Editing Elephant bird

{{Short description|Extinct order of birds}}
{{Use dmy dates|date=May 2021}}
{{Automatic taxobox
| name = Elephant birds
| fossil_range = {{fossil range|Quaternary}}
| image = Aepyornis maximus.jpg
| image_caption = ''[[Aepyornis maximus]]'' skeleton and egg
| display_parents = 2
| parent_authority = 
| taxon = Aepyornithiformes
| authority = [[Alfred Newton|Newton]], 1884<ref name="SN">Brands, S. (2008)</ref>
| type_species = {{extinct}}''[[Aepyornis maximus]]''
| type_species_authority = [[Isidore Geoffroy Saint-Hilaire|Hilaire]], 1851
| subdivision_ranks = Genera
| subdivision = * {{extinct}}''[[Aepyornis]]''
* {{extinct}}''[[Mullerornis]]''
* {{extinct}}''[[Vorombe]]''?
}}

'''Elephant birds''' are extinct [[flightless birds]] belonging to the [[Order (biology)|order]] '''Aepyornithiformes''' that were native to the island of [[Madagascar]]. They are thought to have gone extinct around 1000 CE, likely as a result of human activity. Elephant birds comprised three species, one in the genus ''[[Mullerornis]]'', and two in ''[[Aepyornis]].'' ''Aepyornis maximus'' is possibly the largest bird to have ever lived, with their eggs being the largest known for any [[amniote]]. Elephant birds are [[palaeognaths]] (whose flightless representatives are often known as [[ratite]]s), and their closest living relatives are [[Kiwi (bird)|kiwi]] (found only in New Zealand), suggesting that ratites did not diversify by [[vicariance]] during the breakup of [[Gondwana]] but instead [[convergently evolved]] flightlessness from ancestors that dispersed more recently by flying.

== Discovery ==
Elephant birds have been extinct since at least the 17th century. [[Étienne de Flacourt]], a French governor of Madagascar during the 1640s and 1650s, mentioned an ostrich-like bird, said to inhabit unpopulated regions, although it is unclear whether he was repeating folk tales from generations earlier. In 1659, Flacourt wrote of the "vouropatra&nbsp;– a large bird which haunts the Ampatres and lays eggs like the ostriches; so that the people of these places may not take it, it seeks the most lonely places."<ref name="Flacourt1658">{{cite book|author=Etienne de Flacourt|title=Histoire de la grande isle Madagascar|url= https://books.google.com/books?id=69w_AAAAcAAJ&pg=PA165| page = 165|access-date=21 May 2013|year= 1658|publisher= chez Alexandre Lesselin}}</ref><ref name="ley196608">{{Cite magazine |last=Ley |first=Willy |author-link= Willy Ley |date=August 1966 |title= Scherazade's Island |department=For Your Information |url=https://archive.org/stream/Galaxy_v24n06_1966-08#page/n45/mode/2up |magazine= Galaxy Science Fiction |pages= 45–55}}</ref> There has been speculation, especially popular in the latter half of the 19th century, that the legendary [[Roc (mythology)|roc]] from the accounts of [[Marco Polo]] was ultimately based on elephant birds, but this is disputed.<ref name=":3">{{Cite journal |last=Buffetaut |first=Eric |date=2019-09-06 |title=Early illustrations of Aepyornis eggs (1851–1887): from popular science to Marco Polo's roc bird |url=https://sciencepress.mnhn.fr/en/periodiques/anthropozoologica/54/12 |journal=Anthropozoologica |volume=54 |issue=1 |pages=111 |doi=10.5252/anthropozoologica2019v54a12 |s2cid=203423023 |issn=0761-3032}}</ref>

Between 1830 and 1840, European travelers in Madagascar saw giant eggs and eggshells.{{r|ley196608}} British observers were more willing to believe the accounts of giant birds and eggs because they knew of the [[moa]] in New Zealand.{{r|ley196608}} In 1851 the genus ''Aepyornis'' and species ''A. maximus'' were scientifically described in a paper presented to the [[Paris Academy of Sciences]] by [[Isidore Geoffroy Saint-Hilaire]], based on bones and eggs recently obtained from the island, which resulted in wide coverage in the popular presses of the time, particularly due to their very large eggs.<ref name=":3" />

Two whole eggs have been found in dune deposits in southern [[Western Australia]], one in the 1930s (the Scott River egg) and one in 1992 (the [[Cervantes, Western Australia|Cervantes]] egg); both have been identified as ''Aepyornis maximus'' rather than ''[[Genyornis newtoni]],'' an extinct giant bird known from the Pleistocene of Australia. It is hypothesized that the eggs floated from Madagascar to Australia on the [[Antarctic Circumpolar Current]]. Evidence supporting this is the finding of two fresh [[penguin]] eggs that washed ashore on Western Australia but may have originated in the [[Kerguelen Islands]], and an [[ostrich egg]] found floating in the [[Timor Sea]] in the early 1990s.<ref name="Long1998">{{cite journal |last1=Long |first1=J. A. |last2=Vickers-Rich |first2=P. |last3=Hirsch |first3=K. |last4=Bray |first4=E. |last5=Tuniz |first5=C. |display-authors=3 |date=1998 |title=The Cervantes egg: an early Malagasy tourist to Australia |url=http://museum.wa.gov.au/research/records-supplements/records/cervantes-egg-early-malagsy-tourist-australia |journal=Records of the Western Australian Museum |volume=19 |issue=Part 1 |pages=39–46 |access-date=2014-04-24}}</ref>

