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Elephant birds are members of the extinct
ratite A ratite () is any of a diverse group of flightless, large, long-necked, and long-legged birds of the infraclass Palaeognathae. Kiwi, the exception, are much smaller and shorter-legged and are the only nocturnal extant ratites. The systematics ...
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Aepyornithidae, made up of flightless birds that once lived on the island of
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
. They are thought to have become extinct around 1000-1200 CE, probably as a result of human activity. Elephant birds comprised the genera ''
Mullerornis ''Mullerornis'' is a genus of extinct elephant birds (Aepyornithidae) of Madagascar. Description ''Mullerornis'' is smaller than the more well-known ''Aepyornis''. A bone possibly belonging to ''Mullerornis'' has been radiocarbon dated to about ...
'', '' Vorombe'' and '' Aepyornis''. While they were in close geographical proximity to the ostrich, their closest living relatives are kiwi (found only in New Zealand), suggesting that
ratite A ratite () is any of a diverse group of flightless, large, long-necked, and long-legged birds of the infraclass Palaeognathae. Kiwi, the exception, are much smaller and shorter-legged and are the only nocturnal extant ratites. The systematics ...
s did not diversify by
vicariance Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
during the breakup of Gondwana but instead evolved from ancestors that dispersed more recently by flying. In September 2018, scientists determined that ''Vorombe titan'' reached weights of and stood tall, making it the world's largest and heaviest
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
, slightly larger than the much older '' Dromornis stirtoni''. Other members of the family were also very large, exhibiting the phenomenon of island gigantism.


Description

Elephant birds have been extinct since at least the 17th century.
Étienne de Flacourt Étienne de Flacourt (1607–1660) was a French governor of Madagascar, born in Orléans in 1607. He was named governor of Madagascar by the French East India Company in 1648. Flacourt restored order among the French soldiers, who had mutinied. ...
, 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 – 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."
Marco Polo Marco Polo (, , ; 8 January 1324) was a Venetian merchant, explorer and writer who travelled through Asia along the Silk Road between 1271 and 1295. His travels are recorded in '' The Travels of Marco Polo'' (also known as ''Book of the Marv ...
also mentioned hearing stories of very large birds during his journey to the East during the late 13th century. These accounts are today believed to describe elephant birds. Between 1830 and 1840 European travelers in Madagascar saw giant eggs and egg shells. English observers were more willing to believe the accounts of giant birds and eggs because they knew of the moa in New Zealand. In 1851 the
French Academy of Sciences The French Academy of Sciences (French: ''Académie des sciences'') is a learned society, founded in 1666 by Louis XIV at the suggestion of Jean-Baptiste Colbert, to encourage and protect the spirit of French scientific research. It was at th ...
received three eggs and some bone fragments. In some cases the eggs have a length up to , the largest type of bird egg ever found.Mlíkovsky, J. (2003) The egg weighed about . The egg volume is about 160 times greater than that of a chicken egg.Hawkins, A. F. A. & Goodman, S. M. (2003) ''Aepyornis'' is believed to have been more than tall and weighed perhaps in the range of .Davies, S. J. J. F. (2003)Anderson, J. F., Rahn, H., & Prange, H. D. (1979). ''Scaling of supportive tissue mass''. Quarterly Review of Biology, 139–148. In September 2018, scientists reported that ''Vorombe titan'' reached weights of , and based on a fragmentary femur, possibly up to , making it the world's largest bird. Only the much older species '' Dromornis stirtoni'' from Australia rivals it in size among known fossil birds. In the same report, the upper weight limits for ''A. maximus'' and ''D. stirtoni'' were revised to 540 and 730 kg, respectively.


