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The North Island giant moa (''Dinornis novaezealandiae'') is an extinct
moa Moa are extinct giant flightless birds native to New Zealand. The term has also come to be used for chicken in many Polynesian cultures and is found in the names of many chicken recipes, such as Kale moa and Moa Samoa. Moa or MOA may also refe ...
in the
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 ...
''
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 ...
''. 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 estimated up to 3.6 metres (12 feet).


Taxonomy

It was a
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 ...
and a member of the order
Dinornithiformes Moa are extinct giant flightless birds native to New Zealand. The term has also come to be used for chicken in many Polynesian cultures and is found in the names of many chicken recipes, such as Kale moa and Moa Samoa. Moa or MOA may also refer ...
.Davies, S.J.J.F. (2003). "Moas". In Hutchins, Michael (ed.). Grzimek's Animal Life Encyclopedia. 8 Birds I Tinamous and Ratites to Hoatzins (2 ed.). Farmington Hills, MI: Gale Group. pp. 95–98. ISBN 0-7876-5784-0. The ''Dinornithiformes'' were flightless birds with a
sternum The sternum or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major blood vessels from injury. Sh ...
but without a
keel The keel is the bottom-most longitudinal structural element on a vessel. On some sailboats, it may have a hydrodynamic and counterbalancing purpose, as well. As the laying down of the keel is the initial step in the construction of a ship, in Br ...
. They also had a distinctive
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
.


Origin

The origin of ''ratites'' is becoming clearer, as it is now believed that early ancestors of these birds were able to fly (Davies, 2003). From such fossil evidence it is believed that the early flying ratites originated in the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
and flew to the Southern Hemisphere (Davies, 2003). This movement of flying ancestors is believed to be the cause of the distribution of ''ratites'' across the Southern Hemisphere (Davies, 2003).


Appearance

The ''Dinornis novaezealandiae'' has been described as a bird that was “two-legged, tailless, wingless ndclad in woolly fibres” (Armstrong 2010, p. 327). It had, “long, shaggy hair-like feathers up to 18 cm long” (Szabo, 2013).Szabo, M.J. (2013). (updated 2017). North Island giant moa. In Miskelly, C.M. (ed.) New Zealand Birds Online. Retrieved from https://nzbirdsonline.org.nz/species/north-island-giant-moa Feather and skin fragments indicate that all but the legs were fully feathered and that the wings of this bird were not visible. Their feathers were brown, sometimes with pale edging.


Anatomy

Research shows that the ''D. novaezealandiae'' displayed a large reversed sexual dimorphism whereby the females were much larger than males, estimated to have had up to twice the body mass of males (Turvey & Holdaway 2005, p. 70). ''Dinornis'' had long slim, elongated bones compared to other moa species (Turvey & Holdaway 2005, p. 75). Using algorithms derived from the bone findings of juvenile ''Dinornis'', an adult male has been estimated to weigh between 55 and 88 kg whilst females between 78 and 249 kg (Turvey & Holdaway 2005, p. 76). The growth and development of ''Dinornis'' long bones, compared to that of other ''ratites'' such as the
Ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There are ...
, has been found to be much slower (Turvey & Holdaway 2005, p. 76). The
North Island brown kiwi The North Island brown kiwi (''Apteryx mantelli''; ''Apteryx australis'' or ''Apteryx bulleri'' as before 2000, still used in some sources) is a species of kiwi that is widespread in the northern two-thirds of the North Island of New Zealand an ...
, that does not reach adult body mass until 12 months, is viewed as a more appropriate developmental analog for the ''Dinornis'' due to the similarities that have been drawn between the time taken to reach complete maturity of hindlimbs (5 years), as well as the time upon which tarsals commence fusion with adjacent long bones (4 years) (Turvey & Holdaway 2005, p. 79). Due to extensive land clearance for agriculture during the nineteenth and twentieth century, most land surface containing moa bones was lost throughout New Zealand, making the bones of the ''D. novaezealandiae'' a rarity that are only found in remote, undisturbed and rarely visited sites (Wood & Wilmshurst 2013, p. 254).