== Taxonomy and biogeography ==
[[File:Mullerornis agilis.jpg|thumb|left|''[[Mullerornis modestus]]'']]
[[File:Aepyornis maximus 01 L.D..jpg|upright|thumb|''Aepyornis maximus'' restoration]]
Like the [[ostrich]], [[Rhea (bird)|rhea]], [[cassowary]], [[emu]], [[Kiwi (bird)|kiwi]] and extinct [[moa]], elephant birds were ratites; they could not fly, and their [[sternum|breast bones]] had no [[keel (bird anatomy)|keel]]. Because Madagascar and [[Africa]] separated before the ratite [[lineage (evolution)|lineage]] arose,<ref>Yoder, A. D. & Nowak, M. D. (2006)</ref> elephant birds are thought to have dispersed and become flightless and gigantic ''[[in situ]]''.<ref>van Tuinen, M. et al. (1998)</ref>

More recently, it has been deduced from DNA sequence comparisons that the closest living relatives of elephant birds are New Zealand [[Kiwi (bird)|kiwi]],<ref name="Mitchell2014">{{Cite journal | doi = 10.1126/science.1251981| pmid = 24855267| title = Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution| journal = Science| volume = 344| issue = 6186| pages = 898–900| date = 2014-05-23| display-authors= 3| last1 = Mitchell | first1 = K. J.| last2 = Llamas | first2 = B.| last3 = Soubrier | first3 = J.| last4 = Rawlence | first4 = N. J.| last5 = Worthy | first5 = T. H.| last6 = Wood | first6 = J.| last7 = Lee | first7 = M. S. Y.| last8 = Cooper | first8 = A.| bibcode = 2014Sci...344..898M| url = https://dspace.flinders.edu.au/xmlui/bitstream/2328/35953/1/Mitchell_AncientDNA_AM2014.pdf| hdl = 2328/35953| s2cid = 206555952| hdl-access = free}}</ref> though the split between the two groups is deep, with the two lineages being estimated to have diverged from each other around 54 million years ago.<ref name=":0" />

Placement of Elephant birds within Palaeognathae, after:<ref>{{Cite journal |last=Takezaki |first=Naoko |date=2023-06-01 |editor-last=Holland |editor-first=Barbara |title=Effect of Different Types of Sequence Data on Palaeognath Phylogeny |journal=Genome Biology and Evolution |language=en |volume=15 |issue=6 |doi=10.1093/gbe/evad092 |issn=1759-6653 |pmc=10262969 |pmid=37227001}}</ref><ref>{{Cite journal |last1=Urantówka |first1=Adam Dawid |last2=Kroczak |first2=Aleksandra |last3=Mackiewicz |first3=Paweł |date=December 2020 |title=New view on the organization and evolution of Palaeognathae mitogenomes poses the question on the ancestral gene rearrangement in Aves |journal=BMC Genomics |language=en |volume=21 |issue=1 |page=874 |doi=10.1186/s12864-020-07284-5 |doi-access=free |issn=1471-2164 |pmc=7720580 |pmid=33287726}}</ref> 

{{clade
|label1=[[Paleognathae]]
|1={{clade
|1=[[Struthionidae]] (ostriches)
|2={{clade
|1=[[Rheidae]] (rheas)
|2={{clade
|1=[[Tinamidae]] (tinamou)
|2={{extinct}}[[Dinornithiformes]] (moa)
}}
|3={{clade
|1=[[Apterygidae]] (kiwis)
|2={{extinct}}'''Aepyornithiformes''' (elephant birds)
}}
|4=[[Casuariiformes]] (emu, cassowary)
}}}}}}