Species

Up to ten or eleven species in the genus ''Aepyornis'' have been described,Brodkorb, Pierce (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''. But recent work by Hansford & Turvey 2018 has restricted the number of aepyornithid species to four, with two in ''Aepyornis'', one in ''Mullerornis'', and one in ''Vorombe''. * Order Aepyornithiformes Newton 1884 epyornithes Newton 1884ref name="Brodkorb" /> ** Family Aepyornithidae (Bonaparte 1853) epyornithinae Bonaparte 1853*** Genus '' Aepyornis'' Geoffroy Saint-Hilaire 1850 **** ''Aepyornis hildebrandti'' Burckhardt 1893 (Hildebrandt's elephant-bird) **** ''Aepyornis maximus'' Geoffroy Saint-Hilaire 1851 (Giant elephant-bird) *** Genus ''
Mullerornis ''Mullerornis'' is a genus of extinct elephant birds (Aepyornithidae) of Madagascar. Description ''Mullerornis'' is smaller than the more well-known ''Aepyornis''. A bone possibly belonging to ''Mullerornis'' has been radiocarbon dated to about ...
'' Milne-Edwards & Grandidier 1894 **** ''Mullerornis modestus'' (Milne-Edwards & Grandidier 1869) Hansford & Turvey 2018Brands (a), S. (2008) (Betsile/robust elephant-bird) *** Genus '' Vorombe'' Hansford & Turvey 2018 **** ''Vorombe titan'' (Andrews 1894) Hansford & Turvey 2018 Several ratites outside of Madagascar have been posited as "aepyornithid"-like and could potentially make this clade considerably more speciose. These include '' Eremopezus'' from the
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
of North Africa, unnamed Canary Island remains and several Neogene taxa in Eurasia.


Etymology

''Aepyornis maximus'' is commonly known as the 'elephant bird', a term that apparently originated from
Marco Polo Marco Polo (, , ; 8 January 1324) was a Venetian merchant, explorer and writer who travelled through Asia along the Silk Road between 1271 and 1295. His travels are recorded in '' The Travels of Marco Polo'' (also known as ''Book of the Marv ...
's account of the rukh in 1298, although he was apparently referring to an eagle-like bird strong enough to "seize an elephant with its talons". Sightings of eggs of elephant birds by sailors (e.g. text on the Fra Mauro map of 1467–69, if not attributable to ostriches) could also have been erroneously attributed to a giant raptor from Madagascar. The legend of the roc could also have originated from sightings of such a giant subfossil eagle related to the African crowned eagle, which has been described in the genus ''
Stephanoaetus ''Stephanoaetus'' is a genus of very large birds of prey from Sub-Saharan Africa and Madagascar. Only one of the two known species is extant. Species * Crowned eagle or crowned hawk-eagle (''Stephanoaetus coronatus''). * † Malagasy crowned ea ...
'' from Madagascar, being large enough to carry off large
primates Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
; today, lemurs still retain a fear of aerial predators such as these. Another might be the perception of ratites retaining neotenic features and thus being mistaken for enormous chicks of a presumably more massive bird. The ancient Malagasy name for the bird is vorompatra, meaning "bird of the Ampatres". The Ampatres are today known as the Androy region of southern Madagascar.


