m|ft}} in shoulder height.<ref>{{Cite journal |last=Koenigswald |first=G.H.R. |date=1956 |title=Fossil mammals from the Philippines |journal=Proceedings of the Fourth Far-Eastern Prehistory Congress}}</ref>

=== 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 {{Convert|400|kg|lb}}.<ref>{{Cite book |last=Ong |first=Perry |url=https://www.researchgate.net/publication/303496589 |title=The Philippine Archipelago |publisher=Asia Publishing Co. Ltd. |year=1998 |location=Makati city, Philippines |pages=227–255 |chapter=The Philippine Menagerie}}</ref>

==See also==
* Borneo elephant

==References==<!-- Taxonomy is inconsistent on entire group of Elephant, Elephas, Palaeoloxodon... pages. See also Biol. Lett.2:451 doi:10.1098/rsbl.2006.0467 but see also BiolLett3:55,57,60. First and last papers have digital appendices. -->
{{Reflist}}
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{{DEFAULTSORT:Dwarf Elephant}}
[[Category:Prehistoric elephants]]
[[Category:Holocene extinctions]]

pt:Elefante-pigmeu

{{Sourceattribution|Dwarf elephant}}