Moa (

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

Dinornithiformes) are an extinct group 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 formerly endemic to New Zealand.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 Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division of ...

Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togethe ...

, there were nine

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

(in six genera). The two largest species, ''

Dinornis robustus The South Island giant moa (''Dinornis robustus'') is an extinct moa from the genus ''Dinornis.'' Context The moa were ratites, flightless birds with a sternum without a keel. They also had a distinctive palate. The origin of these birds is b ...

'' 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 Extinction, extinct species of moa from the family Lesser moa, Emeidae (lesser moa). Description It was the smallest known species of moa, only slightly taller ...

(''Anomalopteryx didiformis''), was around the size of a

turkey Turkey ( tr, Türkiye ), officially the Republic of Türkiye ( tr, Türkiye Cumhuriyeti, links=no ), is a list of transcontinental countries, transcontinental country located mainly on the Anatolia, Anatolian Peninsula in Western Asia, with ...

. Estimates of the moa population when

Polynesians Polynesians form an ethnolinguistic group of closely related people who are native to Polynesia (islands in the Polynesian Triangle), an expansive region of Oceania in the Pacific Ocean. They trace their early prehistoric origins to Island Sou ...

settled New Zealand circa 1300 vary between 58,000 and approximately 2.5 million. Moa are traditionally placed in the

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 ...

group. However, genetic studies have found that their closest relatives are the flighted South American

tinamou Tinamous () form an order of birds called Tinamiformes (), comprising a single family called Tinamidae (), divided into two distinct subfamilies, containing 46 species found in Mexico, Central America, and South America. The word "tinamou" come ...

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 t ...

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 adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

s in New Zealand's forest, shrubland, and subalpine

ecosystems An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...

until the arrival of the

Māori Māori or Maori can refer to: Relating to the Māori people * Māori people of New Zealand, or members of that group * Māori language, the language of the Māori people of New Zealand * Māori culture * Cook Islanders, the Māori people of the C ...

, and were hunted only by the

Haast's eagle Haast's eagle (''Hieraaetus moorei'') is an extinct species of eagle that once lived in the South Island of New Zealand, commonly accepted to be the pouakai of Māori legend.

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 accou ...

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), 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 ...

. 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 art is 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 also ...

depicts moa or moa-like birds (likely

geese A goose (plural, : geese) is a bird of any of several waterfowl species in the family (biology), family Anatidae. This group comprises the genera ''Anser (bird), Anser'' (the grey geese and white geese) and ''Branta'' (the black geese). Some o ...

adzebill The adzebills, genus ''Aptornis'', were two closely related bird species, the North Island adzebill, (''Aptornis otidiformis''), and the South Island adzebill, (''Aptornis defossor''), of the extinct family Aptornithidae. The family was endemic ...

s) with necks upright, indicating that moa were more than capable of assuming both neck postures. No records survive of what sounds moa made, though some idea of their

calls Call or Calls may refer to: Arts, entertainment, and media Games * Call, a type of betting in poker * Call, in the game of contract bridge, a bid, pass, double, or redouble in the bidding stage Music and dance * Call (band), from Lahore, Paki ...

can be gained from fossil evidence. The

trachea The trachea, also known as the windpipe, is a Cartilage, cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends ...

of moa were supported by many small rings of bone known as tracheal rings. Excavation of these rings from articulated skeletons has shown that at least two moa genera (''Euryapteryx'' and ''Emeus'') exhibited tracheal elongation, that is, their trachea were up to 1 m (3 ft) long and formed a large loop within the body cavity. 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 family (biology), family Anatidae within the genus ''Cygnus''. 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 t ...

s, cranes, and

guinea fowl 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 feature is associated with deep resonant vocalisations that can travel long distances.

Evolutionary relationships

The moa's closest relatives are small terrestrial South American birds called the tinamous, which can fly.Phillips, et al. (2010) Previously, the

, the Australian

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' ...

, and

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 ...

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 exactly 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 all ...

. Currently, 11 species are formally recognised, although recent studies using

ancient DNA Ancient DNA (aDNA) is DNA isolated from ancient specimens. Due to degradation processes (including cross-linking, deamination and fragmentation) ancient DNA is more degraded in comparison with contemporary genetic material. Even under the bes ...

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 Bergmann's rule is an ecogeographical rule that states that within a broadly distributed taxonomic clade, populations and species of larger size are found in colder environments, while populations and species of smaller size are found in warmer r ...

and Allen’s rule) as well as

sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...

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 North I ...

'' 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 circl ...

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 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 ...

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) period, ...