Taxonomy and biogeography

Like the ostrich, rhea,
cassowary Cassowaries ( tpi, muruk, id, kasuari) are flightless birds of the genus ''Casuarius'' in the order Casuariiformes. They are classified as ratites (flightless birds without a keel on their sternum bones) and are native to the tropical ...
, emu, kiwi and extinct moa, ''Mullerornis'' and ''Aepyornis'' were ratites; they could not fly, and their breast bones had no keel. Because Madagascar and
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
separated before the ratite lineage arose, ''Aepyornis'' has been thought to have dispersed and become flightless and gigantic '' in situ''. More recently, it has been deduced from DNA sequence comparisons that the closest living relatives of elephant birds are New Zealand kiwi, though they are not particularly closely related, being estimated to have diverged from each other 54 million years ago. Elephant birds are actually part of the mid- Cenozoic Australian ratite radiation; their ancestors flew across the Indian Ocean well after Gondwana broke apart. The existence of possible flying palaeognaths in the Miocene such as '' Proapteryx'' further supports the view that ratites did not diversify in response to
vicariance Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
. 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. Molecular clock estimates from complete mitochondrial genomes obtained from ''Aepyornis'' eggshells suggest that ''Aepyornis'' and ''Mullerornis'' diverged from each other during the mid
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
around 27 million years ago, suggesting elephant birds must have been present on Madagascar before this time. Claims of findings of "aepyornithid" egg remains on the eastern
Canary Islands The Canary Islands (; es, Canarias, ), also known informally as the Canaries, are a Spanish autonomous community and archipelago in the Atlantic Ocean, in Macaronesia. At their closest point to the African mainland, they are west of Mo ...
, if valid, would represent a major biogeographical enigma.Sauer and Rothe, 1972 These islands are not thought to have been connected to mainland Africa when elephant birds were alive. There is no indication that elephant birds evolved outside Madagascar, and today, the Canary Island eggshells are considered to belong to extinct North African birds that may not have been ratites (possibly '' Eremopezus''/'' Psammornis'', or even Pelagornithidae, prehistoric seabirds of immense size). Various "aepyornithid-like" eggs and bones occur in
Paleogene The Paleogene ( ; British English, also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period, geologic period and system that spans 43 million years from the end of the Cretaceous Period million yea ...
and
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recent" ...
deposits in
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
and
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located enti ...
. 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 egg); both have been identified as ''Aepyornis maximus'' rather than '' Genyornis''. 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 Penguins (order (biology), order List of Sphenisciformes by population, Sphenisciformes , family (biology), family Spheniscidae ) are a group of Water bird, aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: on ...
eggs that washed ashore on Western Australia but originated in the
Kerguelen Islands The Kerguelen Islands ( or ; in French commonly ' but officially ', ), also known as the Desolation Islands (' in French), are a group of islands in the sub-Antarctic constituting one of the two exposed parts of the Kerguelen Plateau, a large ...
, and an ostrich egg found floating in the Timor Sea in the early 1990s.


Biology

Examination of brain endocasts has shown that both ''A. maximus'' and ''A. hildebrandi'' had greatly reduced optic lobes, similar to those of their closest living relatives, the kiwis, and consistent with a similar
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
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 bulbs than ''A. hildebrandi'', suggesting that the former occupied forested habitats where the sense of smell is more useful while the latter occupied open habitats.


Diet

Because there is no
rainforest Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainfores ...
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
record in Madagascar, it is not known for certain if there were species adapted to dense forest dwelling, like the
cassowary Cassowaries ( tpi, muruk, id, kasuari) are flightless birds of the genus ''Casuarius'' in the order Casuariiformes. They are classified as ratites (flightless birds without a keel on their sternum bones) and are native to the tropical ...
in Australia and
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; id, Papua, or , historically ) is the world's second-largest island with an area of . Located in Oceania in the southwestern Pacific Ocean, the island is separated from Australia by the wide Torr ...
today. However, some rainforest fruits with thick, highly sculptured endocarps, such as that of the currently undispersed and highly threatened forest coconut palm ''
Voanioala gerardii ''Voanioala gerardii'', commonly known as the forest coconut, is a species of flowering plant in the family Arecaceae. It is a relative of the coconut, and is generally regarded as monotypic within the genus ''Voanioala''. However, a team of gen ...
'', may have been adapted for passage through ratite guts, and the fruit of some palm species are indeed dark bluish purple (e.g. ''
Ravenea louvelii ''Ravenea louvelii'' is a species of flowering plant in the Arecaceae family. It is a palm endemic to Madagascar, where it is known from a single location. There are fewer than 10 individuals left. They are believed to be in Andasibe-Mantadia N ...
'' and ''
Satranala decussilvae ''Satranala decussilvae'' is a species of flowering plant in the Arecaceae family. It is a palm endemic to Madagascar. It is the only species in the genus ''Satranala'', and is threatened by habitat loss Habitat destruction (also termed hab ...
''), just like many cassowary-dispersed fruits.