The ancestors of elephant birds are thought to have arrived in Madagascar well after [[Gondwana]] broke apart. The existence of possible flying [[palaeognathae]] in the Miocene such as ''[[Proapteryx]]'' further supports the view that ratites did not diversify in response to [[vicariance]]. Gondwana broke apart in the Cretaceous and their phylogenetic tree does not match the process of [[continental drift]]. Madagascar has a notoriously poor Cenozoic terrestrial fossil record, with essentially no fossils between the end of the Cretaceous ([[Maevarano Formation]]) and the Late Pleistocene.<ref>{{Cite journal|last1=Samonds|first1=Karen E.|last2=Zalmout|first2=Iyad S.|last3=Irwin|first3=Mitchell T. |last4= Krause| first4= David W.|last5=Rogers|first5=Raymond R.|last6=Raharivony|first6=Lydia L.| display-authors= 3| date=2009-12-12|title=Eotheroides lambondrano, new middle Eocene seacow (Mammalia, Sirenia) from the Mahajanga Basin, northwestern Madagascar|journal=Journal of Vertebrate Paleontology|language=en|volume=29|issue=4|pages=1233–1243|doi=10.1671/039.029.0417|bibcode=2009JVPal..29.1233S |s2cid=59466434|issn=0272-4634}}</ref> Complete mitochondrial genomes obtained from elephant birds eggshells suggest that ''Aepyornis'' and ''Mullerornis'' are significantly genetically divergent from each other, with  [[molecular clock]] analyses estimating the split at around 27-30 million years ago, during the [[Oligocene]] epoch.<ref name=":0">{{Cite journal |display-authors= 3| last1= Grealy| first1= Alicia| last2= Phillips| first2= Matthew |last3= Miller| first3= Gifford|last4=Gilbert|first4=M. Thomas P.|last5=Rouillard|first5=Jean-Marie |last6= Lambert |first6= David| last7= Bunce| first7= Michael |last8=Haile|first8=James|date=April 2017|title=Eggshell palaeogenomics: Palaeognath evolutionary history revealed through ancient nuclear and mitochondrial DNA from Madagascan elephant bird (Aepyornis sp.) eggshell|journal=Molecular Phylogenetics and Evolution |language= en |volume=109|pages=151–163|doi=10.1016/j.ympev.2017.01.005|pmid=28089793| bibcode= 2017MolPE.109..151G}}</ref><ref>{{cite journal |last1=Grealy |first1=Alicia |last2=Miller |first2=Gifford H. |last3=Phillips |first3=Matthew J. |last4=Clarke |first4=Simon J. |last5=Fogel |first5=Marilyn |last6=Patalwala |first6=Diana |last7=Rigby |first7=Paul |last8=Hubbard |first8=Alysia |last9=Demarchi |first9=Beatrice |last10=Collins |first10=Matthew |last11=Mackie |first11=Meaghan |last12=Sakalauskaite |first12=Jorune |last13=Stiller |first13=Josefin |last14=Clarke |first14=Julia A. |last15=Legendre |first15=Lucas J. |last16=Douglass |first16=Kristina |last17=Hansford |first17=James |last18=Haile |first18=James |last19=Bunce |first19=Michael |title=Molecular exploration of fossil eggshell uncovers hidden lineage of giant extinct bird |journal=Nature Communications |date=28 February 2023 |volume=14 |issue=1 |page=914 |doi=10.1038/s41467-023-36405-3 |pmid=36854679 |pmc=9974994 |bibcode=2023NatCo..14..914G }}</ref>

=== Species ===
Up to 10 or 11 species in the genus ''Aepyornis'' have been described,{{sfn|Brodkorb|1963}} but the validity of many have been disputed, with numerous authors treating them all in just one species, ''A. maximus''. Up to three species have been described in ''Mullerornis''.<ref name="Davies">Davies, S. J. J. F. (2003)</ref> Recent work has restricted the number of elephant bird species to three, with two in ''Aepyornis'', one in ''Mullerornis''.<ref name="Molecular exploration of fossil eggshell uncovers hidden lineage of giant extinct bird">{{Cite journal |last1=Grealy |first1=Alicia |last2=Miller |first2=Gifford H. |last3=Phillips |first3=Matthew J. |last4=Clarke |first4=Simon J. |last5=Fogel |first5=Marilyn |last6=Patalwala |first6=Diana |last7=Rigby |first7=Paul |last8=Hubbard |first8=Alysia |last9=Demarchi |first9=Beatrice |last10=Collins |first10=Matthew |last11=Mackie |first11=Meaghan |last12=Sakalauskaite |first12=Jorune |last13=Stiller |first13=Josefin |last14=Clarke |first14=Julia A. |last15=Legendre |first15=Lucas J. |date=2023-02-28 |title=Molecular exploration of fossil eggshell uncovers hidden lineage of giant extinct bird |journal=Nature Communications |language=en |volume=14 |issue=1 |pages=914 |doi=10.1038/s41467-023-36405-3 |issn=2041-1723|doi-access=free |pmid=36854679 |pmc=9974994 |bibcode=2023NatCo..14..914G }}</ref> 
* '''Order Aepyornithiformes''' <small>Newton 1884</small> [Aepyornithes <small>Newton 1884</small>]{{sfn|Brodkorb|1963}}
** '''Genus ''[[Aepyornis]]''''' <small>Geoffroy Saint-Hilaire 1850</small><ref>{{cite book |last1=Hume |first1=Julian P. |url=https://books.google.com/books?id=z7RIAkGw0-UC&q=greater+elephant-bird&pg=PA26 |title=Extinct birds |last2=Walters |first2=Michael |publisher=T&AD Poyser |year=2012 |isbn=978-1-4081-5861-6 |pages=544}}</ref> (Synonym: ''Vorombe'' <small>Hansford & Turvey 2018</small>)
*** ''Aepyornis hildebrandti'' <small>[[Carlos Burckhardt|Burckhardt]], 1893</small> (Possibly divided into two subspecies<ref name="Molecular exploration of fossil eggshell uncovers hidden lineage of giant extinct bird" />) 
*** ''Aepyornis maximus'' <small>[[Isidore Geoffroy Saint-Hilaire|Hilaire]], 1851</small> 
** '''Genus ''[[Mullerornis]]''''' <small>Milne-Edwards & Grandidier 189</small>4
*** ''Mullerornis modestus'' <small>(Milne-Edwards & Grandidier 1869) Hansford & Turvey 2018</small>
All elephant birds are usually placed in the single family Aepyornithidae, but some authors suggest ''Aepyornis'' and ''Mullerornis'' should be placed in separate families within the Aepyornithiformes, with the latter placed into Mullerornithidae.<ref name="Molecular exploration of fossil eggshell uncovers hidden lineage of giant extinct bird" />

