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Elephant birds are members of the
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
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 Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Aepyornithidae, made up of
flightless bird Flightless birds are birds that through evolution lost the ability to fly. There are over 60 extant species, including the well known ratites (ostriches, emu, cassowaries, rheas, and kiwi) and penguins. The smallest flightless bird is the In ...
s 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 Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
''
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 abo ...
'', ''
Vorombe ''Vorombe'' is one of three genera of elephant birds, an extinct family of large ratite birds endemic to Madagascar. Originally considered to be large ''Aepyornis'' specimens, it is now thought ''Vorombe'' are the largest and heaviest birds kno ...
'' and ''
Aepyornis ''Aepyornis'' is a genus of aepyornithid, one of three genera of ratite birds endemic to Madagascar until their extinction sometime around 1000 CE. The species ''A. maximus'' weighed up to , and until recently was regarded as the largest known ...
''. While they were in close geographical proximity to the
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There are ...
, their closest living relatives are
kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), a nickname for New Zealanders * Kiwifruit, an edible berry * Kiwi dollar or New Zealand dollar, a unit of currency Kiwi or KIWI may also refe ...
(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 Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
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 lightweigh ...
, slightly larger than the much older ''
Dromornis stirtoni ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, if ...
''. Other members of the family were also very large, exhibiting the phenomenon of
island gigantism Island gigantism, or insular gigantism, is a biological phenomenon in which the size of an animal species isolated on an island increases dramatically in comparison to its mainland relatives. Island gigantism is one aspect of the more general "i ...
.


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 Moa are extinct giant flightless birds native to New Zealand. The term has also come to be used for chicken in many Polynesian cultures and is found in the names of many chicken recipes, such as Kale moa and Moa Samoa. Moa or MOA may also refe ...
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 of France, Louis XIV at the suggestion of Jean-Baptiste Colbert, to encourage and protect the spirit of French Scientific me ...
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 ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, if ...
'' 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 ''Aepyornis'' is a genus of aepyornithid, one of three genera of ratite birds endemic to Madagascar until their extinction sometime around 1000 CE. The species ''A. maximus'' weighed up to , and until recently was regarded as the largest known ...
'' Geoffroy Saint-Hilaire 1850 **** ''Aepyornis hildebrandti''
Burckhardt Burckhardt, or (de) Bourcard in French, is a family of the Basel patriciate, descended from Christoph (Stoffel) Burckhardt (1490–1578), a merchant in cloth and silk originally from Münstertal, Black Forest, who received Basel citizenship i ...
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 abo ...
'' Milne-Edwards & Grandidier 1894 **** ''Mullerornis modestus'' (Milne-Edwards & Grandidier 1869) Hansford & Turvey 2018Brands (a), S. (2008) (Betsile/robust elephant-bird) *** Genus ''
Vorombe ''Vorombe'' is one of three genera of elephant birds, an extinct family of large ratite birds endemic to Madagascar. Originally considered to be large ''Aepyornis'' specimens, it is now thought ''Vorombe'' are the largest and heaviest birds kno ...
'' 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 ''Eremopezus'' is a prehistoric bird genus, possibly a palaeognath. It is known only from the fossil remains of a single species, the huge and presumably flightless ''Eremopezus eocaenus''. This was found in Upper Eocene Jebel Qatrani Formation ...
'' from the
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
of
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
, unnamed
Canary Island 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 Morocco ...
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 The Fra Mauro map is a map of the world made around 1450 by the Venetian cartographer Fra Mauro, which is “considered the greatest memorial of medieval cartography." It is a circular planisphere drawn on parchment and set in a wooden frame ...
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 The crowned eagle, also known as the African crowned eagle or the crowned hawk-eagle (''Stephanoaetus coronatus''), is a large bird of prey found in sub-Saharan Africa; in Southern Africa it is restricted to eastern areas.Sinclair & Ryan (2003) ...
, 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,
lemur Lemurs ( ) (from Latin ''lemures'' – ghosts or spirits) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea (), divided into 8 Family (biology), families and consisting of 15 genera and around 100 exist ...
s still retain a fear of aerial predators such as these. Another might be the perception of ratites retaining
neotenic Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared ...
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 Androy is the most southerly region of Madagascar. It covers an area of 19,540 km2, and had a population of 903,376 inhabitants in 2018. The administrative capital is Ambovombe-Androy, and the chief administrator is Michael Andrianirina (sinc ...
region of southern Madagascar.