. Known from multiple eggshells and hind limb elements, these represent at least two already fairly large-sized species.

Classification

Taxonomy

The currently recognised genera and species are: *Order †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 North I ...

Owen 1843 alapteryginae Bonaparte 1854; Palapterygidae Haast 1874; Dinornithnideae Stejneger 1884(giant moa) ***Genus ''

'' ****

North Island giant moa The North Island giant moa (''Dinornis novaezealandiae'') is an extinct moa in the genus ''Dinornis''. Even though it might have walked with a lowered posture, standing upright, it would have been the tallest bird ever to exist, with a height ...

, ''Dinornis novaezealandiae'' (North Island, New Zealand) ****

South Island giant moa The South Island giant moa (''Dinornis robustus'') is an extinct moa from the genus ''Dinornis.'' Context The moa were Ratite, ratites, flightless birds with a sternum without a Keel (bird anatomy), keel. They also had a distinctive palate. T ...

, ''Dinornis robustus'' (South Island, New Zealand) **Family Emeidae (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 Extinction, extinct species of moa from the family Lesser moa, Emeidae (lesser moa). Description It was the smallest known species of moa, only slightly taller ...

, ''Anomalopteryx didiformis'' (North and South Island, New Zealand) ***Genus '' Emeus'' ****

Eastern moa The eastern moa (''Emeus crassus'') is an extinct species of moa. When the first specimens were originally described by Richard Owen, they were placed within the genus ''Dinornis'' as three different species, but, was later split off into their ...

, ''Emeus crassus'' (South Island, New Zealand) ***Genus ''

Euryapteryx The broad-billed, stout-legged moa or coastal moa (''Euryapteryx curtus'') is an extinct species of moa. These moa lived in both the North Island, North and the South Island, South Islands of New Zealand, and on Stewart Island. Its habitat was ...

'' **** Broad-billed moa, ''Euryapteryx curtus'' (North and South Island, New Zealand) ***Genus ''

Pachyornis ''Pachyornis'' is an extinct genus of ratites from New Zealand which belonged to the moa family. Like all ratites it was a member of the Order (biology), order Struthioniformes. The Struthioniformes are flightless birds with a sternum without a k ...

'' ****

Heavy-footed moa The heavy-footed moa (''Pachyornis elephantopus'') is a species of moa from the lesser moa family. The heavy-footed moa was widespread only in the South Island of New Zealand, and its habitat was the lowlands (shrublands, dunelands, grasslands, ...

, ''Pachyornis elephantopus'' (South Island, New Zealand) ****

Mantell's moa Mantell's moa (''Pachyornis geranoides'') also known as Mappin's moa is an extinct species of moa from the North Island of New Zealand. Its habitat was the lowlands (shrublands, grasslands, dunelands, and forests). The moa were ratites, flight ...

, ''Pachyornis geranoides'' (North Island, New Zealand) **** Crested moa, '' Pachyornis australis'' (South Island, New Zealand)Stephenson, Brent (2009) **Family Megalapterygidae ***

Genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

Megalapteryx The upland moa (''Megalapteryx didinus'') was a species of moa endemic to New Zealand. It was a ratite, a grouping of flightless birds with no keel on the sternum. It was the last moa species to become extinct, vanishing in 1445 CE, and was pr ...

'' ****

Upland moa The upland moa (''Megalapteryx didinus'') was a species of moa endemic to New Zealand. It was a ratite, a grouping of flightless birds with no keel on the sternum. It was the last moa species to become extinct, vanishing in 1445 CE, and was pr ...

, ''Megalapteryx didinus'' (South Island, New Zealand) Two unnamed species from the Saint Bathans Fauna.

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

-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 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 ...

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) is a mountain range extending along much of the length of New Zealand's South Island, reaching its greatest elevations near the range's western side. The name "Southern ...

about 6 Mya, and the habitat fragmentation on both islands resulting from Pleistocene glacial cycles,

volcanism Volcanism, vulcanism or volcanicity is the phenomenon of eruption of molten rock (magma) onto the surface of the Earth or a solid-surface planet or moon, where lava, pyroclastics, and volcanic gases erupt through a break in the surface called ...

, 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:

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 in southern South America (Chile, Argentina) and Australasia (east and southeast Australia, New Zealand, New Gui ...

'') 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

that included ''Pachyornis elephantopus'' (heavy-footed moa), ''Euryapteryx gravis'', ''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 (also kitchen midden or shell heap) is an old dump for domestic waste which may consist of animal bone, human excrement, botanical material, mollusc shells, potsherds, lithics (especially debitage), and other artifacts and ecofact ...