== Description ==
[[File:Giantbirds.png|thumb|left|Size of ''Aepyornis maximus'' (center, in purple) compared to a human, a common ostrich (''Struthio camelus,'' second from right, in maroon), and several non-avian [[theropod]] [[dinosaur]]s. Grid spacings are 1.0 m.]]Elephant birds were large sized birds (the largest reaching {{Convert|3|m|ft}} tall in normal standing posture) that had vestigial wings, long legs and necks, with small heads relative to body size, which bore straight, thick conical beaks that were not hooked. The tops of elephant bird skulls display punctuated marks, which may have been attachment sites for fleshy structures or head feathers.<ref>{{Citation |last1=Angst |first1=Delphine |title=Aepyornithiformes |date=2017 |url=https://linkinghub.elsevier.com/retrieve/pii/B9781785481369500039 |work=Palaeobiology of Extinct Giant Flightless Birds |pages=65–94 |access-date=2023-05-02 |publisher=Elsevier |language=en |doi=10.1016/b978-1-78548-136-9.50003-9 |isbn=978-1-78548-136-9 |last2=Buffetaut |first2=Eric|url-access=subscription }}</ref> ''Mullerornis'' is the smallest of the elephant birds, with a body mass of around {{Convert|80|kg|lb}},<ref name="Molecular exploration of fossil eggshell uncovers hidden lineage of giant extinct bird" /> with its skeleton much less robustly built than ''Aepyornis''.<ref name=":1" /> ''A. hildebrandti'' is thought to have had a body mass of around {{Convert|230-285|kg|lb}}.<ref name="Molecular exploration of fossil eggshell uncovers hidden lineage of giant extinct bird" /> Estimates of the body mass of ''Aepyornis maximus'' span from around {{Convert|275|kg|lb}}<ref name="Extinct Birds">{{cite book
| last1 = Hume
| first1 = J. P.
| first2 = M.
| last2 = Walters
|year= 2012
|title= Extinct Birds
|publisher= A & C Black
|location= London
|pages= 19–21
|isbn=978-1-4081-5725-1}}</ref> to {{Convert|700-1000|kg|lb}}<ref name="Molecular exploration of fossil eggshell uncovers hidden lineage of giant extinct bird" /> making it one of the largest birds ever, alongside ''[[Dromornis stirtoni]]'' and ''[[Pachystruthio dmanisensis]]''.<ref name=":5">{{Cite journal |last1=Handley |first1=Warren D. |last2=Chinsamy |first2=Anusuya |last3=Yates |first3=Adam M. |last4=Worthy |first4=Trevor H. |date=2016-09-02 |title=Sexual dimorphism in the late Miocene mihirung Dromornis stirtoni (Aves: Dromornithidae) from the Alcoota Local Fauna of central Australia |url=https://doi.org/10.1080/02724634.2016.1180298 |journal=Journal of Vertebrate Paleontology |volume=36 |issue=5 |pages=e1180298 |doi=10.1080/02724634.2016.1180298 |bibcode=2016JVPal..36E0298H |s2cid=88784039 |issn=0272-4634|url-access=subscription }}</ref><ref>{{Cite journal |last1=Zelenkov |first1=Nikita V. |last2=Lavrov |first2=Alexander V. |last3=Startsev |first3=Dmitry B. |last4=Vislobokova |first4=Innessa A. |last5=Lopatin |first5=Alexey V. |date=2019-03-04 |title=A giant early Pleistocene bird from eastern Europe: unexpected component of terrestrial faunas at the time of early Homo arrival |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2019.1605521 |journal=Journal of Vertebrate Paleontology |language=en |volume=39 |issue=2 |pages=e1605521 |doi=10.1080/02724634.2019.1605521 |bibcode=2019JVPal..39E5521Z |s2cid=198384367 |issn=0272-4634|url-access=subscription }}</ref> Females of ''A. maximus'' are suggested to have been larger than the males, as is observed in other ratites.<ref name="Molecular exploration of fossil eggshell uncovers hidden lineage of giant extinct bird" />