Taxonomy and biogeography

Like the
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There are ...
, 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 forest ...
,
emu The emu () (''Dromaius novaehollandiae'') is the second-tallest living bird after its ratite relative the ostrich. It is endemic to Australia where it is the largest native bird and the only extant member of the genus ''Dromaius''. The emu' ...
,
kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), a nickname for New Zealanders * Kiwifruit, an edible berry * Kiwi dollar or New Zealand dollar, a unit of currency Kiwi or KIWI may also refe ...
and extinct
moa Moa are extinct giant flightless birds native to New Zealand. The term has also come to be used for chicken in many Polynesian cultures and is found in the names of many chicken recipes, such as Kale moa and Moa Samoa. Moa or MOA may also refe ...
, ''Mullerornis'' and ''Aepyornis'' were ratites; they could not fly, and their breast bones had no
keel The keel is the bottom-most longitudinal structural element on a vessel. On some sailboats, it may have a hydrodynamic and counterbalancing purpose, as well. As the laying down of the keel is the initial step in the construction of a ship, in Br ...
. 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 Lineage may refer to: Science * Lineage (anthropology), a group that can demonstrate its common descent from an apical ancestor or a direct line of descent from an ancestor * Lineage (evolution), a temporal sequence of individuals, populati ...
arose, ''Aepyornis'' has been thought to have dispersed and become flightless and gigantic ''
in situ ''In situ'' (; often not italicized in English) is a Latin phrase that translates literally to "on site" or "in position." It can mean "locally", "on site", "on the premises", or "in place" to describe where an event takes place and is used in ...
''. More recently, it has been deduced from DNA sequence comparisons that the closest living relatives of elephant birds are New Zealand
kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), a nickname for New Zealanders * Kiwifruit, an edible berry * Kiwi dollar or New Zealand dollar, a unit of currency Kiwi or KIWI may also refe ...
, 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 The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
Australian ratite radiation; their ancestors flew across the Indian Ocean well after
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
broke apart. The existence of possible flying palaeognaths in the Miocene such as ''
Proapteryx ''Proapteryx micromeros'' is an extinct kiwi known from the 16–19 million-year-old early Miocene sediments of the St Bathans Fauna of Otago, New Zealand. Features ''P. micromeros'' is considerably smaller than modern kiwis, weighing around 23 ...
'' 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 Continental drift is the hypothesis that the Earth's continents have moved over geologic time relative to each other, thus appearing to have "drifted" across the ocean bed. The idea of continental drift has been subsumed into the science of pla ...
. Madagascar has a notoriously poor Cenozoic terrestrial fossil record, with essentially no fossils between the end of the Cretaceous (
Maevarano Formation The Maevarano Formation is a Late Cretaceous sedimentary rock formation found in the Mahajanga Province of northwestern Madagascar. It is most likely Maastrichtian in age, and records a seasonal, semiarid environment with rivers that had greatly v ...
) 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 Morocc ...
, if valid, would represent a major
biogeographical Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, i ...
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 Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
n birds that may not have been ratites (possibly ''
Eremopezus ''Eremopezus'' is a prehistoric bird genus, possibly a palaeognath. It is known only from the fossil remains of a single species, the huge and presumably flightless ''Eremopezus eocaenus''. This was found in Upper Eocene Jebel Qatrani Formation ...
''/'' Psammornis'', or even
Pelagornithidae The Pelagornithidae, commonly called pelagornithids, pseudodontorns, bony-toothed birds, false-toothed birds or pseudotooth birds, are a prehistoric family of large seabirds. Their fossil remains have been found all over the world in rocks dating ...
, prehistoric
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
s 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 recen ...
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 Continent#Subcontinents, subcontinent of Eurasia ...
. Two whole eggs have been found in dune deposits in southern
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to th ...
, one in the 1930s (the Scott River egg) and one in 1992 (the
Cervantes Miguel de Cervantes Saavedra (; 29 September 1547 (assumed) – 22 April 1616 Old Style and New Style dates, NS) was an Early Modern Spanish writer widely regarded as the greatest writer in the Spanish language and one of the world's pre-emin ...
egg); both have been identified as ''Aepyornis maximus'' rather than ''
Genyornis ''Genyornis newtoni'', also known as thunder bird and mihirung paringmal (meaning "giant bird"), is an extinct species of large, flightless bird that lived in Australia during the Pleistocene epoch until around 50,000 years ago. Over two met ...
''. It is hypothesized that the eggs floated from Madagascar to Australia on the
Antarctic Circumpolar Current The Antarctic Circumpolar Current (ACC) is an ocean current that flows clockwise (as seen from the South Pole) from west to east around Antarctica. An alternative name for the ACC is the West Wind Drift. The ACC is the dominant circulation feat ...
. 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 The egg of the ostrich (genus ''Struthio'') is the largest of any living bird. The shell has a long history of use by humans as a container and for decorative artwork. The eggs are not commonly eaten. Biology The female common ostrich lays her f ...
found floating in the
Timor Sea The Timor Sea ( id, Laut Timor, pt, Mar de Timor, tet, Tasi Mane or ) is a relatively shallow sea bounded to the north by the island of Timor, to the east by the Arafura Sea, and to the south by Australia. The sea contains a number of reefs, ...
in the early 1990s.


