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Moa (
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
Dinornithiformes) are an extinct group of
flightless bird Flightless birds are birds that cannot Bird flight, fly, as they have, through evolution, lost the ability to. There are over 60 extant species, including the well-known ratites (ostriches, emus, cassowary, cassowaries, Rhea (bird), rheas, an ...
s formerly endemic to
New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
.OSNZ (2009) During the
Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as the Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division ...
-
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
, there were nine
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
(in six genera). The two largest species, ''
Dinornis robustus The South Island giant moa (''Dinornis robustus'') is an extinct species of moa in the genus '' Dinornis'', known in Māori by the name moa nunui. It was one of the tallest-known bird species to walk the Earth, exceeded in weight only by the ...
'' and '' Dinornis novaezelandiae'', reached about in height with neck outstretched, and weighed about Davies, S.J.J.F. (2003) while the smallest, the
bush moa The bush moa, little bush moa, or lesser moa (''Anomalopteryx didiformis'') is an extinct species of moa from the family Emeidae (lesser moa) endemic to New Zealand. It is the only species in the genus ''Anomalopteryx'', from Ancient Greek ἀ ...
(''Anomalopteryx didiformis''), was around the size of a
turkey Turkey, officially the Republic of Türkiye, is a country mainly located in Anatolia in West Asia, with a relatively small part called East Thrace in Southeast Europe. It borders the Black Sea to the north; Georgia (country), Georgia, Armen ...
. Estimates of the moa population when
Polynesians Polynesians are an ethnolinguistic group comprising closely related ethnic groups native to Polynesia, which encompasses the islands within the Polynesian Triangle in the Pacific Ocean. They trace their early prehistoric origins to Island Sout ...
settled New Zealand circa 1300 vary between 58,000 and approximately 2.5 million. Moa are traditionally placed in the
ratite Ratites () are a polyphyletic group consisting of all birds within the infraclass Palaeognathae that lack keels and cannot fly. They are mostly large, long-necked, and long-legged, the exception being the kiwi, which is also the only nocturnal ...
group. Genetic studies have found that their closest relatives are the flighted South American
tinamou Tinamous () are members of the order Tinamiformes (), and family Tinamidae (), divided into two distinct subfamily, subfamilies, containing 46 species found in Mexico, Central America, and South America. The word "tinamou" comes from the Carib la ...
s, once considered a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to ratites. The nine species of moa were the only wingless birds, lacking even the
vestigial Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...
wings that all other ratites have. They were the largest terrestrial animals and dominant
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
s in New Zealand's forest, shrubland, and subalpine
ecosystems An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
until the arrival of the Māori, and were hunted only by
Haast's eagle Haast's eagle (''Hieraaetus moorei'') is an Extinction, extinct species of eagle that lived in the South Island of New Zealand, commonly accepted to be the of Māori mythology.
. Moa extinction occurred within 100 years of human settlement of New Zealand, primarily due to overhunting.


Etymology

The word ''moa'' is a Polynesian term for domestic fowl. The name was not in common use among the Māori by the time of European contact, likely because the bird it described had been extinct for some time, and traditional stories about it were rare. The earliest record of the name was by missionaries William Williams and
William Colenso William Colenso (17 November 1811 – 10 February 1899) FRS was a Cornish Christian missionary to New Zealand, and also a printer, botanist, explorer and politician. He attended the signing of the Treaty of Waitangi and later wrote an acco ...
in January 1838; Colenso speculated that the birds may have resembled gigantic fowl. In 1912, Māori chief Urupeni Pūhara claimed that the moa's traditional name was "te kura" (the red bird).


Description

Moa skeletons were traditionally reconstructed in an upright position to create impressive height, but analysis of their vertebral articulations indicates that they probably carried their heads forward,Worthy & Holdaway (2002) in the manner of a
kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), an informal name for New Zealanders * Kiwifruit, an edible hairy fruit with many seeds * Kiwi dollar or New Zealand dollar, a unit of curren ...
. The spine was attached to the rear of the head rather than the base, indicating the horizontal alignment. This would have let them graze on low vegetation, while being able to lift their heads and browse trees when necessary. This has resulted in a reconsideration of the height of larger moa. However, Māori
rock art In archaeology, rock arts are human-made markings placed on natural surfaces, typically vertical stone surfaces. A high proportion of surviving historic and prehistoric rock art is found in caves or partly enclosed rock shelters; this type al ...
depicts moa or moa-like birds with necks upright, indicating that moa were capable of assuming both neck postures. No records survive of what sounds moa made, though some idea of their calls can be gained from fossil evidence. The
trachea The trachea (: tracheae or tracheas), also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all animals' lungs. The trachea extends from ...
of moa were supported by many small rings of bone known as tracheal rings. These rings were preserved within some articulated skeletons, and shows that at least two moa genera (''Euryapteryx'' and ''Emeus'') exhibited tracheal elongation, where their trachea, up to long, formed a large loop within the
body cavity A body cavity is any space or compartment, or potential space, in an animal body. Cavities accommodate organs and other structures; cavities as potential spaces contain fluid. The two largest human body cavities are the ventral body cavity, a ...
. They are the only ratites known to exhibit this feature, which is also present in several other bird groups, including
swan Swans are birds of the genus ''Cygnus'' within the family Anatidae. The swans' closest relatives include the goose, geese and ducks. Swans are grouped with the closely related geese in the subfamily Anserinae where they form the tribe (biology) ...
s, cranes, and
guinea fowl Guinea fowl () (or guineahen) are birds of the family Numididae in the order Galliformes. They are endemic to Africa and rank among the oldest of the gallinaceous birds. Phylogenetics, Phylogenetically, they branched off from the core Galliforme ...
. The feature is associated with deep resonant vocalisations that can travel long distances.