== Biology ==
[[File:Aepyornis skull.JPG|thumb|''Aepyornis'' skull]]
Examination of brain [[endocast]]s has shown that both ''A. maximus'' and ''A. hildebrandti'' had greatly reduced [[Superior colliculus|optic lobe]]s, similar to those of their closest living relatives, the kiwis, and consistent with a similar [[nocturnal]] lifestyle. The optic lobes of ''Mullerornis'' were also reduced, but to a lesser degree, suggestive of a nocturnal or [[crepuscular]] lifestyle. ''A. maximus'' had relatively larger [[olfactory bulb]]s than ''A. hildebrandti'', suggesting that the former occupied forested habitats where the sense of smell is more useful while the latter occupied open habitats.<ref name="Torres2018">{{cite journal|last1= Torres|first1=C. R.|last2= Clarke|first2=J. A.|title= Nocturnal giants: evolution of the sensory ecology in elephant birds and other palaeognaths inferred from digital brain reconstructions|journal= Proceedings of the Royal Society B: Biological Sciences|volume= 285|issue= 1890|year= 2018|pages= 20181540|doi= 10.1098/rspb.2018.1540|pmid=30381378|pmc=6235046}}</ref>

=== Diet ===
A 2022 isotope analysis study suggested that some specimens of ''Aepyornis'' ''hildebrandti'' were mixed feeders that had a large (~48%) [[Grazing (behaviour)|grazing]] component to their diets, similar to that of the living ''[[Rhea americana]]'', while the other species (''A. maximus'', ''Mullerornis modestus'') were probably [[Browsing (herbivory)|browsers]].<ref>{{Cite journal |last1=Hansford |first1=James P. |last2=Turvey |first2=Samuel T. |date=April 2022 |title=Dietary isotopes of Madagascar's extinct megafauna reveal Holocene browsing and grazing guilds |journal=Biology Letters |language=en |volume=18 |issue=4 |pages=20220094 |doi=10.1098/rsbl.2022.0094 |issn=1744-957X |pmc=9006009 |pmid=35414222}}</ref> It has been suggested that ''Aepyornis'' straightened its legs and brought its torso into an erect position in order to browse higher vegetation.<ref name=":6" /> Some rainforest fruits with thick, highly sculptured [[endocarp]]s, such as that of the currently undispersed and highly threatened [[forest coconut]] palm (''Voanioala gerardii''), may have been adapted for passage through ratite guts and consumed by elephant birds, and the fruit of some palm species are indeed dark bluish-purple (e.g., ''[[Ravenea louvelii]]'' and ''[[Satranala decussilvae]]''), just like many cassowary-dispersed fruits, suggesting that they too may have been eaten by elephant birds.<ref>Dransfield, J. & Beentje, H. (1995)</ref> 

=== Growth and reproduction ===
Elephant birds are suggested to have grown in periodic spurts rather than having continuous growth.<ref name=":1">{{Cite journal |last1=Chinsamy |first1=Anusuya |last2=Angst |first2=Delphine |last3=Canoville |first3=Aurore |last4=Göhlich |first4=Ursula B |date=2020-06-01 |title=Bone histology yields insights into the biology of the extinct elephant birds (Aepyornithidae) from Madagascar |url=https://academic.oup.com/biolinnean/article/130/2/268/5815707 |journal=Biological Journal of the Linnean Society |language=en |volume=130 |issue=2 |pages=268–295 |doi=10.1093/biolinnean/blaa013 |issn=0024-4066|doi-access=free }}</ref> An embryonic skeleton of ''Aepyornis'' is known from an intact egg, around 80–90% of the way through incubation before it died. This skeleton shows that even at this early ontogenetic stage that the skeleton was robust, much more so than comparable hatchling ostriches or rheas,<ref>{{Cite journal |last1=Balanoff |first1=Amy M. |last2=Rowe |first2=Timothy |date=2007-12-12 |title=Osteological description of an embryonic skeleton of the extinct elephant bird, Aepyornis (Palaeognathae: Ratitae) |url=http://www.tandfonline.com/doi/abs/10.1671/0272-4634%282007%2927%5B1%3AODOAES%5D2.0.CO%3B2 |journal=Journal of Vertebrate Paleontology |language=en |volume=27 |issue=sup4 |pages=1–53 |doi=10.1671/0272-4634(2007)27[1:ODOAES]2.0.CO;2 |s2cid=85920479 |issn=0272-4634|url-access=subscription }}</ref> which may suggest that hatchlings were [[precocial]].<ref name=":1" />