Habitat

This moa lived on the
North Island The North Island, also officially named Te Ika-a-Māui, is one of the two main islands of New Zealand, separated from the larger but much less populous South Island by the Cook Strait. The island's area is , making it the world's 14th-largest ...
of
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...
, and lived in the lowlands (
shrubland Shrubland, scrubland, scrub, brush, or bush is a plant community characterized by vegetation dominated by shrubs, often also including grasses, herbs, and geophytes. Shrubland may either occur naturally or be the result of human activity. It m ...
s,
grassland A grassland is an area where the vegetation is dominated by grasses (Poaceae). However, sedge (Cyperaceae) and rush (Juncaceae) can also be found along with variable proportions of legumes, like clover, and other herbs. Grasslands occur natur ...
s, dunelands, and forests). The ''Dinornis'' was a widely represented genus of moa in the North Island of New Zealand (Scarlett 1974, p. 1).Scarlett, R. J. (1974). Moa and man in New Zealand. Notornis, 21(1), 1-12. The habitat of the ''Dinornis novaezealandiae'' is thought to have remained relatively unchanged for hundreds of thousands of years (McCallum et al. 2013, p.e50732). This was in part due to the bird's inability to fly, as this restricted its movement between islands and therefore confined the species to the North Island of New Zealand (Lomolino et al. 2021, p. 5). Shifts in this moa's habitat have been noted during particular times of changing climates and vegetative zones (Lomolino et al. 2021, p. 1) and in reaction to the influence of anthropogenic factors such as the introduction of
Polynesian rat The Polynesian rat, Pacific rat or little rat (''Rattus exulans''), known to the Māori as ''kiore'', is the third most widespread species of rat in the world behind the brown rat and black rat. The Polynesian rat originated in Southeast Asia, a ...
(''Rattus exulans'') and
Polynesian dog The Polynesian Dog refers to a few extinct varieties of domesticated dogs from the islands of Polynesia. These dogs were used for both companionship and food and were introduced alongside poultry and pigs to various islands. They became extinct ...
(''Canis familiaris'') (Lomolino et al. 2021, p. 1). Later, the human environmental impact of fires made by the
Māori people The Māori (, ) are the indigenous Polynesian people of mainland New Zealand (). Māori originated with settlers from East Polynesia, who arrived in New Zealand in several waves of canoe voyages between roughly 1320 and 1350. Over several ce ...
also contributed to habitat alterations (Lomolino et al. 2021, p. 10). Based on recent knowledge, the Kahikatea-Pukatea-tawa forest,
Waikato Waikato () is a Regions of New Zealand, local government region of the upper North Island of New Zealand. It covers the Waikato District, Waipa District, Matamata-Piako District, South Waikato District and Hamilton, New Zealand, Hamilton City ...
, was home to the majority of ''Dinornis novaezealandiae'' (Lomolino et al. 2021, p. 5). However, bone discoveries also reveal it to have been in nearby places, such as Opito,
Auckland Auckland (pronounced ) ( mi, Tāmaki Makaurau) is a large metropolitan city in the North Island of New Zealand. The List of New Zealand urban areas by population, most populous urban area in the country and the List of cities in Oceania by po ...
, where it was the predominant genus of moa (Scarlett 1974, p. 11).


Diet

''Dinornis novaezealandie'' was one of the largest
herbivores 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 ...
in New Zealand's terrestrial ecosystem (Wood et al. 2020, p. 15). Their diet is described as diverse, consuming a wide range of plant taxa (Wood et al. 2020, p. 14). It is difficult to be certain of the exact diet that this species of moa would have eaten, as
coprolites 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 ...
and
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 ...
content for this species of moa have not yet been found (Wood et al. 2020, p. 17). However, studies propose that the diet would have been similar to that of the ''Dinornis robustus'' (
South Island giant moa 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 ...
) due to the similarities in morphology and the landscapes that they roamed (Wood et al. 2020, p. 17). This would suggest that like the ''Dinornis robustus'', the ''Dinornis novaezealandie'' diet would consist of forest trees, especially
Southern Beech ''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 ...
(''Nothofagaceae''), seeds and leaves of small shrubs in forest areas (Wood et al. 2020, p. 9). In non-forest areas there is speculation that they may have grazed on herbs (Wood et al. 2020, p. 9). There is also theories that with such a diverse range of plants restricted to the North Island, if coprolites and gizzard content for the ''Dinornis novaezealandiae'' are found, this would likely expand the number of plant taxa known to have been eaten by moa (Wood et al. 2020, P.17). Research into moa beak shapes and jaw muscle size provide some evidence for the diverse diets of moa (Baker et al. 2005, p. 8261). From such research it has been found that large ''Dinornis'' browsed primarily on coarse twigs (Baker et al. 2005, p. 8261). Further speculation suggests that in order to ferment their plant diet in accordance with their large body size, moa such as ''D. novaezealandiae'' may have evolved to have long intestines (Baker et al. 2005, p. 8261).