Feathers and soft tissues

Several examples of moa remains have been found with soft tissues (
muscle Muscle is a soft tissue, one of the four basic types of animal tissue. There are three types of muscle tissue in vertebrates: skeletal muscle, cardiac muscle, and smooth muscle. Muscle tissue gives skeletal muscles the ability to muscle contra ...
,
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other animal coverings, such as the arthropod exoskeleton, have different ...
,
feathers Feathers are epidermis (zoology), epidermal growths that form a distinctive outer covering, or plumage, on both Bird, avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in ...
) preserved through
desiccation Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. The ...
after the bird died at a dry site (for example, a cave with a constant dry breeze blowing through it); these specimens are a form of
natural mummy A mummy is a dead human or an animal whose soft tissues and Organ (biology), organs have been preserved by either intentional or accidental exposure to Chemical substance, chemicals, extreme cold, very low humidity, or lack of air, so that the ...
. Most were found in the
semiarid A semi-arid climate, semi-desert climate, or steppe climate is a dry climate sub-type. It is located on regions that receive precipitation below potential evapotranspiration, but not as low as a desert climate. There are different kinds of sem ...
Central Otago region, the driest part of New Zealand, but others are known from outside the Central Otago region. These include: * Dried muscle on bones of a female ''Dinornis robustus'' found at Tiger Hill in the
Manuherikia River The Manuherikia River is located in Otago in the South Island of New Zealand. It rises in the far north of the Maniototo, with the West Branch draining the eastern side of the St Bathans Range, and the East Branch draining the western flanks of ...
Valley by gold miners in 1864Owen, R. (1879) (currently held by
Yorkshire Museum The Yorkshire Museum is a museum in York, England. It was opened in 1830, and has five permanent collections, covering biology, geology, archaeology, numismatics and astronomy. History The museum was founded by the Yorkshire Philosophical Soci ...
) * Several bones of ''Emeus crassus'' with muscle attached, and a row of neck vertebrae with muscle, skin, and feathers collected from Earnscleugh Cave near the town of Alexandra in 1870 (currently held by the
Otago Museum Otago (, ; ) is a region of New Zealand located in the southern half of the South Island and administered by the Otago Regional Council. It has an area of approximately , making it the country's second largest local government region. Its po ...
) *An articulated foot of a male ''D. giganteus'' with skin and foot pads preserved, found in a crevice on the Knobby Range in 1874Buller, W.L. (1888) (currently held by the
Otago Museum Otago (, ; ) is a region of New Zealand located in the southern half of the South Island and administered by the Otago Regional Council. It has an area of approximately , making it the country's second largest local government region. Its po ...
) * The type specimen of ''Megalapteryx didinus'' found near Queenstown in 1878 (currently held by
Natural History Museum A natural history museum or museum of natural history is a scientific institution with natural history scientific collection, collections that include current and historical records of animals, plants, Fungus, fungi, ecosystems, geology, paleo ...
, London; see photograph of foot on this page) * The lower leg of ''Pachyornis elephantopus'', with skin and muscle, from the Hector Range in 1884; (currently held by the Zoology Department,
Cambridge University The University of Cambridge is a Public university, public collegiate university, collegiate research university in Cambridge, England. Founded in 1209, the University of Cambridge is the List of oldest universities in continuous operation, wo ...
) * The complete feathered leg of a ''M. didinus'' from Old Man Range in 1894 (currently held by the
Otago Museum Otago (, ; ) is a region of New Zealand located in the southern half of the South Island and administered by the Otago Regional Council. It has an area of approximately , making it the country's second largest local government region. Its po ...
) * The head of a ''M. didinus'' found near Cromwell sometime before 1949 (currently held by
Te Papa Tongarewa The Museum of New Zealand Te Papa Tongarewa is New Zealand's national museum and is located in Wellington. Usually known as Te Papa ( Māori for ' the treasure box'), it opened in 1998 after the merging of the National Museum of New Zealand a ...
). * A complete foot of ''M. didinus'' found in a cave on Mount Owen near Nelson in the 1980s (currently held by
Te Papa Tongarewa The Museum of New Zealand Te Papa Tongarewa is New Zealand's national museum and is located in Wellington. Usually known as Te Papa ( Māori for ' the treasure box'), it opened in 1998 after the merging of the National Museum of New Zealand a ...
) * A skeleton of ''Anomalopteryx didiformis'' with muscle, skin, and feather bases collected from a cave near
Te Anau Te Anau is a town in the Southland, New Zealand, Southland List of regions in New Zealand, region of the South Island of New Zealand. In Māori language, Māori, Te-Anau means the Place of the Swirling Waters. It is on the eastern shore of Lake ...
in 1980. In addition to these specimens, loose moa feathers have been collected from caves and rock shelters in the southern South Island, and based on these remains, some idea of the moa plumage has been achieved. The preserved leg of ''M. didinus'' from the Old Man Range reveals that this species was feathered right down to the foot. This is likely to have been an adaptation to living in high-altitude, snowy environments, and is also seen in the Darwin’s rhea, which lives in a similar seasonally snowy habitat. Moa feathers are up to long, and a range of colours has been reported, including reddish-brown, white, yellowish, and purplish. Dark feathers with white or creamy tips have also been found, and indicate that some moa species may have had plumage with a speckled appearance.


Classification


Evolutionary relationships

The moa's closest relatives are small terrestrial South American birds called the
tinamou Tinamous () are members of the order Tinamiformes (), and family Tinamidae (), divided into two distinct subfamily, subfamilies, containing 46 species found in Mexico, Central America, and South America. The word "tinamou" comes from the Carib la ...
s, which can fly.Phillips, et al. (2010) Previously, the
kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), an informal name for New Zealanders * Kiwifruit, an edible hairy fruit with many seeds * Kiwi dollar or New Zealand dollar, a unit of curren ...
, the Australian
emu The emu (; ''Dromaius novaehollandiae'') is a species of flightless bird endemism, endemic to Australia, where it is the Tallest extant birds, tallest native bird. It is the only extant taxon, extant member of the genus ''Dromaius'' and the ...
, and
cassowary Cassowaries (; Biak: ''man suar'' ; ; Papuan: ''kasu weri'' ) 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. Cassowaries a ...
Turvey et al. (2005) were thought to be most closely related to moa. Although dozens of species were described in the late 19th and early 20th centuries, many were based on partial skeletons and turned out to be
synonyms A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
. Currently, 11 species are formally recognised, although recent studies using
ancient DNA Ancient DNA (aDNA) is DNA isolated from ancient sources (typically Biological specimen, specimens, but also environmental DNA). Due to degradation processes (including Crosslinking of DNA, cross-linking, deamination and DNA fragmentation, fragme ...
recovered from bones in museum collections suggest that distinct lineages exist within some of these. One factor that has caused much confusion in moa taxonomy is the intraspecific variation of bone sizes, between glacial and interglacial periods (see Bergmann’s rule and Allen’s rule), as well as
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
being evident in several species. ''
Dinornis The giant moa (''Dinornis'') is an extinct genus of birds belonging to the moa family. As with other moa, it was a member of the order Dinornithiformes. It was endemic to New Zealand. Two species of ''Dinornis'' are considered valid, the Nort ...
'' seems to have had the most pronounced sexual dimorphism, with females being up to 150% as tall and 280% as heavy as males—so much bigger that they were classified as separate species until 2003.Huynen, L.J., et al. (2003)Bunce, M., et al. (2003) A 2009 study showed that ''Euryapteryx curtus'' and ''E. gravis'' were synonyms. A 2010 study explained size differences among them as sexual dimorphism. A 2012 morphological study interpreted them as subspecies, instead. Analyses of ancient DNA have determined that a number of cryptic evolutionary lineages occurred in several moa genera. These may eventually be classified as species or subspecies; ''Megalapteryx benhami'' (Archey) is synonymised with ''M. didinus'' (Owen) because the bones of both share all essential characters. Size differences can be explained by a north–south
cline Cline may refer to: Science * Cline (biology), a measurable gradient in a single trait in a species across its geographical range * Cline (hydrology), a fluid layer with a property that varies * Cline (mathematics) or generalised circle, a ci ...
combined with temporal variation such that specimens were larger during the Otiran glacial period (the last ice age in New Zealand). Similar temporal size variation is known for the North Island's '' Pachyornis mappini''. Some of the other size variation for moa species can probably be explained by similar geographic and temporal factors. The earliest moa remains come from the
Miocene The Miocene ( ) is the first epoch (geology), 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 mea ...
Saint Bathans Fauna The St Bathans fauna is found in the lower Bannockburn Formation of the Manuherikia Group of Central Otago, in the South Island of New Zealand. It comprises a suite of fossilised prehistoric animals from the late Early Miocene (Altonian) perio ...
. Known from multiple eggshells and hind limb elements, these represent at least two already fairly large-sized species.