The eggs of ''Aepyornis'' are the largest known for any [[amniote]], and have a volume of around {{Convert|5.6–13|L|USpt}}, a length of approximately {{Convert|26–40|cm|in}} and a width of {{Convert|19–25|cm|in}}.<ref name=":1" /> The largest ''Aepyornis'' eggs are on average {{Convert|3.3|mm|in|abbr=on|frac=64}} thick, with an estimated weight of approximately {{Convert|10.5|kg|lb}}.<ref name="Molecular exploration of fossil eggshell uncovers hidden lineage of giant extinct bird" /> Eggs of ''Mullerornis'' were much smaller, estimated to be only {{Convert|1.1|mm|in|abbr=on|frac=64}} thick, with a weight of about {{Convert|0.86|kg|lb}}.<ref name="Molecular exploration of fossil eggshell uncovers hidden lineage of giant extinct bird" /> The large size of elephant bird eggs means that they would have required substantial amounts of calcium, which is usually taken from a reservoir in the medullary bone in the femurs of female birds. Possible remnants of this tissue have been described from the femurs of ''A. maximus.''<ref name=":1" />{{gallery|Animals of the past (Fig. 16) (5985156816) (cropped).jpg|Eggs of ''Aepyornis'' (top left) chicken (bottom left) [[moa]] (right) and ostrich (centre)|Denis Bourez - Natural History Museum, London (8900355257).jpg|Elephant bird egg (far left, 1) in comparison to other eggs, including ostrich egg (centre, 3) and chicken egg (third from right, 6)|Mus Nat Hist Nat 25022013 Aepyornis egg.jpg|Well preserved egg showing porous surface|Oeuf d'Aepyornis maximus (musée des Douanes).JPG|Egg of ''Aepyornis'' standing upright|||.png||||||width=200|height=180|align=center}}

== Extinction ==
It is widely believed that the extinction of elephant birds was a result of human activity. The birds were initially widespread, occurring from the northern to the southern tip of [[Madagascar]].<ref name="Hawkins">Hawkins, A. F. A. & Goodman, S. M. (2003)</ref> The late Holocene also witnessed the extinction of other Malagasy animals, including several species of [[Malagasy hippopotamus]], two species of giant tortoise (''[[Aldabrachelys abrupta]]'' and ''[[Aldabrachelys grandidieri]]''),  the [[giant fossa]], over a dozen species of [[giant lemurs]], the aardvark-like animal ''[[Plesiorycteropus]],'' and the crocodile ''[[Voay]]''.''<ref name=":6" />'' Several elephant bird bones with incisions have been dated to approximately 10,000 BCE which some authors suggest are cut marks, which have been proposed as evidence of a long history of coexistence between elephant birds and humans;<ref name="Hansford2018">{{cite journal |last1=Hansford |first1=J. |last2=Wright |first2=P. C. |last3=Rasoamiaramanana |first3=A. |last4=Pérez |first4=V. R. |last5=Godfrey |first5=L. R. |last6=Errickson |first6=D. |last7=Thompson |first7=T. |last8=Turvey |first8=S. T. |year=2018 |title=Early Holocene human presence in Madagascar evidenced by exploitation of avian megafauna |url=http://discovery.ucl.ac.uk/10052571/35/Hansford_eaat6925.full.pdf |journal=Science Advances |volume=4 |issue=9 |pages=eaat6925 |bibcode=2018SciA....4.6925H |doi=10.1126/sciadv.aat6925 |pmc=6135541 |pmid=30214938}}</ref> however, these conclusions conflict with more commonly accepted evidence of a much shorter history of human presence on the island and remain controversial. The oldest securely dated evidence for humans on Madagascar dates to the mid-first millennium AD.<ref name=":4">{{Cite journal |last=Mitchell |first=Peter |date=2020-10-01 |title=Settling Madagascar: When Did People First Colonize the World's Largest Island? |url=https://www.tandfonline.com/doi/full/10.1080/15564894.2019.1582567 |journal=The Journal of Island and Coastal Archaeology |language=en |volume=15 |issue=4 |pages=576–595 |doi=10.1080/15564894.2019.1582567 |issn=1556-4894 |s2cid=195555955|url-access=subscription }}</ref>

A 2021 study suggested that elephant birds, along with the Malagasy hippopotamus species, became extinct in the interval 800–1050 CE (1150–900 years [[Before Present]]), based on the timing of the latest radiocarbon dates. The timing of the youngest radiocarbon dates co-incided with major environmental alteration across Madagascar by humans changing forest into grassland, probably for cattle [[pastoralism]], with the environmental change likely being induced by the use of fire. This reduction of forested area may have had cascade effects, like making elephant birds more likely to be encountered by hunters,<ref name=":2">{{Cite journal |last1=Hansford |first1=James P. |last2=Lister |first2=Adrian M. |last3=Weston |first3=Eleanor M. |last4=Turvey |first4=Samuel T. |date=July 2021 |title=Simultaneous extinction of Madagascar's megaherbivores correlates with late Holocene human-caused landscape transformation |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379121002031 |journal=Quaternary Science Reviews |language=en |volume=263 |pages=106996 |doi=10.1016/j.quascirev.2021.106996|bibcode=2021QSRv..26306996H |s2cid=236313083 }}</ref> though there is little evidence of human hunting of elephant birds. Humans may have utilized elephant bird eggs. Introduced diseases ([[Late Pleistocene extinctions#Disease|hyperdisease]]) have been proposed as a cause of extinction, but the plausibility for this is weakened due to the evidence of centuries of overlap between humans and elephant birds on Madagascar.<ref name=":6">{{Cite journal |last1=Clarke |first1=Simon J. |last2=Miller |first2=Gifford H. |last3=Fogel |first3=Marilyn L. |last4=Chivas |first4=Allan R. |last5=Murray-Wallace |first5=Colin V. |date=September 2006 |title=The amino acid and stable isotope biogeochemistry of elephant bird (Aepyornis) eggshells from southern Madagascar |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379106000758 |journal=Quaternary Science Reviews |language=en |volume=25 |issue=17–18 |pages=2343–2356 |doi=10.1016/j.quascirev.2006.02.001|bibcode=2006QSRv...25.2343C |url-access=subscription }}</ref>