Reproduction

Occasionally subfossil eggs are found intact. The National Geographic Society in Washington, D.C. holds a specimen of an ''Aepyornis'' egg which was given to Luis Marden in 1967. The specimen is intact and contains the skeleton of the unhatched bird. The Denver Museum of Nature and Science in Colorado holds two intact eggs, one of which is currently on display. Another giant ''Aepyornis'' egg is on display at the Harvard Museum of Natural History in Cambridge, Massachusetts, and a complete, unbroken egg is held at
Leeds Discovery Centre Leeds Discovery Centre is the purpose-built display storage facility built for Leeds Museums & Galleries in 2007. It was funded by Leeds City Council and the National Lottery Heritage Fund. The facility stores over one million objects in climate ...
in the UK. A cast of the egg is preserved at the Grant Museum of Zoology at
London University The University of London (UoL; abbreviated as Lond or more rarely Londin in post-nominals) is a federal public research university located in London, England, United Kingdom. The university was established by royal charter in 1836 as a degr ...
. An intact specimen also exists in
Kuleli Military High School Kuleli Military High School was the oldest military high school in Turkey, located in Çengelköy, Istanbul, on the Asian shore of the Bosphorus strait. It was founded on September 21, 1845, by Ottoman Sultan Abdülmecid I. After the 2016 Turki ...
Museum, Istanbul, Turkey.
David Attenborough Sir David Frederick Attenborough (; born 8 May 1926) is an English broadcaster, biologist, natural historian and author. He is best known for writing and presenting, in conjunction with the BBC Natural History Unit, the nine natural histor ...
owned an almost complete eggshell, dating from 600 to 700 AD, which he pieced together from fragments that were given to him while making his 1961 BBC series '' Zoo Quest to Madagascar''. In March 2011, the BBC broadcast the 60-minute documentary ''Attenborough and the Giant Egg'', presented by Attenborough, about his personal scientific quest to discover the secrets of the elephant bird and its egg. A complete eggshell is also available in the collection of the University of Wrocław Museum of Natural History. There is also an intact specimen of an elephant bird's egg (contrasted with the eggs from other bird species, including a hummingbird's) on display at the Delaware Museum of Natural History, just outside Wilmington,
Delaware Delaware ( ) is a state in the Mid-Atlantic region of the United States, bordering Maryland to its south and west; Pennsylvania to its north; and New Jersey and the Atlantic Ocean to its east. The state takes its name from the adjacen ...
, US, and another in the Natural History Museum, London. The Melbourne Museum has two ''Aepyornis'' eggs. The first was acquired for £100 by Professor Frederick McCoy in June 1862, and is an intact example. In 1950 it was subjected to radiological examination, which revealed no traces of embryonic material. A second, side- blown ''Aepyornis'' egg was acquired at a later date. The Western Foundation of Vertebrate Zoology, with one of the world's largest collections of avian eggs, has seven ''Aepyornis'' egg specimens. A specimen is also held by the science department at
Stowe School , motto_translation = I stand firm and I stand first , established = , closed = , type = Public schoolIndependent school, day & boarding , religion = Church of England , president = , head_label = Headmaster ...
in Buckinghamshire, UK. In the collections of the department of geology at the Field Museum of Natural History in
Chicago (''City in a Garden''); I Will , image_map = , map_caption = Interactive Map of Chicago , coordinates = , coordinates_footnotes = , subdivision_type = List of sovereign states, Count ...
there is a complete, side-blown egg collected, in about 1917, by Rev. Peter A. Bjelde. A specimen of egg is also held at Regional Museum of Natural History, Bhubaneswar which was donated by former Indian Ambassador Abasar Beuria and his wife Tripti Beuria from their collection. In April 2013 a specimen was sold at Christie's in London for £66,675 The pre-sale estimate had been "more than $45,000". In April 2018, staff at the
Buffalo Museum of Science The Buffalo Museum of Science is a science museum located at Martin Luther King Jr. Park in Buffalo, New York, United States, northeast of the downtown district, near the Kensington Expressway. The historic building was designed by August Es ...
radiographed what had long been thought to be a model of an elephant bird egg. The cream-colored object, measuring in length and in circumference and weighing over , was found to be a mislabeled real egg (the museum does also have such a model in its collections). Two eggs have been discovered on the Western Australia coast, one discovered in the 1930s and another in 1992. A study concluded that the egg found at Cervantes was 2000 years old and had likely drifted across the Indian Ocean.