Phylogeny

Because moa are a group of flightless birds with no vestiges of wing bones, questions have been raised about how they arrived in New Zealand, and from where. Many theories exist about the moa's arrival and radiation in New Zealand, but the most recent theory suggests that they arrived in New Zealand about 60 million years ago (Mya) and split from the "basal" (see below) moa species, ''Megalapteryx'', about 5.8 Mya instead of the 18.5 Mya split suggested by Baker et al. (2005). This does not necessarily mean there was no speciation between the arrival 60 Mya and the basal split 5.8 Mya, but the fossil record is lacking and most likely the early moa lineages existed, but became extinct before the basal split 5.8 Mya. The presence of
Miocene The Miocene ( ) is the first epoch (geology), 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 mea ...
-aged species certainly suggests that moa diversification began before the split between ''Megalapteryx'' and the other taxa. The Oligocene Drowning Maximum event, which occurred about 22 Mya, when only 18% of present-day New Zealand was above sea level, is very important in the moa radiation. Because the basal moa split occurred so recently (5.8 Mya), it was argued that ancestors of the Quaternary moa lineages could not have been present on both the South and North Island remnants during the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
drowning. This does not imply that moa were previously absent from the North Island, but that only those from the South Island survived, because only the South Island was above sea level. Bunce et al. (2009) argued that moa ancestors survived on the South Island and then recolonised the North Island about 2 Myr later, when the two islands rejoined after 30 Myr of separation. The presence of Miocene moa in the Saint Bathans fauna seems to suggest that these birds increased in size soon after the Oligocene drowning event, if they were affected by it at all. Bunce et al. also concluded that the highly complex structure of the moa lineage was caused by the formation of the
Southern Alps The Southern Alps (; officially Southern Alps / Kā Tiritiri o te Moana) are a mountain range extending along much of the length of New Zealand, New Zealand's South Island, reaching its greatest elevations near the range's western side. The n ...
about 6 Mya, and the habitat fragmentation on both islands resulting from Pleistocene glacial cycles,
volcanism Volcanism, vulcanism, volcanicity, or volcanic activity is the phenomenon where solids, liquids, gases, and their mixtures erupt to the surface of a solid-surface astronomical body such as a planet or a moon. It is caused by the presence of a he ...
, and landscape changes. The cladogram below is a phylogeny of Palaeognathae generated by Mitchell (2014) with some clade names after Yuri ''et al.'' (2013). It provides the position of the moa (Dinornithiformes) within the larger context of the "ancient jawed" (Palaeognathae) birds: The cladogram below gives a more detailed, species-level phylogeny, of the moa branch (Dinornithiformes) of the "ancient jawed" birds (Palaeognathae) shown above:


Taxonomy

The currently recognised genera and species are: *Order
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
Dinornithiformes (Gadow 1893) Ridgway 1901 inornithes Gadow 1893; Immanes Newton 1884(moa) **Family
Dinornithidae The giant moa (''Dinornis'') is an extinct genus of birds belonging to the moa family. As with other moa, it was a member of the order Dinornithiformes. It was endemic to New Zealand. Two species of ''Dinornis'' are considered valid, the No ...
Owen 1843 alapteryginae Bonaparte 1854; Palapterygidae Haast 1874; Dinornithnideae Stejneger 1884(giant moa) ***Genus ''
Dinornis The giant moa (''Dinornis'') is an extinct genus of birds belonging to the moa family. As with other moa, it was a member of the order Dinornithiformes. It was endemic to New Zealand. Two species of ''Dinornis'' are considered valid, the Nort ...
'' ****
North Island giant moa The North Island giant moa (''Dinornis novaezealandiae'') is an extinct moa in the genus ''Dinornis'', known in Māori as kuranui. It was a large, herbivorous bird belonging to the order Dinornithiformes, and exhibited a strong sexual dimorph ...
, ''Dinornis novaezealandiae'' (North Island, New Zealand) ****
South Island giant moa The South Island giant moa (''Dinornis robustus'') is an extinct species of moa in the genus ''Dinornis'', known in Māori language, Māori by the name moa nunui. It was one of the tallest-known bird species to walk the Earth, exceeded in weigh ...
, ''Dinornis robustus'' (South Island, New Zealand) **Family
Emeidae The lesser moa (family Emeidae) were a family in the moa order Dinornithiformes. About two-thirds of all moa species are in the lesser moa family. The moa were ratites from New Zealand: flightless birds with a sternum but without a keel. They al ...
(Bonaparte 1854) meinae Bonaparte 1854; Anomalopterygidae Oliver 1930; Anomalapteryginae Archey 1941(lesser moa) ***Genus '' Anomalopteryx'' ****
Bush moa The bush moa, little bush moa, or lesser moa (''Anomalopteryx didiformis'') is an extinct species of moa from the family Emeidae (lesser moa) endemic to New Zealand. It is the only species in the genus ''Anomalopteryx'', from Ancient Greek ἀ ...
, ''Anomalopteryx didiformis'' (North and South Island, New Zealand) ***Genus '' Emeus'' ****
Eastern moa The eastern moa (''Emeus crassus'') is an extinct species of moa that was endemic to New Zealand. Taxonomy When the first specimens were originally described by Richard Owen in 1846, they were placed within the genus ''Dinornis'' as three d ...
, ''Emeus crassus'' (South Island, New Zealand) ***Genus '' Euryapteryx'' ****
Broad-billed moa The broad-billed moa, stout-legged moa, or coastal moa (''Euryapteryx curtus'') is an extinct species of moa that was endemic to New Zealand. Taxonomy ''Euryapteryx curtus'' is a ratite and a member of the lesser moa family. The ratites a ...
, ''Euryapteryx curtus'' (North and South Island, New Zealand) ***Genus ''
Pachyornis ''Pachyornis'' is an extinct genus of ratites from New Zealand which belongs to the moa family. Like all ratites, ''Pachyornis'' is a flightless bird with a sternum that lacks a keel. They also have a distinctive palate. The genus currently cont ...
'' ****
Heavy-footed moa The heavy-footed moa (''Pachyornis elephantopus'') is an extinct species of moa from the lesser moa family. The heavy-footed moa was widespread across the South Island of New Zealand, and inhabited lowland environments like shrublands, dunelan ...
, ''Pachyornis elephantopus'' (South Island, New Zealand) ****
Mantell's moa Mantell's moa (''Pachyornis geranoides'') also known as Mappin's moa or moa ruarangi, is an extinct species of moa from the North Island of New Zealand. It inhabited lowland environments like shrublands, grasslands, dunelands, and forests.Davie ...
, ''Pachyornis geranoides'' (North Island, New Zealand) ****
Crested moa The crested moa (''Pachyornis australis'') is an extinct species of moa. It is one of the 9 known species of moa to have existed. Moa are grouped together with emus, ostriches, kiwi, cassowaries, rheas, and tinamous in the clade Palaeognathae. ...
, '' Pachyornis australis'' (South Island, New Zealand)Stephenson, Brent (2009) **Family Megalapterygidae ***
Genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''
Megalapteryx The upland moa (''Megalapteryx didinus'') is an extinct species of moa that was endemic to New Zealand. The species was named by Richard Owen in 1883, and belongs to the ratites, a group of flightless birds with no keel on the sternum. Of all m ...
'' ****
Upland moa The upland moa (''Megalapteryx didinus'') is an extinct species of moa that was endemic to New Zealand. The species was named by Richard Owen in 1883, and belongs to the ratites, a group of flightless birds with no keel on the sternum. Of all m ...
, ''Megalapteryx didinus'' (South Island, New Zealand) Two unnamed species are also known from the Saint Bathans Fauna.