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 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. Apart from a narrow coastal strip, the town of Karam ...

districts (such as

Honeycomb Hill Cave Honeycomb Hill Cave is located in the Oparara Basin, on the northwestern edge of the South Island of New Zealand. The Oparara River flows through a section of the cave. The cave was discovered in 1976 by the Buller Caving Group. Since then, 14 ...

), 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/Mata-Au and is the gateway to Mount Aspiring National Park. ...

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 of the South Island of New Zealand. It is located between Westport and Greymouth on , the only through-road on the West Coast. Punakaiki is immediately adjacent to Paparoa National Park, and is a ...

on the west coast and

Central Otago Central Otago is 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 and tributa ...

) 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. It is located on State Highway 1, 180 km north of Christchurch. The town has an estimated permanent resident population of (as of ). The town is the government ...

Otago Peninsula The Otago Peninsula ( mi, Muaūpoko) 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 sou ...

, 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 a paucity 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 gravis'', ''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 gree ...

had made a land bridge across the

Cook Strait Cook Strait ( mi, Te Moana-o-Raukawa) separates the 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, A H, ...

. 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 gravis'', ''E. curtus'', and ''Pachyornis geranoides'') tended to inhabit drier forest and shrubland habitats. ''P. geranoides'' occurred throughout the North Island. The distributions of ''E. gravis'' and ''E. curtus'' were almost mutually exclusive, the former having only been found in coastal sites around the southern half of the North Island.

Behaviour and ecology

About eight moa

trackway Historic roads (historic trails in USA 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 trackway ...

s, with fossilised moa footprint impressions in fluvial silts, have been found in the North Island, including

Waikanae Waikanae (, ) is a town on the Kapiti Coast, 60 kilometres north of the Wellington CBD. The name is a Māori word meaning "waters" (''wai'') "of the grey mullet". The town lies between Paraparaumu, eight kilometres to the southwest, and Ōtak ...

Creek (1872),

Napier Napier may refer to: People * Napier (surname), including a list of people with that name * Napier baronets, five baronetcies and lists of the title holders Given name * Napier Shaw (1854–1945), British meteorologist * Napier Waller (1893–19 ...

(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 Foxton, ...

(1895), Marton (1896),

Palmerston North Palmerston North (; mi, Te Papa-i-Oea, known colloquially as Palmy) is a city in the North Island of New Zealand and the seat of the Manawatū-Whanganui region. Located in the eastern Manawatu Plains, the city is near the north bank of the ...

(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 Bu ...

(1939), and under water in

Lake Taupō Lake Taupō (also spelled Taupo; mi, Taupō-nui-a-Tia or ) is a large crater lake in New Zealand's North Island, located in the caldera of the Taupō Volcano. The lake is the namesake of the town of Taupō, which sits on a bay in the lake's nor ...

(1973). Analysis of the spacing of these tracks indicates walking speeds between 3 and 5 km/h (1.75–3 mph).

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 preserved ...

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 (pterosaurs, crocodiles, alligators, dinosaurs, birds), earthworms, some gastropods, so ...

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 is de ...

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 ''

elephantopus'' was analogous to a pair of secateurs, 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 n ...

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 resources and competitors (for ...

occupied in other countries by large browsing mammals such as

antelope The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia. Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mammals ...

and

llamas The llama (; ) (''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 soft ...

. Some biologists contend that a number of plant species evolved to avoid moa browsing. Divaracating plants such as ''

Pennantia corymbosa ''Pennantia corymbosa'', commonly known as kaikomako (from the Māori ), is a small dioecious forest tree of New Zealand. Small creamy, white flowers are produced between November and February, followed by a shiny black fruit in autumn. They ar ...

'' (the kaikōmako), which have small leaves and a dense mesh of branches, and ''

Pseudopanax crassifolius ''Pseudopanax crassifolius'', the horoeka or lancewood, is a New Zealand native tree belonging to the family Araliaceae. It is found throughout New Zealand from sea level up to about 750 m. The juvenile form, which lasts for between 15 and ...

'' (the horoeka or lancewood), which has tough juvenile leaves, are possible examples of plants that evolved in such a way. 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 othe ...

s), which were retained in their muscular gizzards, providing a grinding action that allowed them to eat coarse plant material. 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, ''r''/''K'' selection theory relates to the selection of combinations of traits in an organism that trade off between quantity and quality of offspring. The focus on either an increased quantity of offspring at the expense of individ ...

, 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 natural capability to pr ...