Relationship with humans


Extinction

It is widely believed that the extinction of ''Aepyornis'' was a result of human activity. The birds were initially widespread, occurring from the northern to the southern tip of
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
. One theory states that humans hunted the elephant birds to extinction in a very short time for such a large landmass (the blitzkrieg hypothesis). There is indeed evidence that they were hunted and their preferred habitats destroyed. Eggs may have been particularly vulnerable. 19th-century travelers saw eggshells used as bowls, and a recent archaeological study found remains of eggshells among the remains of human fires,Pearson and Godden (2002) suggesting that the eggs regularly provided meals for entire families. The exact period when they died out is also not certain; tales of these giant birds may have persisted for centuries in folk memory. There is archaeological evidence of ''Aepyornis'' from a radiocarbon-dated bone at 1880 ± 70  BP (approximately 120  CE) with signs of butchering, and on the basis of radiocarbon dating of shells, about 1000 BP (approximately 1000 AD). An alternative theory is that the extinction was a secondary effect of human impact resulting from transfer of hyperdiseases from human commensals such as
chicken The chicken (''Gallus gallus domesticus'') is a domestication, domesticated junglefowl species, with attributes of wild species such as the grey junglefowl, grey and the Ceylon junglefowl that are originally from Southeastern Asia. Rooster ...
s and guineafowl. The bones of these domesticated fowl have been found in
subfossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
sites in the island (MacPhee and Marx, 1997: 188), such as Ambolisatra (Madagascar), where ''Mullerornis'' sp. and ''Aepyornis maximus'' have been reported. Also reported by these authors, ratite remains have been found in west and south west Madagascar, at Belo-sur-Mer (''A. medius'', ''Mullerornis rudis''), Bemafandry (''M. agilis'') and Lamboharana (''Mullerornis'' sp.). Several elephant bird bones with tool marks have been dated to approximately 10,000 BC and interpreted as evidence of a long history of coexistence between elephant birds and humans; however, these conclusions conflict with more commonly accepted evidence of a much shorter history of human presence on the island and remain controversial.


In art and literature

* The roc (rukh) is known from Sindbad the Sailor's encounter with one in ''
One Thousand and One Nights ''One Thousand and One Nights'' ( ar, أَلْفُ لَيْلَةٍ وَلَيْلَةٌ, italic=yes, ) is a collection of Middle Eastern folk tales compiled in Arabic during the Islamic Golden Age. It is often known in English as the ''Arabian ...
''. Some scholars think the roc is a distorted account of ''Aepyornis''. Historical evidence for this can be found in Megiser (1623). * H. G. Wells wrote a short story " Æpyornis Island" (1894) about the bird. It was first collected in '' The Stolen Bacillus and Other Incidents'' (1895). * Wildlife artist
Walton Ford Walton Ford (born 1960 in Larchmont, New York) is an American artist who makes paintings and prints in the style of naturalist illustrations, often depicting extinct species. Each of his paintings is a meticulous, realistic study in flora and fa ...
created a painting called ''Madagascar'' about the elephant bird in 2002.


See also

* Late Quaternary prehistoric birds * New World Pleistocene extinctions * Pleistocene megafauna


Notes


References

* * * * * * * * * * * * * * * * * * * * *


External links


Digimorph.org




excavations of elephant bird eggshells



* ttps://www.bbc.co.uk/news/entertainment-arts-21960778# Giant egg from extinct elephant bird up for auction {{Authority control Elephant birds Prehistoric animals of Madagascar Pleistocene first appearances Holocene extinctions Taxa named by Charles Lucien Bonaparte Roc (mythology)