Distribution and habitat

Analyses of fossil moa bone assemblages have provided detailed data on the habitat preferences of individual moa species, and revealed distinctive regional moa faunas:


South Island

The two main faunas identified in the South Island include: : The fauna of the high-rainfall west coast beech (''
Nothofagus ''Nothofagus'', also known as the southern beeches, is a genus of 43 species of trees and shrubs native to the Southern Hemisphere, found across southern South America (Chile, Argentina) and east and southeast Australia, New Zealand, New Guin ...
'') forests that included ''Anomalopteryx didiformis'' (bush moa) and ''Dinornis robustus'' (South Island giant moa), and : The fauna of the dry rainshadow forest and shrublands east of the
Southern Alps The Southern Alps (; officially Southern Alps / Kā Tiritiri o te Moana) are a mountain range extending along much of the length of New Zealand, New Zealand's South Island, reaching its greatest elevations near the range's western side. The n ...
that included ''Pachyornis elephantopus'' (heavy-footed moa), ''Euryapteryx curtus'', ''Emeus crassus'', and ''Dinornis robustus''. A '
subalpine Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial f ...
fauna' might include the widespread ''D. robustus'', and the two other moa species that existed in the South Island: :: ''Pachyornis australis'', the rarest moa species, the only moa species not yet found in Māori
midden A midden is an old dump for domestic waste. It may consist of animal bones, human excrement, botanical material, mollusc shells, potsherds, lithics (especially debitage), and other artifacts and ecofacts associated with past human oc ...
s. Its bones have been found in caves in the northwest
Nelson Nelson may refer to: Arts and entertainment * ''Nelson'' (1918 film), a historical film directed by Maurice Elvey * ''Nelson'' (1926 film), a historical film directed by Walter Summers * ''Nelson'' (opera), an opera by Lennox Berkeley to a lib ...
and
Karamea Karamea is a town on the West Coast, New Zealand, West Coast of the South Island of New Zealand. It is the northernmost settlement of any real size on the West Coast, and is located northeast by road from Westport, New Zealand, Westport. Apart ...
districts (such as Honeycomb Hill Cave), and some sites around the
Wānaka Wānaka () is a popular ski and summer resort town in the Otago region of the South Island of New Zealand. At the southern end of Lake Wānaka, it is at the start of the Clutha River and is the gateway to Mount Aspiring National Park. Wānaka ...
district. :: ''Megalapteryx didinus'', more widespread, named "upland moa" because its bones are commonly found in the subalpine zone. However, it also occurred down to sea level, where suitable steep and rocky terrain (such as
Punakaiki Punakaiki is a small village on the West Coast, New Zealand, West Coast of the South Island of New Zealand. It is located between Westport, New Zealand, Westport and Greymouth on , the only through-road on the West Coast. Punakaiki is immediatel ...
on the west coast and
Central Otago Central Otago is an area located in the inland part of the Otago region in the South Island of New Zealand. The motto for the area is "A World of Difference". The area is dominated by mountain ranges and the upper reaches of the Clutha River ...
) existed. Their distributions in coastal areas have been rather unclear, but were present at least in several locations such as on
Kaikōura Kaikōura (; ) is a town on the east coast of the South Island of New Zealand, located on New Zealand State Highway 1, State Highway 1, north of Christchurch. The town has an estimated permanent resident population of as of . Kaikōura is th ...
,
Otago Peninsula The Otago Peninsula () is a long, hilly indented finger of land that forms the easternmost part of Dunedin, New Zealand. Volcanic in origin, it forms one wall of the eroded valley that now forms Otago Harbour. The peninsula lies south-east of Ot ...
, and
Karitane The small town of Karitane is located within the limits of the city of Dunedin in New Zealand, 35 kilometres to the north of the city centre. Set in rolling country near the mouth of the Waikouaiti River, the town is a popular holiday retreat f ...
.


North Island

Significantly less is known about North Island paleofaunas, due to the scarcity of fossil sites compared to the South Island, but the basic pattern of moa-habitat relationships was the same. The South Island and the North Island shared some moa species (''Euryapteryx curtus'', ''Anomalopteryx didiformis''), but most were exclusive to one island, reflecting divergence over several thousand years since lower sea level in the
Ice Age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages, and g ...
had made a land bridge across the
Cook Strait Cook Strait () is a strait that separates the North Island, North and South Islands of New Zealand. The strait connects the Tasman Sea on the northwest with the South Pacific Ocean on the southeast. It is wide at its narrowest point,McLintock, ...
. In the North Island, ''Dinornis novaezealandiae'' and ''Anomalopteryx didiformis'' dominated in high-rainfall forest habitat, a similar pattern to the South Island. The other moa species present in the North Island (''Euryapteryx'' ''curtus'', and ''Pachyornis geranoides'') tended to inhabit drier forest and shrubland habitats. ''P. geranoides'' occurred throughout the North Island.