Behaviour and ecology

New Zealand plants and moa were in co-evolution (Wood et al. 2020, p. 2). Moa have been found to filiramulate growth habit in plants such as divarication,
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 ...
, deciduousness, spines or spine like structures (enlarged stinging hairs), leaf loss and photosynthetic stems, mimicry and reduced visual apparency, tough and fibrous leaves, distasteful compounds and low nutrient status (Wood et al. 2020, p. 2). Though moa ate flowers, it is unlikely that they contributed to
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds ...
processes (Wood et al. 2020, p. 15). On the contrary, this would have been a more destructive process than other impacts the ''Dinornis novaezealandiae'' may have had on the North Island's ecology (Wood et al. 2020, p. 15). The deep, longstanding interconnectedness between plants and moa means that the consequences of the extinction of the ''Dinornis novaezealandiae'' may still be largely unknown (Wood et al. 2020, p. 2).


Spore disposal

There are theories that due to the moa's longstanding prevalence in New Zealand's landscape, they would have come to form certain roles in New Zealand's ecology. One example that has been proposed is the possibility of their ability to spread seeds (Wood et al. 2020, p. 15). This stems from findings in moa specimen examinations that show different species of fungi that moa ingested, such as ''
Cortinarius ''Cortinarius'' is a globally distributed genus of mushrooms in the family Cortinariaceae. It is suspected to be the largest genus of agarics, containing over 2,000 widespread species. A common feature among all species in the genus ''Cortinariu ...
'', because this fungus is known in particular to be associated with higher spore dispersal by birds (Wood et al. 2020, p. 15).


Moa dung

There is also theories that moa defecation and their herbivorous diet may have contributed to nutrient spreading and cycling, though this is not easy to validate (Wood et al.2020, p. 16). However, large dung from the ''Dinornis novaezealandie'' would have likely nurtured the existence of
dung beetles Dung beetles are beetles that feed on feces. Some species of dung beetles can bury dung 250 times their own mass in one night. Many dung beetles, known as ''rollers'', roll dung into round balls, which are used as a food source or breeding cha ...
and dung mosses (
Splachnaceae Splachnaceae is a family of mosses, containing around 70 species in 6 genera. Around half of those species are entomophilous, using insects to disperse their spores, a characteristic found in no other seedless land plants. Many species in this f ...
) in the North Island (Wood et al. 2020, p. 16). Some of the dung mosses may have included those of the ''Tayloria'' genus (Wood et al. 2020, p. 16).


Moa tracks

The enormous, flightless birds caused large paths to be cleared through the landscape. It was observed in the
Poukawa Poukawa is a rural community in the Hastings District, New Zealand, Hastings District and Hawke's Bay Region of New Zealand's North Island. It is located south of Hastings, New Zealand, Hastings, off New Zealand State Highway 2, State Highway ...
region that these would often lead to freshwater springs and the bottom of rocky cliffs; where they would tend to nest and roost (Horn 1989, p. 46). Once this observation was made these paths became particularly useful for humans when searching for fresh water sources and were continued to be used for these purposes long after the moa's extinction (Horn 1989, p. 47).


Reproduction

Whole moa eggs are a rare find in archaeology (Huynen et al. 2010, p. 16201), however the abundance of fragments suggest that when fresh, the egg of the ''Dinornis novaezealandiae'' is estimated to have weighed over 3 kg and 190 x 150mm (Szabo, 2013). Ancient DNA results reveal that surfaces of the outer shell of eggs that belonged to Dinornis novaezealandiae yielded DNA from males only (Huynen et al. 2010, p. 16203). This indicates that the males were the likely incubators of eggs (Huynen et al. 2010, p. 16203). Findings also show that the inside of these eggs, and the remains on the outer surface, matched female DNA, which is thought to be from the egg laying process (Huynen et al. 2010, p. 16203). The eggs of Dinornis have also been shown to be more susceptible to breakage than any of the 3,434 avian species measured to date (Huynen et al. 2010, p. 16204). Though the male D. novaezealandiae were lighter than females, questions have been raised as to how birds of such weight, even those that were smaller, could manage to incubate the fragile eggs successfully since the possibility of breakage is many times greater than that of any other bird (Huynen et al. 2010, p. 16204). For this reason, it is unlikely that larger moa, such as the ''D. novaezealandiae'', would have been able to incubate their eggs using the same contact method that is practiced by almost all birds (Huynen et al. 2010, p. 16204). Whilst the exact structure remains unclear, it is more likely they would have formed a special nest that would support their body weight in some way (Huynen et al. 2010, p. 16204).