== See also ==
{{Portal|Paleontology}}
* [[Late Quaternary prehistoric birds]]
* [[Holocene extinction]]

== References ==
{{Reflist|30em}}

== Further reading ==
{{Refbegin}}
* {{cite web |url=http://news.bbc.co.uk/2/hi/in_pictures/7963484.stm |access-date=26 March 2009|work= BBC News|title=One minute world news |date=2009-03-25}}
* {{cite serial| title= BBC-2 Presents: Attenborough and the Giant Egg |url=http://www.bbc.co.uk/programmes/b00z6dsg#broadcasts |credits=Presenter: David Attenborough; Director: Sally Thomson; Producer: Sally Thomson; Executive Producer: Michael Gunton |network=BBC |station=BBC Two |air-date=2 March 2011}}
* {{cite web |url=http://sn2000.taxonomy.nl/Taxonomicon/TaxonTree.aspx?id=51284 |title=The Taxonomicon : Taxon: Order Aepyornithiformes |access-date=21 January 2010 |last=Brands |first=Sheila J. |year=1989 |publisher=Universal Taxonomic Services |location=Zwaag, Netherlands |archive-url=https://web.archive.org/web/20080421213745/http://sn2000.taxonomy.nl/Taxonomicon/TaxonTree.aspx?id=51284 |archive-date=2008-04-21 |url-status=dead}}
* {{cite web |url=http://www.taxonomy.nl/Main/Classification/51286.htm |title=Systema Naturae 2000 / Classification, Genus Aepyornis |access-date=4 February 2009 |last=Brands(a) |first=Sheila |date=14 August 2008 |work=Project: The Taxonomicon |url-status=dead |archive-url=https://web.archive.org/web/20090303044044/http://www.taxonomy.nl/Main/Classification/51286.htm |archive-date=2009-03-03}}
* {{cite journal |last1=Brodkorb |first1=Pierce |year=1963 |title= Catalogue of Fossil Birds Part 1 (Archaeopterygiformes through Ardeiformes)|journal=Bulletin of the Florida State Museum, Biological Sciences |volume=7 |issue=4 |pages=179–293 |publisher=University of Florida |location= Gainesville, FL |url=http://ufdc.ufl.edu/UF00001514/00001/98j }}
* {{cite journal |last1=Cooper |first1=A. |last2=Lalueza-Fox |first2=C. |last3=Anderson |first3=S. |last4=Rambaut |first4=A. |last5=Austin |first5=J. |last6=Ward |first6=R. |title=Complete Mitochondrial Genome Sequences of Two Extinct Moas Clarify Ratite Evolution |journal=Nature |volume=409 |pages=704–707 |date=2001-02-08 |doi=10.1038/35055536 |pmid=11217857 |issue=6821|bibcode=2001Natur.409..704C |s2cid=4430050}}
* {{cite encyclopedia |last=Davies|first=S. J. J. F.|editor=Hutchins, Michael|encyclopedia=Grzimek's Animal Life Encyclopedia, Vol. 8: Birds I Tinamous and Ratites to Hoatzins |title=Elephant birds |edition=2nd |year=2003 |publisher=Gale Group |location=Farmington Hills, MI|isbn=978-0-7876-5784-0 |pages=103–104}}
* {{cite book |last1=Dransfield|first1= John |last2=Beentje |first2= Henk |title=The Palms of Madagascar |year=1995 |publisher=Royal Botanic Gardens |location=Kew, Victoria, Australia |isbn=978-0-947643-82-9 }}
* {{cite book |last1=Flacourt |first1=Etienne de. |last2=Allibert |first2=Claude |title=Histoire de la grande île de Madagascar |year=2007 |publisher=Karthala |location=Paris, FR |language=fr |isbn=978-2-84586-582-2 |url-access=registration |url=https://archive.org/details/histoiredelagran0000flac }}
* {{cite journal |last1=Goodman|first1=Steven M. |year=1994 |title=Description of a new species of subfossil eagle from Madagascar: ''Stephanoaetus'' (Aves: Falconiformes) from the deposits of Ampasambazimba |url=https://archive.org/details/biostor-81217|journal=Proceedings of the Biological Society of Washington |issue=107 |pages=421–428 }}
* {{cite book |last1=Goodman|first1=S. M. |last2=Rakotozafy |first2=L. M. A. |editor1-first=S. M. |editor1-last=Goodman|editor2-first=B. D. |editor2-last=Patterson|title=Natural Change and Human Impact in Madagascar |year=1997 |publisher=Smithsonian Institution Press |location=Washington, DC |isbn=978-1-56098-683-6 |pages=257–279 |chapter=Subfossil birds from coastal sites in western and southwestern Madagascar }}
* {{cite book |last1=Hawkins|first1=A. F. A. |last2=Goodman |first2=S. M. |editor1-first=S. M. |editor1-last=Goodman |editor2-first= J. P.|editor2-last=Benstead |title=The Natural History of Madagascar |year=2003 |publisher=University of Chicago Press |isbn=978-0-226-30307-9 |pages=1026–1029 }}
* {{cite book |last1=Hay|first1=W. W. | last2=DeConto|first2=R. M. |last3=Wold|first3=C. N. |last4=Wilson |first4=K. M.|last5=Voigt|first5=S.|editor1-first=E. |editor1-last=Barrera |editor2-first= C. C. |editor2-last=Johnson |title=Evolution of the Cretaceous Ocean Climate System |year=1999 |publisher=Geological Society of America |location=Boulder, CO |isbn=978-0-8137-2332-7 |pages=1–47 |chapter=Alternative global Cretaceous paleogeography }}
* {{cite web| url=http://avibase.bsc-eoc.org/avibase.jsp?pg=search&fam=4.2&lang=EN | title=Aepyornithidae | access-date=4 February 2009 |last=LePage |first=Dennis |year=2008 |work=Avibase, the World Bird Database}}
* {{cite book |last1=MacPhee|first1=R. D. E. |last2=Marx |first2=P. A. |editor1-first=S. M. |editor1-last=Goodman |editor2-first= B. D. |editor2-last=Patterson|title=Natural Change and Human Impact in Madagascar |year=1997 |publisher=Smithsonian Institution Press |location=Washington, DC |pages=169–217 |chapter=The 40,000 year plague: humans, hyperdisease, and first-contact extinctions }}
* {{cite book |last1=Megiser|first1=H. |title=Warhafftige ... so wol Historische als Chorographische Beschreibung der ... Insul Madagascar, sonsten S. Laurentii genandt (etc.) |year=1623 |publisher=Groß |location=Leipzig }}
* {{cite journal |last1=Mlíkovsky|first1=J. |year=2003 |title=Eggs of extinct aepyornithids (Aves: Aepyornithidae) of Madagascar: size and taxonomic identity |journal=Sylvia |volume=39 |pages=133–138 }}
* {{cite book |last1=Pearson|first1=Mike Parker |last2=Godden |first2=K. |title=In search of the Red Slave: Shipwreck and Captivity in Madagascar |year=2002 |publisher=The History Press |location=Stroud, Gloucestershire |isbn=978-0-7509-2938-7 }}
* {{cite journal |last1=van Tuinen|first1=Marcel | last2=Sibley|first2=Charles G. |last3=Hedges|first3=S. Blair |year=1998 |title=Phylogeny and Biogeography of Ratite Birds Inferred from DNA Sequences of the Mitochondrial Ribosomal Genes |journal=Molecular Biology and Evolution |volume=15 |issue=4 |pages=370–376 |pmid=9549088 |doi= 10.1093/oxfordjournals.molbev.a025933|doi-access=free }}
* {{cite journal |last1=Yoder|first1=Anne D. | last2=Nowak|first2=Michael D. |year=2006 |title= Has Vicariance or Dispersal Been the Predominant Biogeographic Force in Madagascar? Only Time Will Tell |journal=Annual Review of Ecology, Evolution, and Systematics |volume=37 |pages=405–431 |doi=10.1146/annurev.ecolsys.37.091305.110239 }}
{{Refend}}