Behaviour and ecology

About eight moa
trackway Historic roads (or historic trails in the US and Canada) are paths or routes that have historical importance due to their use over a period of time. Examples exist from prehistoric times until the early 20th century. They include ancient track ...
s, with fossilised moa footprint impressions in fluvial silts, have been found in the North Island, including
Waikanae Waikanae (, ) is a town on the Kāpiti Coast, north of the Wellington, New Zealand. The name is a Māori language, Māori word meaning "waters" (''wai'') "of the flathead grey mullet, grey mullet". The area surrounding the town is notable fo ...
Creek (1872), Napier (1887),
Manawatū River The Manawatū River is a major river of the lower North Island of New Zealand. The river flows from the Ruahine Ranges, through both the Manawatū Gorge and the city of Palmerston North, and across the Manawatū Plains to the Tasman Sea at Foxto ...
(1895), Marton (1896),
Palmerston North Palmerston North (; , colloquially known as Palmerston or Palmy) is a city in the North Island of New Zealand and the seat of the Manawatū-Whanganui region. Located in the eastern Manawatū Plains, the city is near the north bank of the Manaw ...
(1911) (see photograph to left),
Rangitīkei River The Rangitīkei River is one of New Zealand's longest rivers, long. Its headwaters are to the southeast of Lake Taupō in the Kaimanawa Ranges. It flows from the Central Plateau south past Taihape, Mangaweka, Hunterville, Marton, and B ...
(1939), and under water in
Lake Taupō Lake Taupō (also spelled Taupo; or ) is a large crater lake in New Zealand's North Island, located in the caldera of Taupō Volcano. The lake is the namesake of the town of Taupō, which sits on a bay in the lake's northeastern shore. With ...
(1973). Analysis of the spacing of these tracks indicates walking speeds between . In 2022, an additional trackway was discovered at the entrance of
Kaipara Harbour Kaipara Harbour is a large enclosed harbour estuary complex on the north western side of the North Island of New Zealand. The northern part of the harbour is administered by the Kaipara District and the southern part is administered by the Auck ...
. In 2019, the first known trackway in the South Island was found in a riverbed at
Kyeburn Kyeburn is a small settlement in Otago, in the South Island of New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and L ...
, Otago. A second set of footprints were discovered at
Pareora Pareora is a small town in the South Island of New Zealand. It is located close to State Highway 1, which bypasses the western edge of the town, and close to the Pacific Ocean coast, five kilometres north of Saint Andrews and 10 kilometres sou ...
, Canterbury, in 2022. Studies of accumulated dried vegetation in the pre-human mid-late Holocene period suggests a low ''
Sophora microphylla ''Sophora microphylla'', commonly known as weeping kōwhai and small-leaved kōwhai, is a species of flowering tree in the family Fabaceae native to New Zealand. It is the most widespread of the eight species of kōwhai (the New Zealand ''Sophora ...
'' or Kōwai forest ecosystem in
Central Otago Central Otago is an area located in the inland part of the Otago region in the South Island of New Zealand. The motto for the area is "A World of Difference". The area is dominated by mountain ranges and the upper reaches of the Clutha River ...
that was used by — and perhaps maintained by — moa for both nesting material and food. By the 1850s, when European settlers came to the area, neither the forests nor moa were present, vanishing prior to settlement by colonists.


Diet

Their diet has been deduced from
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 preserve ...
ised contents of their
gizzard The gizzard, also referred to as the ventriculus, gastric mill, and gigerium, is an organ found in the digestive tract of some animals, including archosaurs (birds and other dinosaurs, crocodiles, alligators, pterosaurs), earthworms, some gast ...
sWood (2007) and
coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name ...
s, as well as indirectly through morphological analysis of skull and beak, and
stable isotope analysis Isotope analysis is the identification of isotopic signature, abundance of certain stable isotopes of chemical elements within organic and inorganic compounds. Isotopic analysis can be used to understand the flow of energy through a food web ...
of their bones. Moa fed on a range of plant species and plant parts, including fibrous twigs and leaves taken from low trees and shrubs. The beak of ''
Pachyornis ''Pachyornis'' is an extinct genus of ratites from New Zealand which belongs to the moa family. Like all ratites, ''Pachyornis'' is a flightless bird with a sternum that lacks a keel. They also have a distinctive palate. The genus currently cont ...
elephantopus'' was analogous to a pair of
secateur Pruning shears, also called hand pruners (in American English) or secateurs (in British English), are a type of scissors used for plants. They are strong enough to prune hard branches of trees and shrubs, sometimes up to two centimetres thick. T ...
s, and could clip the fibrous leaves of New Zealand flax (''
Phormium ''Phormium'' is a genus of two plant species in the family Asphodelaceae. One species is endemic to New Zealand and the other is native to New Zealand and Norfolk Island. The two species are widely known in New Zealand as flax or their Māori l ...
tenax'') and twigs up to at least 8 mm in diameter. Moa filled the
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of Resource (biology), resources an ...
occupied in other countries by large browsing mammals such as
antelope The term antelope refers to numerous extant or recently extinct species of the ruminant artiodactyl family Bovidae that are indigenous to most of Africa, India, the Middle East, Central Asia, and a small area of Eastern Europe. Antelopes do ...
and
llamas The llama (; or ) (''Lama glama'') is a domesticated South American camelid, widely used as a meat and pack animal by Andean cultures since the pre-Columbian era. Llamas are social animals and live with others as a herd. Their wool is so ...
. Some biologists contend that a number of plant species evolved to avoid moa browsing. Divaricating plants such as ''
Pennantia corymbosa ''Pennantia corymbosa'', commonly known as kaikomako (from the Māori ), is a small dioecious tree endemic to New Zealand. Small, creamy flowers are produced between November and February, followed by a shiny black fruit in autumn. They are a ...
'' (the kaikōmako), which have small leaves and a dense mesh of branches, and ''
Pseudopanax crassifolius ''Pseudopanax crassifolius'', also known as horoeka or lancewood, is a Heteroblasty, heteroblastic tree belonging to the family Araliaceae. It is Endemism, endemic to New Zealand and found throughout the country from sea level up to about in l ...
'' (the horoeka or lancewood), which has tough juvenile leaves, are possible examples of plants that evolved in such a way. Likewise, it has been suggested that
heteroblasty Heteroblasty is the significant and abrupt change in form and function, that occurs over the lifespan of certain plants. Characteristics affected include internode length and stem structure as well as leaf form, size and arrangement. It should no ...
might be a response to moa browsing. Like many other birds, moa swallowed gizzard stones (
gastrolith A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In ...
s), which were retained in their muscular
gizzards The gizzard, also referred to as the ventriculus, gastric mill, and gigerium, is an organ found in the digestive tract of some animals, including archosaurs (birds and other dinosaurs, crocodiles, alligators, pterosaurs), earthworms, some gastro ...
, providing a grinding action that allowed them to eat coarse plant material. This grinding action suggests that moa were not effective seed dispersers, with only the smallest seeds passing through their gut intact. These stones were commonly smooth rounded quartz pebbles, but stones over long have been found among preserved moa gizzard contents. ''Dinornis'' gizzards could often contain several kilograms of stones. Moa likely exercised a certain selectivity in the choice of gizzard stones and chose the hardest pebbles.