Relationship with humans

Some cultural depictions of moa focus on how the moa was best cooked and enjoyed as a food, such as, ''He koromiko te wahie i taona ai te moa'' (“The moa was cooked with the wood of the koromiko”) (Wehi & Cox, 2018).Wehi, P., & Cox, M. (2018). Dead as the Moa. Waikato.ac.nz. Retrieved from https://www.waikato.ac.nz/news-opinion/media/2018/dead-as-the-moa Other depictions, however, focus on the development and fate of their extinction. This is because the moa was used as a metaphor for the
Māori people The Māori (, ) are the indigenous Polynesian people of mainland New Zealand (). Māori originated with settlers from East Polynesia, who arrived in New Zealand in several waves of canoe voyages between roughly 1320 and 1350. Over several ce ...
to express fears of their own extinction that developed; as illness, disease and deforestation, by European settlers, posed severe threat to their survival (Wehi & Cox, 2018). This is seen in sayings such as ''Huna I te huna a te moa'' ("Hidden as the moa hid") and "Dead as the Moa" (Wehi & Cox, 2018), as well as depictions of moa whereby Māori describe it as “having a human face and living in a cave,” (Armstrong 2010, p. 329). Whilst hunting ''Dinornis novaezealandie'' was largely for consumption, findings have also shown that ''Dinornis'' bones were used to make many one-piece fish-hooks (Scarlett 1974, p. 11).


Extinction

The disappearance and eventually extinction of the moa occurred around the 15th c, 200 years after human settlement in New Zealand (Perry et al. 2014, p. 131). Before the settlement of humans, ''Dinornis novaezealandiae'' had few natural predators, meaning there was little threat that the species would become extinct (Huynen et al. 2014, p. 4). However, after the arrival of the
Māori people The Māori (, ) are the indigenous Polynesian people of mainland New Zealand (). Māori originated with settlers from East Polynesia, who arrived in New Zealand in several waves of canoe voyages between roughly 1320 and 1350. Over several ce ...
and their human activities of hunting and fires, humans soon became a threat to all species of moa, including ''Dinornis novaezealandiae'', as it has been found that there did not appear to be a preferred moa size for hunting (Perry et al. 2014, p. 131). Though geographically restricted to the North Island, the diversity of habitat that the moa could survive in has dismantled theories that its extinction could have been a result of habitat loss (Perry 2014, p. 133). Meanwhile, radiocarbon data shows that the vast spread of the highly mobile Māori people across the country, highly correlates with the time upon which moa populations were plummeting (Perry et al. 2014, pp. 131–134). This was the case across all geographical areas, not just those that were being deforested, but also other areas where human activities, such as hunting, were carried out (Perry et al. 2014, pp. 131–134). Though human settlement and hunting activities played the most significant role, there are some factors that may have inhibited ''Dinornis novaezealandiae'' abilities to reproduce at the rate that they were being culled, such as the introduction of Polynesian dogs (Szabo, 2013). This is because it is believed that their extinction was at least in part due Polynesian dogs ''Canis familiaris'' eating moa chicks (Szabo, 2013).


Footnotes


References

* Allen, E. W., Erxleben, J., Hanhart, M., Hanhart, N., Keulemans, J. G., Mintern B., Rowley, G. D., Smith, J., & Walther, T.(1878). Ornithological miscellany. Retrieved from. https://commons.wikimedia.org/wiki/File:Ornithological_miscellany_(5982107548).jpg January 11, 2022. * * * * * * * * * * * * Scarlett, R. J. (1974). Moa and man in New Zealand. Notornis, 21(1), 1–12. * Szabo, M.J. (2013). (updated 2017). North Island giant moa. In Miskelly, C.M. (ed.) New Zealand Birds Online. Retrieved from https://nzbirdsonline.org.nz/species/north-island-giant-moa * * Wehi, P., & Cox, M. (2018). Dead as the Moa. Waikato.ac.nz. Retrieved from https://www.waikato.ac.nz/news-opinion/media/2018/dead-as-the-moa * *


External links


''North Island Giant Moa. Dinornis Novaezealandiae''.
by Paul Martinson. Artwork produced for the book Extinct Birds of New Zealand, by Alan Tennyson, Te Papa Press, Wellington, 2006
Holotypes
of Dinornis Novaezealandiae in the collection of the Museum of New Zealand Te Papa Tongarewa
''Ornithological miscellany''.
by Allen, Edgar W.; Erxleben, J.; Hanhart, Michael; Hanhart, N; Keulemans, J. G.; Mintern Bros.; Rowley, George Dawson; Smith, J.; Walther, T., Public domain, via Wikimedia Commons. {{Taxonbar, from=Q4691773
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 ...
Bird extinctions since 1500 Birds of the North Island Dinornithidae Extinct birds of New Zealand Extinct flightless birds Higher-level bird taxa restricted to New Zealand Holocene extinctions Late Quaternary prehistoric birds Ratites Species made extinct by human activities