== External links ==
* [http://digimorph.org/specimens/Aepyornis_maximus/ Digimorph.org]
* [https://web.archive.org/web/20060307211250/http://www.ornitaxa.com/SM/Fossil/FossilAepyorn.htm Fossil Aepyornithidae]
* [https://web.archive.org/web/20110606164414/http://www.shef.ac.uk/archaeology/research/madagascar/madagascar4.html University of Sheffield] excavations of elephant bird eggshells
* [https://web.archive.org/web/20010411230801/http://www.geocities.com/vorompatra/index.html Vorompatra info]
* [https://web.archive.org/web/20100323070258/http://www.independent.co.uk/news/science/archaeology/fossilised-egg-gives-clue-to-fate-of-ancient-birds-1919138.html Fossilised egg gives clue to fate of ancient birds The Independent]
* [https://www.bbc.co.uk/news/entertainment-arts-21960778# Giant egg from extinct elephant bird up for auction]

{{Palaeognathae|N.}}
{{Taxonbar|from=Q214508}}
{{Authority control}}

[[Category:Elephant birds]]
[[Category:Novaeratitae]]
[[Category:Prehistoric animals of Madagascar]]
[[Category:Pleistocene first appearances]]
[[Category:Species made extinct by human activities]]
[[Category:Holocene extinctions]]
[[Category:Taxa named by Charles Lucien Bonaparte]]