Reproduction

The pairs of species of moa described as ''Euryapteryx curtus'' / ''E. exilis'', ''Emeus huttonii'' / ''E. crassus'', and ''Pachyornis septentrionalis'' / ''P. mappini'' have long been suggested to constitute males and females, respectively. This has been confirmed by analysis for sex-specific genetic markers of DNA extracted from bone material. For example, before 2003, three species of ''Dinornis'' were recognised: South Island giant moa (''D. robustus''), North Island giant moa (''D. novaezealandiae''), and slender moa (''D. struthioides''). However, DNA showed that all ''D. struthioides'' were males, and all ''D. robustus'' were females. Therefore, the three species of ''Dinornis'' were reclassified as two species, one each formerly occurring on New Zealand's North Island (''D. novaezealandiae'') and South Island (''D. robustus''); ''D. robustus'' however, comprises three distinct genetic lineages and may eventually be classified as many species, as discussed above. Examination of growth rings in moa cortical bone has revealed that these birds were
K-selected In ecology, selection theory relates to the natural selection, selection of combinations of Trait (biological), traits in an organism that trade off between quantity and quality of offspring. The focus on either an increased quantity of offsprin ...
, as are many other large endemic New Zealand birds. They are characterised by having a low
fecundity Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the capability to produc ...
and a long maturation period, taking about 10 years to reach adult size. The large ''Dinornis'' species took as long to reach adult size as small moa species, and as a result, had fast skeletal growth during their juvenile years. No evidence has been found to suggest that moa were colonial nesters. Moa nesting is often inferred from accumulations of eggshell fragments in
caves Caves or caverns are natural voids under the Earth's surface. Caves often form by the weathering of rock and often extend deep underground. Exogene caves are smaller openings that extend a relatively short distance underground (such as rock ...
and rock shelters, little evidence exists of the
nest A nest is a structure built for certain animals to hold Egg (biology), eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of ...
s themselves. Excavations of rock shelters in the eastern North Island during the 1940s found moa nests, which were described as "small depressions obviously scratched out in the soft dry
pumice Pumice (), called pumicite in its powdered or dust form, is a volcanic rock that consists of extremely vesicular rough-textured volcanic glass, which may or may not contain crystals. It is typically light-colored. Scoria is another vesicula ...
". Moa nesting material has also been recovered from rock shelters in the Central Otago region of the South Island, where the dry climate has preserved plant material used to build the nesting platform (including twigs clipped by moa bills).Wood, J.R. (2008)
Seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s and
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
within moa coprolites found among the nesting material provide evidence that the nesting season was late spring to summer. Fragments of moa eggshell are often found in
archaeological Archaeology or archeology is the study of human activity through the recovery and analysis of material culture. The archaeological record consists of Artifact (archaeology), artifacts, architecture, biofact (archaeology), biofacts or ecofacts, ...
sites and
sand dunes A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
around the New Zealand coast. Thirty-six whole moa eggs exist in museum collections and vary greatly in size (from in length and wide). The outer surface of moa eggshell is characterised by small, slit-shaped pores. The eggs of most moa species were white, although those of the upland moa (''Megalapteryx didinus'') were blue-green.Huynen, Leon; Gill, Brian J.; Millar, Craig D.; and Lambert, David M. (2010) A 2010 study by Huynen et al. found that the eggs of certain species were fragile, only around a millimetre in shell thickness: "Unexpectedly, several thin-shelled eggs were also shown to belong to the heaviest moa of the genera ''Dinornis'', ''Euryapteryx'', and ''Emeus'', making these, to our knowledge, the most fragile of all avian eggs measured to date. Moreover, sex-specific DNA recovered from the outer surfaces of eggshells belonging to species of ''Dinornis'' and ''Euryapteryx'' suggest that these very thin eggs were likely to have been incubated by the lighter males. The thin nature of the eggshells of these larger species of moa, even if incubated by the male, suggests that egg breakage in these species would have been common if the typical contact method of avian egg incubation was used." Despite the bird's extinction, the high yield of DNA available from recovered fossilised eggs has allowed the moa's genome to be sequenced.


Relationship with humans


Discovery

Before the arrival of humans, the moa's only predator was the massive
Haast's eagle Haast's eagle (''Hieraaetus moorei'') is an Extinction, extinct species of eagle that lived in the South Island of New Zealand, commonly accepted to be the of Māori mythology.
. New Zealand had been isolated for 80 million years and had few predators before human arrival, meaning that not only were its ecosystems extremely vulnerable to perturbation by outside species, but also the native species were ill-equipped to cope with human predators. Polynesians arrived sometime before 1300, and all moa genera were soon driven to extinction by hunting and, to a lesser extent, by habitat reduction due to forest clearance. By 1445, all moa had become extinct, along with Haast's eagle, which had relied on them for food. Recent research using
carbon-14 dating Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for Chronological dating, determining the age of an object containing organic material by using the properties of carbon-14, radiocarbon, a radioactive Isotop ...
of
midden A midden is an old dump for domestic waste. It may consist of animal bones, human excrement, botanical material, mollusc shells, potsherds, lithics (especially debitage), and other artifacts and ecofacts associated with past human oc ...
s strongly suggests that the events leading to extinction took less than a hundred years, rather than a period of exploitation lasting several hundred years as previously hypothesised. An expedition in the 1850s under Lieutenant A. Impey reported two emu-like birds on a hillside in the South Island; an 1861 story from the ''Nelson Examiner'' told of three-toed footprints measuring between
Tākaka Tākaka is a small town situated at the southeastern end of Golden Bay / Mohua, Golden Bay, at the northern end of New Zealand's South Island, located on the lower reaches of the Tākaka River. State Highway 60 (New Zealand), State Highway 60 r ...
and
Riwaka Riwaka () is a small settlement in the Tasman District of New Zealand's South Island. It lies beside Tasman Bay / Te Tai-o-Aorere, five kilometres north of Motueka, and close to the mouth of the Riuwaka River. The land where the town is based w ...
that were found by a surveying party; and finally in 1878, the ''
Otago Witness The ''Otago Witness'' was a prominent illustrated weekly newspaper in the early years of the European settlement of New Zealand, produced in Dunedin, the provincial capital of Otago. Published weekly, it existed from 1851 to 1932. The introduct ...
'' published an additional account from a farmer and his shepherd. An 86-year-old woman, Alice McKenzie, claimed in 1959 that she had seen a moa in
Fiordland Fiordland (, "The Pit of Tattooing", and also translated as "the Shadowlands"), is a non-administrative geographical region of New Zealand in the south-western corner of the South Island, comprising the western third of Southland. Most of F ...
bush in 1887, and again on a Fiordland beach when she was 17 years old. She claimed that her brother had also seen a moa on another occasion. In childhood, Mackenzie saw a large bird that she believed to be a
takahē The South Island takahē (''Porphyrio hochstetteri'') is a Flightless bird, flightless swamphen indigenous to New Zealand and the largest living member of the Rail (bird), rail family. It is often known by the abbreviated name takahē, whic ...
, but after its rediscovery in the 1940s, she saw a picture of it and concluded that she had seen something else. Some authors have speculated that a few ''Megalapteryx didinus'' may have persisted in remote corners of New Zealand until the 18th and even 19th centuries, but this view is not widely accepted.Anderson (1989) Some Māori hunters claimed to be in pursuit of the moa as late as the 1770s; however, these accounts possibly did not refer to the hunting of actual birds as much as a now-lost ritual among South Islanders.
Whaler A whaler or whaling ship is a specialized vessel, designed or adapted for whaling: the catching or processing of whales. Terminology The term ''whaler'' is mostly historic. A handful of nations continue with industrial whaling, and one, Jap ...
s and
sealers Sealer may refer either to a person or ship engaged in seal hunting, or to a sealant; associated terms include: Seal hunting * Sealer Hill, South Shetland Islands, Antarctica * Sealers' Oven, bread oven of mud and stone built by sealers around 1800 ...
recalled seeing monstrous birds along the coast of the South Island, and in the 1820s, a man named George Pauley made an unverified claim of seeing a moa in the Otago region of New Zealand. Occasional speculation since at least the late 19th century, and as recently as 2008, has suggested that some moa may still exist, particularly in the wilderness of
South Westland Westland District is a territorial authority district on the West Coast of New Zealand's South Island. It is administered by the Westland District Council. The district's population is History Westland was originally a part of Canterbury Pr ...
and
Fiordland Fiordland (, "The Pit of Tattooing", and also translated as "the Shadowlands"), is a non-administrative geographical region of New Zealand in the south-western corner of the South Island, comprising the western third of Southland. Most of F ...
. A 1993 report initially interested the Department of Conservation, but the animal in a blurry photograph was identified as a
red deer The red deer (''Cervus elaphus'') is one of the largest deer species. A male red deer is called a stag or Hart (deer), hart, and a female is called a doe or hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Ir ...
.
Cryptozoologists Cryptozoology is a pseudoscience and subculture that searches for and studies unknown, legendary, or extinct animals whose present existence is disputed or unsubstantiated, particularly those popular in folklore, such as Bigfoot, the Loch Ness M ...
continue to search for them, but their claims and supporting evidence (such as of purported footprints)Laing, Doug (2008) have earned little attention from experts and are
pseudoscientific Pseudoscience consists of statements, beliefs, or practices that claim to be both scientific and factual but are incompatible with the scientific method. Pseudoscience is often characterized by contradictory, exaggerated or unfalsifiable cl ...
. The rediscovery of the takahē in 1948 after none had been seen since 1898 showed that rare birds can exist undiscovered for a long time. However, the takahē is a much smaller bird than the moa, and was rediscovered after its tracks were identified—yet no reliable evidence of moa tracks has ever been found, and experts still contend that moa survival is extremely unlikely, since they would have to be living unnoticed for over 500 years in a region visited often by hunters and hikers.