Biology

Examination of brain
endocast An endocast is the internal cast of a hollow object, often referring to the cranial vault in the study of brain development in humans and other organisms. Endocasts can be artificially made for examining the properties of a hollow, inaccessible sp ...
s 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 In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
lifestyle. ''A. maximus'' had relatively larger
olfactory bulb The olfactory bulb (Latin: ''bulbus olfactorius'') is a grey matter, neural structure of the vertebrate forebrain involved in olfaction, the sense of odor, smell. It sends olfactory information to be further processed in the amygdala, the orbitof ...
s 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 forest ...
in
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
and
New Guinea New Guinea (; Hiri Motu Hiri Motu, also known as Police Motu, Pidgin Motu, or just Hiri, is a language of Papua New Guinea, which is spoken in surrounding areas of Port Moresby (Capital of Papua New Guinea). It is a simplified version of ...
today. However, some rainforest fruits with thick, highly sculptured
endocarp Fruit anatomy is the plant anatomy of the internal structure of fruit. Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Agg ...
s, such as that of the currently undispersed and highly threatened forest
coconut The coconut tree (''Cocos nucifera'') is a member of the palm tree family ( Arecaceae) and the only living species of the genus ''Cocos''. The term "coconut" (or the archaic "cocoanut") can refer to the whole coconut palm, the seed, or the ...
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'' and '' Satranala decussilvae''), just like many cassowary-dispersed fruits.