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 A cave or cavern is a natural void in the ground, specifically a space large enough for a human to enter. Caves often form by the weathering of rock and often extend deep underground. The word ''cave'' can refer to smaller openings such as sea ...

and rock shelters, little evidence exists of the

nest A nest is a structure built for certain animals to hold 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 organic materia ...

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 highly vesicular rough-textured volcanic glass, which may or may not contain crystals. It is typically light-colored. Scoria is another vesicular vol ...

". Moa nesting material has also been recovered from rock shelters in the

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)

Seeds A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosperm pl ...

and

pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...

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 scientific study of human activity through the recovery and analysis of material culture. The archaeological record consists of artifacts, architecture, biofacts or ecofacts, sites, and cultural landscap ...

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, fl ...

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. File:The skeleton of female upland moa.JPG, The skeleton of female upland moa with egg in unlaid position within the pelvic cavity in Otago Museum File:Emeus egg and embryo.jpg, An egg and embryo fragments of ''Emeus crassus'' File:Megalapteryx.png, Restoration of an

upland moa The upland moa (''Megalapteryx didinus'') was a species of moa endemic to New Zealand. It was a ratite, a grouping of flightless birds with no keel on the sternum. It was the last moa species to become extinct, vanishing in 1445 CE, and was pr ...

Pre-human forests

Studies of accumulated dried vegetation in the pre-human mid-late

period suggests a low ''

Sophora microphylla ''Sophora microphylla'', common name kōwhai, is a species of flowering plant in the family Fabaceae, native to New Zealand. Growing to tall and broad, it is an evergreen shrub or small tree. Each leaf is long with up to 40 pairs of shiny oval ...

'' or Kōwai forest ecosystem in

that was used and perhaps maintained by moa, for both nesting material and food. Neither the forests nor moa existed when European settlers came to the area in the 1850s.

Relationship with humans

Extinction

Before the arrival of humans, the moa's only predator was the massive

Haast's eagle Haast's eagle (''Hieraaetus moorei'') is an extinct species of eagle that once lived in the South Island of New Zealand, commonly accepted to be the pouakai of Māori legend.carbon-14 dating Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for determining the age of an object containing organic material by using the properties of radiocarbon, a radioactive isotope of carbon. The method was dev ...

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, at the northern end of New Zealand's South Island, located on the lower reaches of the Tākaka River. State Highway 60 runs through Takaka and follows the river valley b ...

and

Riwaka Riwaka ( mi, Riuwaka) 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 tow ...

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 introduction ...

'' published an additional account from a farmer and his shepherd. An 80-year-old woman, Alice McKenzie, claimed in 1959 that she had seen a moa in

Fiordland Fiordland is a geographical region of New Zealand in the south-western corner of the South Island, comprising the westernmost third of Southland. Most of Fiordland is dominated by the steep sides of the snow-capped Southern Alps, deep lakes, ...

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 swamphen indigenous to New Zealand and the largest living member of the rail family. It is often known by the abbreviated name takahē, which it shares with the recently ...

, 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, Japa ...

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 180 ...

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 Prov ...

and

. 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, and a female is called a hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Iran, and parts of wes ...

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 claim ...

. 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.

Surviving remains

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 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 ...

" 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: ''Tūranganui-a-Kiwa'') 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 from Young Nick's Head in the southwest to Tuaheni Point in the north ...

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 anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Owe ...

, 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 a ...

in London. Owen puzzled over the fragment for almost four years. He established it was part of the

femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...

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 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 ...

, 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 collections that include current and historical records of animals, plants, fungi, ecosystems, geology, paleontology, climatology, and more. ...

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. Kapua Swamp

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). It follows the Neogene Period and spans from 2.58 million years ...

and

sedimentary Sedimentary rocks are types of rock (geology), rock that are formed by the accumulation or deposition of mineral or organic matter, organic particles at Earth#Surface, Earth's surface, followed by cementation (geology), cementation. Sedimentati ...

deposits, but are most common in three main types of site:

cave A cave or cavern is a natural void in the ground, specifically a space large enough for a human to enter. Caves often form by the weathering of rock and often extend deep underground. The word ''cave'' can refer to smaller openings such as sea ...

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, f ...

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 locally also known as ''vrtače'' and shakeholes, and to openi ...

, 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 ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...

marble Marble is a metamorphic rock composed of recrystallized carbonate minerals, most commonly calcite or Dolomite (mineral), dolomite. Marble is typically not Foliation (geology), foliated (layered), although there are exceptions. In geology, the ...

areas of northwest Nelson,

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 region of the South Island of New Zealand. In Maori, Te-Anau means the Place of the Swirling Waters. It is on the eastern shore of Lake Te Anau in Fiordland. Te Anau is 155 kilometres north of Invercargill an ...