Research history

Joel Polack, a trader who lived on the East Coast of the North Island from 1834 to 1837, recorded in 1838 that he had been shown "several large fossil ossifications" found near Mt Hikurangi. He was certain that these were the bones of a species of emu or ostrich, noting that "the Natives add that in times long past they received the traditions that very large birds had existed, but the scarcity of animal food, as well as the easy method of entrapping them, has caused their extermination". Polack further noted that he had received reports from Māori that a "species of
Struthio Ostriches are large flightless birds. Two living species are recognised, the common ostrich, native to large parts of sub-Saharan Africa, and the Somali ostrich, native to the Horn of Africa. They are the heaviest and largest living birds, w ...
" still existed in remote parts of the South Island. Dieffenbach also refers to a fossil from the area near Mt Hikurangi, and surmises that it belongs to "a bird, now extinct, called Moa (or Movie) by the natives". 'Movie' is the first transcribed name for the bird. In 1839, John W. Harris, a
Poverty Bay Poverty Bay (Māori language, Māori: ''Tūranganui-a-Kiwa''), officially named Tūranganui-a-Kiwa / Poverty Bay, is the largest of several small bays on the east coast of New Zealand's North Island to the north of Hawke Bay. It stretches for ...
flax trader who was a natural-history enthusiast, was given a piece of unusual bone by a Māori who had found it in a river bank. He showed the fragment of bone to his uncle, John Rule, a Sydney surgeon, who sent it to
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...
, who at that time was working at the Hunterian Museum at the
Royal College of Surgeons The Royal College of Surgeons is an ancient college (a form of corporation) established in England to regulate the activity of surgeons. Derivative organisations survive in many present and former members of the Commonwealth. These organisations ...
in London. Owen puzzled over the fragment for almost four years. He established it was part of the
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
of a big animal, but it was uncharacteristically light and honeycombed. Owen announced to a skeptical scientific community and the world that it was from a giant extinct bird like an
ostrich Ostriches are large flightless birds. Two living species are recognised, the common ostrich, native to large parts of sub-Saharan Africa, and the Somali ostrich, native to the Horn of Africa. They are the heaviest and largest living birds, w ...
, and named it ''Dinornis''. His deduction was ridiculed in some quarters, but was proved correct with the subsequent discoveries of considerable quantities of moa bones throughout the country, sufficient to reconstruct skeletons of the birds.Fuller, Errol (1987) In July 2004, the
Natural History Museum A natural history museum or museum of natural history is a scientific institution with natural history scientific collection, collections that include current and historical records of animals, plants, Fungus, fungi, ecosystems, geology, paleo ...
in London placed on display the moa bone fragment Owen had first examined, to celebrate 200 years since his birth, and in memory of Owen as founder of the museum. Since the discovery of the first moa bones in the late 1830s, thousands more have been found. They occur in a range of late
Quaternary The Quaternary ( ) is the current and most recent of the three periods of the Cenozoic Era in the geologic time scale of the International Commission on Stratigraphy (ICS), as well as the current and most recent of the twelve periods of the ...
and
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
sedimentary Sedimentary rocks are types of rock formed by the cementation of sediments—i.e. particles made of minerals (geological detritus) or organic matter (biological detritus)—that have been accumulated or deposited at Earth's surface. Sedime ...
deposits, but are most common in three main types of site:
cave Caves or caverns are natural voids under the Earth's Planetary surface, surface. Caves often form by the weathering of rock and often extend deep underground. Exogene caves are smaller openings that extend a relatively short distance undergrou ...
s,
dune A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
s, and
swamp A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...
s. Bones are commonly found in caves or ''tomo'' (the Māori word for doline or
sinkhole A sinkhole is a depression or hole in the ground caused by some form of collapse of the surface layer. The term is sometimes used to refer to doline, enclosed depressions that are also known as shakeholes, and to openings where surface water ...
, often used to refer to pitfalls or vertical cave shafts). The two main ways that the moa bones were deposited in such sites were birds that entered the cave to nest or escape bad weather, and subsequently died in the cave and birds that fell into a vertical shaft and were unable to escape. Moa bones (and the bones of other extinct birds) have been found in caves throughout New Zealand, especially in the
limestone Limestone is a type of carbonate rock, carbonate sedimentary rock which is the main source of the material Lime (material), lime. It is composed mostly of the minerals calcite and aragonite, which are different Polymorphism (materials science) ...
/
marble Marble is a metamorphic rock consisting of carbonate minerals (most commonly calcite (CaCO3) or Dolomite (mineral), dolomite (CaMg(CO3)2) that have recrystallized under the influence of heat and pressure. It has a crystalline texture, and is ty ...
areas of northwest Nelson,
Karamea Karamea is a town on the West Coast, New Zealand, West Coast of the South Island of New Zealand. It is the northernmost settlement of any real size on the West Coast, and is located northeast by road from Westport, New Zealand, Westport. Apart ...
,
Waitomo Waitomo is a rural community in the King Country region of New Zealand's North Island. There are several solutional cave systems in the area around the village, which are popular tourist attractions. Restaurants and accommodation are centred in ...
, and
Te Anau Te Anau is a town in the Southland, New Zealand, Southland List of regions in New Zealand, region of the South Island of New Zealand. In Māori language, Māori, Te-Anau means the Place of the Swirling Waters. It is on the eastern shore of Lake ...
. Moa bones and eggshell fragments sometimes occur in active coastal sand dunes, where they may erode from
paleosol In Earth science, geoscience, paleosol (''palaeosol'' in Great Britain and Australia) is an ancient soil that formed in the past. The definition of the term in geology and paleontology is slightly different from its use in soil science. In geo ...
s and concentrate in ' blowouts' between dune ridges. Many such moa bones antedate human settlement, although some originate from Māori midden sites, which frequently occur in dunes near harbours and river mouths (for example the large moa hunter sites at Shag River, Otago, and
Wairau Bar The Wairau Bar, or Te Pokohiwi, is a gravel bar formed where the Wairau River meets the sea in Cloudy Bay, Marlborough, north-eastern South Island, New Zealand. It is an important archaeological site, settled by explorers from East Polynesia ...
,
Marlborough Marlborough or the Marlborough may refer to: Places Australia * Marlborough, Queensland * Principality of Marlborough, a short-lived micronation in 1993 * Marlborough Highway, Tasmania; Malborough was an historic name for the place at the sou ...
). Densely intermingled moa bones have been encountered in swamps throughout New Zealand. The most well-known example is at Pyramid Valley in north Canterbury, where bones from at least 183 individual moa have been excavated, mostly by
Roger Duff Roger Shepherd Duff (11 July 1912 – 30 October 1978) was a New Zealand ethnologist and museum director. Biography Duff was born in Invercargill, New Zealand, on 11 July 1912. He was the son of Oliver Duff, the founding editor of the New Z ...
of Canterbury Museum. Many explanations have been proposed to account for how these deposits formed, ranging from poisonous spring waters to floods and wildfires. However, the currently accepted explanation is that the bones accumulated slowly over thousands of years, from birds that entered the swamps to feed and became trapped in the soft sediment. Many New Zealand and international museums hold moa bone collections.
Auckland War Memorial Museum The Auckland War Memorial Museum (), also known as Auckland Museum, is one of New Zealand's most important museums and war memorials. Its neoclassical architecture, neoclassical building constructed in the 1920s and 1950s, stands on Observatory ...
– Tāmaki Paenga Hira has a significant collection, and in 2018 several moa skeletons were imaged and 3D scanned to make the collections more accessible. There is also a major collection in
Otago Museum Otago (, ; ) is a region of New Zealand located in the southern half of the South Island and administered by the Otago Regional Council. It has an area of approximately , making it the country's second largest local government region. Its po ...
in
Dunedin Dunedin ( ; ) is the second-most populous city in the South Island of New Zealand (after Christchurch), and the principal city of the Otago region. Its name comes from ("fort of Edin"), the Scottish Gaelic name for Edinburgh, the capital of S ...
.