Reproduction

Occasionally subfossil eggs are found intact. The
National Geographic Society The National Geographic Society (NGS), headquartered in Washington, D.C., United States, is one of the largest non-profit scientific and educational organizations in the world. Founded in 1888, its interests include geography, archaeology, and ...
in
Washington, D.C. ) , image_skyline = , image_caption = Clockwise from top left: the Washington Monument and Lincoln Memorial on the National Mall, United States Capitol, Logan Circle, Jefferson Memorial, White House, Adams Morgan, ...
holds a specimen of an ''Aepyornis'' egg which was given to
Luis Marden Luis Marden (born Annibale Luigi Paragallo) (January 25, 1913 – March 3, 2003) was an American photographer, explorer, writer, filmmaker, diver, navigator, and linguist who worked for ''National Geographic Magazine''. He worked as a photographer ...
in 1967. The specimen is intact and contains the skeleton of the unhatched bird. The
Denver Museum of Nature and Science The Denver Museum of Nature & Science is a municipal natural history and science museum in Denver, Colorado. It is a resource for informal science education in the Rocky Mountain region. A variety of exhibitions, programs, and activities help mus ...
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 The Harvard Museum of Natural History is a natural history museum housed in the University Museum Building, located on the campus of Harvard University in Cambridge, Massachusetts. It features 16 galleries with 12,000 speciments drawn from the col ...
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 The Grant Museum of Zoology and Comparative Anatomy is a natural history museum that is part of University College London in London, England. It was established by Robert Edmond Grant in 1828 as a teaching collection of zoological specimens and ...
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 degree- ...
. 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 Tur ...
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 A natural history museum or museum of natural history is a scientific institution with natural history collections that include current and historical records of animals, plants, fungi, ecosystems, geology, paleontology, climatology, and more. ...
. 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 The Delaware Museum of Nature & Science (DMNS) is a museum located since January 1, 2022. The museum was founded in 1957 by John Eleuthere du Pont near Greenville, Delaware; it opened in 1972 on a site near Winterthur, Delaware. It is known for i ...
, 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 adjacent Del ...
, US, and another in the
Natural History Museum, London The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum an ...
. The
Melbourne Museum The Melbourne Museum is a natural and cultural history museum located in the Carlton Gardens, Melbourne, Carlton Gardens in Melbourne, Australia. Located adjacent to the Royal Exhibition Building, the museum was opened in 2000 as a project of t ...
has two ''Aepyornis'' eggs. The first was acquired for £100 by Professor
Frederick McCoy Sir Frederick McCoy (1817 – 13 May 1899), was an Irish palaeontologist, zoologist, and museum administrator, active in Australia. He is noted for founding the Botanic Garden of the University of Melbourne in 1856. Early life McCoy was the so ...
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 The Western Foundation of Vertebrate Zoology (WFVZ) is a non-profit charitable organization based in Camarillo, California focused on research and education on bird conservation Bird conservation is a field in the science of conservation biolog ...
, 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 Buckinghamshire (), abbreviated Bucks, is a ceremonial county in South East England that borders Greater London to the south-east, Berkshire to the south, Oxfordshire to the west, Northamptonshire to the north, Bedfordshire to the north-ea ...
, UK. In the collections of the department of geology at the
Field Museum of Natural History The Field Museum of Natural History (FMNH), also known as The Field Museum, is a natural history museum in Chicago, Illinois, and is one of the largest such museums in the world. The museum is popular for the size and quality of its educational ...
in
Chicago (''City in a Garden''); I Will , image_map = , map_caption = Interactive Map of Chicago , coordinates = , coordinates_footnotes = , subdivision_type = Country , subdivision_name ...
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 The Regional Museum of Natural History, Bhubaneswar is a museum in Bhubaneswar, Odisha, India with exhibits on plants, animals and geology of the eastern region of India. The Regional Museum of Natural History at Bhubaneswar, was inaugurated in ...
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 Christie's is a British auction house founded in 1766 by James Christie (auctioneer), James Christie. Its main premises are on King Street, St James's in London, at Rockefeller Center in New York City and at Alexandra House in Hong Kong. It is ...
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 Miguel de Cervantes Saavedra (; 29 September 1547 (assumed) – 22 April 1616 Old Style and New Style dates, NS) was an Early Modern Spanish writer widely regarded as the greatest writer in the Spanish language and one of the world's pre-emin ...
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 Folk memory, also known as folklore or myths, refers to past events that have been passed orally from generation to generation. The events described by the memories may date back hundreds, thousands, or even tens of thousands of years and often hav ...
. 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
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fro ...
s such as
chicken The chicken (''Gallus gallus domesticus'') is a domesticated junglefowl species, with attributes of wild species such as the grey and the Ceylon junglefowl that are originally from Southeastern Asia. Rooster or cock is a term for an adult m ...
s and
guineafowl Guineafowl (; sometimes called "pet speckled hens" or "original fowl") are birds of the family Numididae in the order Galliformes. They are endemic to Africa and rank among the oldest of the gallinaceous birds. Phylogenetically, they branched o ...
. 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 Sinbad the Sailor (; ar, سندباد البحري, Sindibādu al-Bahriyy; fa, سُنباد بحری, Sonbād-e Bahri or Sindbad) is a fictional mariner and the hero of a story-cycle of Persian origin. He is described as hailing from Baghdad ...
'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 Herbert George Wells"Wells, H. G."
Revised 18 May 2015. ''
Æpyornis Island "Æpyornis Island", or "Aepyornis Island", is a short story by H. G. Wells, first published in 1894 in the ''Pall Mall Budget''. It was included in ''The Stolen Bacillus and Other Incidents'', the first collection of short stories by Wells, first ...
" (1894) about the bird. It was first collected in ''
The Stolen Bacillus and Other Incidents ''The Stolen Bacillus and Other Incidents'' is a collection of fifteen fantasy and science fiction short stories written by the English author H. G. Wells between 1893 and 1895. It was first published by Methuen & Co. in 1895 and was Wells's fir ...
'' (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 Late Quaternary prehistoric birds are avian taxa that became extinct during the Late Quaternary – the Holocene or Late Pleistocene – and before recorded history, or more precisely, before they could be studied alive by ornithol ...
*
New World Pleistocene extinctions The Quaternary period (from 2.588 ± 0.005 million years ago to the present) has seen the extinctions of numerous predominantly megafaunal species, which have resulted in a collapse in faunal density and diversity and the extinction of key ecol ...
*
Pleistocene megafauna Pleistocene megafauna is the set of large animals that lived on Earth during the Pleistocene epoch. Pleistocene megafauna became extinct during the Quaternary extinction event resulting in substantial changes to ecosystems globally. The role of hu ...


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)