. Moa bones and eggshell fragments sometimes occur in active coastal sand dunes, where they may erode from

paleosol In the geosciences, paleosol (''palaeosol'' in Great Britain and Australia) is an ancient soil that formed in the past. The precise definition of the term in geology and paleontology is slightly different from its use in soil science. In geolo ...

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,

Marlborough Marlborough may refer to: Places United Kingdom * Marlborough, Wiltshire, England ** Marlborough College, public school * Marlborough School, Woodstock in Oxfordshire, England * The Marlborough Science Academy in Hertfordshire, England Austral ...

). 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 in 1912. He is the son of Oliver Duff founding editor of the New Zealand Listener, and ...

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 Tāmaki Paenga Hira (or simply the Auckland Museum) is one of New Zealand's most important museums and war memorials. Its collections concentrate on New Zealand history (and especially the history of the Aucklan ...

– 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 Tūhura Otago Museum is located in the city centre of Dunedin, New Zealand. It is adjacent to the University of Otago campus in Dunedin North, 1,500 metres northeast of the city centre. It is one of the city's leading attractions and has one of t ...

Dunedin Dunedin ( ; mi, Ōtepoti) is the second-largest city in the South Island of New Zealand (after Christchurch), and the principal city of the Otago region. Its name comes from , the Scottish Gaelic name for Edinburgh, the capital of Scotland. Th ...

Feathers and soft tissues

Several remarkable examples of moa remains have been found which exhibit soft tissues (

muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...

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 cuticle, animal coverings, such as the arthropod exoskeleton, have diffe ...

feathers Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier e ...

), that were 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. ...

when the bird died in a naturally dry site (for example, a cave with a constant dry breeze blowing through it). Most of these specimens have been 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 semi-ar ...

region, the driest part of New Zealand. 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 t ...

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

) *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 type specimen of ''Megalapteryx didinus'' found near Queenstown in 1878 (currently held by

, 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 , mottoeng = Literal: From here, light and sacred draughts. Non literal: From this place, we gain enlightenment and precious knowledge. , established = , other_name = The Chancellor, Masters and Schola ...

) * The complete feathered leg of a ''M. didinus'' from Old Man Range in 1894 (currently held by

) * The head of a ''M. didinus'' found near Cromwell sometime before 1949 (currently held by the

Museum of New Zealand The Museum of New Zealand Te Papa Tongarewa is New Zealand's national museum and is located in Wellington. ''Te Papa Tongarewa'' translates literally to "container of treasures" or in full "container of treasured things and people that spring fr ...

). Two specimens are known from outside the Central Otago region: * A complete foot of ''M. didinus'' found in a cave on Mount Owen near Nelson in the 1980s (currently held by the

) * A skeleton of ''Anomalopteryx didiformis'' with muscle, skin, and feather bases collected from a cave near

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.

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 extinct flightless bird that was endemic to the island of Mauritius, which is east of Madagascar in the Indian Ocean. The dodo's closest genetic relative was the also-extinct Rodrigues solitaire. The ...

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 extinct species. There are several ways to carry out the process of de-extinction. Cloning is th ...

. Preliminary work involving the extraction of DNA 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 Trevor Colin Mallard (born 17 June 1954) is a New Zealand politician and member of the Labour Party. First elected to Parliament in 1984, he was Speaker of the New Zealand House of Representatives from 2017 until 2022. Mallard was a Cabinet m ...

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 literature and culture

Heinrich Harder Heinrich Harder (2 June 1858 – 5 February 1935) was a German artist and an art professor at the Prussian Academy of Arts in Berlin best known for his depictions of extinct animals. Life Heinrich Harder was born in Putzar, Pomerania, the son ...

portrayed moa being hunted by Māori in the classic German collecting cards about extinct and prehistoric animals, "Tiere der Urwelt", in the early 1900s.

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.Curnow, Allen (1944). ''Sailing or Drowning''. Wellington: Progressive Publishing Society.

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 Higher-level bird taxa restricted to 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

Etymology

Description

Evolutionary relationships

Classification

Taxonomy

Phylogeny

Distribution and habitat

South Island

North Island

Behaviour and ecology

Diet

Reproduction

Pre-human forests

Relationship with humans

Extinction

Surviving remains

Feathers and soft tissues

Potential revival

In literature and culture

See also

Footnotes

Notes

References

External links