In literature and culture

Heinrich Harder portrayed moa being hunted by Māori in the classic German collecting cards about extinct and prehistoric animals, "Tiere der Urwelt", in the early 1900s. The moa was the most commonly used animal as a symbol of New Zealand before it was replaced by the kiwi in the early 20th century.
Allen Curnow Thomas Allen Monro Curnow (17 June 1911 – 23 September 2001) was a New Zealand poet and journalist. Life Curnow was born in Timaru, New Zealand, the son of a fourth generation New Zealander, an Anglican clergyman, and he grew up in a relig ...
's poem, "The Skeleton of the Great Moa in the Canterbury Museum, Christchurch" was published in 1943.


Potential revival

The creature has frequently been mentioned as a potential candidate for revival by cloning. Its iconic status, coupled with the facts that it only became extinct a few hundred years ago and that substantial quantities of moa remains exist, mean that it is often listed alongside such creatures as the
dodo The dodo (''Raphus cucullatus'') is an extinction, extinct flightless bird that was endemism, endemic to the island of Mauritius, which is east of Madagascar in the Indian Ocean. The dodo's closest relative was the also-extinct and flightles ...
as leading candidates for
de-extinction De-extinction (also known as resurrection biology, or species revivalism) is the process of generating an organism that either resembles or is an Extinction, extinct organism. There are several ways to carry out the process of de-extinction. Cloni ...
. Preliminary work involving the extraction of
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
has been undertaken by Japanese geneticist Ankoh Yasuyuki Shirota. Interest in the moa's potential for revival was further stirred in mid-2014 when New Zealand Member of Parliament
Trevor Mallard Sir Trevor Colin Mallard (born 17 June 1954) is a New Zealand politician and diplomat. Since 2023, he has served as Ambassador of New Zealand to Ireland. He was a Member of Parliament from 1984 to 1990 and again from 1993 to 2022. He served a ...
suggested that bringing back some smaller species of moa within 50 years was a viable idea. The idea was ridiculed by many, but gained support from some natural history experts. In July 2025, American biotechnology
Colossal Biosciences Colossal Biosciences Inc. is an American biotechnology and genetic engineering company working to de-extinct several extinct animals, including the woolly mammoth, the Tasmanian tiger, the northern white rhinoceros, the dire wolf, and the dodo. ...
announced early phases of plans to "revive" the South Island giant moa by adding moa genes to a related species in collaboration with the Ngāi Tahu Research Centre with funding from
Peter Jackson Sir Peter Robert Jackson (born 31 October 1961) is a New Zealand filmmaker. He is best known as the director, writer, and producer of the ''Lord of the Rings'' trilogy (2001–2003) and the ''Hobbit'' trilogy (2012–2014), both of which ar ...
.


See also

* List of New Zealand species extinct in the Holocene *
Moa-nalo The moa-nalo are a group of extinct aberrant, goose-like ducks that lived on the larger Hawaiian Islands, except Hawaii (island), Hawaii itself, in the Pacific. They were the major herbivores on most of these islands until they became extinct a ...
, several flightless ducks from the Hawaiian Islands that grew to be as large as geese. *
Elephant bird 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. Eleph ...
s, flightless
ratite Ratites () are a polyphyletic group consisting of all birds within the infraclass Palaeognathae that lack keels and cannot fly. They are mostly large, long-necked, and long-legged, the exception being the kiwi, which is also the only nocturnal ...
s up to over 3 metres tall that once lived on the island of
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
. General: *
Late Quaternary prehistoric birds Late Quaternary prehistoric birds are Bird, avian taxa that became extinct during the Late Quaternary – the Late Pleistocene or Early Holocene – and before recorded history, specifically before they could be studied alive by orni ...
*
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 "is ...
*
Megafauna In zoology, megafauna (from Ancient Greek, Greek μέγας ''megas'' "large" and Neo-Latin ''fauna'' "animal life") are large animals. The precise definition of the term varies widely, though a common threshold is approximately , this lower en ...


Footnotes


Notes


References

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


External links


TerraNature list of New Zealand's extinct birds



Tree of Life classification and references

Moa article
in Te Ara – the Encyclopedia of New Zealand
3D model of a moa skull
{{Authority control * Extinct flightless birds Endemic birds of New Zealand Late Quaternary prehistoric birds Extinct birds of New Zealand Ratites Bird extinctions since 1500 Holocene extinctions Species made extinct by human activities Animals with only two limbs Miocene first appearances Taxa named by Charles Lucien Bonaparte Notopalaeognathae