Moa
Moa | |
---|---|
North Island giant moa skeleton | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Aves |
Infraclass: | Palaeognathae |
Clade: | Notopalaeognathae |
Order: | †Dinornithiformes Bonaparte, 1853[1] |
Type species | |
† Dinornis novaezealandiae
, 1843 | |
Subgroups | |
See text | |
Diversity[2] | |
6 genera, 9 species | |
Synonyms[3] | |
|
Moa
Moa are traditionally placed in the
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 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).[10]
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,
No records survive of what sounds moa made, though some idea of their
Evolutionary relationships
The moa's closest relatives are small terrestrial South American birds called the tinamous, which can fly.[9][14][15][16] Previously, the kiwi, the Australian emu, and cassowary[17] 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
Analyses of ancient DNA have determined that a number of cryptic evolutionary lineages occurred in several moa genera.[23] 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 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.[24] Some of the other size variation for moa species can probably be explained by similar geographic and temporal factors.[25]
The earliest moa remains come from the
Classification
Taxonomy
The currently recognised genera and species are:[5]
- Order †Dinornithiformes (Gadow 1893) Ridgway 1901 [Dinornithes Gadow 1893; Immanes Newton 1884] (moa)
- Family Dinornithidae Owen 1843 [Palapteryginae Bonaparte 1854; Palapterygidae Haast 1874; Dinornithnideae Stejneger 1884] (giant moa)
- Genus Dinornis
- North Island giant moa, Dinornis novaezealandiae (North Island, New Zealand)
- South Island giant moa, Dinornis robustus (South Island, New Zealand)
- Genus
- Family Emeidae (Bonaparte 1854) [Emeinae Bonaparte 1854; Anomalopterygidae Oliver 1930; Anomalapteryginae Archey 1941] (lesser moa)
- Genus Anomalopteryx
- Bush moa, Anomalopteryx didiformis (North and South Island, New Zealand)
- Genus Emeus
- Eastern moa, Emeus crassus (South Island, New Zealand)
- Genus Euryapteryx
- Broad-billed moa, Euryapteryx curtus (North and South Island, New Zealand)
- Genus Pachyornis
- Heavy-footed moa, Pachyornis elephantopus (South Island, New Zealand)
- Mantell's moa, Pachyornis geranoides (North Island, New Zealand)
- Crested moa, Pachyornis australis (South Island, New Zealand)[2]
- Genus
- Family Megalapterygidae
- Megalapteryx
- Upland moa, Megalapteryx didinus (South Island, New Zealand)
- Family Dinornithidae Owen 1843 [Palapteryginae Bonaparte 1854; Palapterygidae Haast 1874; Dinornithnideae Stejneger 1884] (giant moa)
Two unnamed species are also known from the Saint Bathans Fauna.[26]
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[27] 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.[28] The presence of Miocene-aged species certainly suggests that moa diversification began before the split between Megalapteryx and the other taxa.[26]
The
Bunce et al. also concluded that the highly complex structure of the moa lineage was caused by the formation of the Southern Alps about 6 Mya, and the habitat fragmentation on both islands resulting from Pleistocene glacial cycles, volcanism, and landscape changes.[19] The cladogram below is a phylogeny of Palaeognathae generated by Mitchell (2014)[15] with some clade names after Yuri et al. (2013).[30] It provides the position of the moa (Dinornithiformes) within the larger context of the "ancient jawed" (Palaeognathae) birds:
Palaeognathae |
| ||||||||||||||||||||||||||||||||||||||||||
The cladogram below gives a more detailed, species-level phylogeny, of the moa branch (Dinornithiformes) of the "ancient jawed" birds (Palaeognathae) shown above:[19]
†Dinornithiformes |
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||
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:[11][31][32][33][34][35][36]
South Island
The two main faunas identified in the South Island include:
- The fauna of the high-rainfall west coast beech (Nothofagus) 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 that included Pachyornis elephantopus (heavy-footed moa), Euryapteryx gravis, Emeus crassus, and Dinornis robustus.
A '
- Pachyornis australis, the rarest moa species, the only moa species not yet found in Māori middens. Its bones have been found in caves in the northwest Nelson and Karamea districts (such as Honeycomb Hill Cave), and some sites around the 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 on the west coast and Central Otago) existed. Their distributions in coastal areas have been rather unclear, but were present at least in several locations such as on Kaikōura, Otago Peninsula,[37] and Karitane.[38]
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.[11] 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 had made a land bridge across the Cook Strait.[11]
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.[11]
Behaviour and ecology
About eight moa
Diet
Their diet has been deduced from
Moa filled the
Like many other birds, moa swallowed gizzard stones (
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.[18]
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);[18][44] 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
No evidence has been found to suggest that moa were colonial nesters. Moa nesting is often inferred from accumulations of eggshell fragments in
Fragments of moa eggshell are often found in
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."[48] Despite the bird's extinction, the high yield of DNA available from recovered fossilised eggs has allowed the moa's genome to be sequenced.[49]
-
The skeleton of female upland moa with egg in unlaid position within the pelvic cavity in Otago Museum
-
An egg and embryo fragments of Emeus crassus
-
Restoration of an upland moa
Pre-human forests
Studies of accumulated dried vegetation in the pre-human mid-late Holocene period suggests a low Sophora microphylla or Kōwai forest ecosystem in Central Otago 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.[50]
Relationship with humans
Extinction
Before the arrival of humans, the moa's only predator was the massive
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 36 cm (14 in) between Tākaka and Riwaka that were found by a surveying party; and finally in 1878, the Otago Witness published an additional account from a farmer and his shepherd.[54] An 80-year-old woman, Alice McKenzie, claimed in 1959 that she had seen a moa in Fiordland 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.[55] In childhood, Mackenzie saw a large bird that she believed to be a takahē, but after its rediscovery in the 1940s, she saw a picture of it and concluded that she had seen something else.[56]
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.
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.[62]
Surviving remains
Dieffenbach[67] 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.[68][69] In 1839, John W. Harris, a Poverty Bay 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 15 cm (6 in) fragment of bone to his uncle, John Rule, a Sydney surgeon, who sent it to Richard Owen, who at that time was working at the Hunterian Museum at the Royal College of Surgeons in London.[54]
Owen puzzled over the fragment for almost four years. He established it was part of the femur 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, 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.[54]
In July 2004, the Natural History Museum 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
.Bones are commonly found in caves or tomo (the Māori word for doline or sinkhole, 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/marble areas of northwest Nelson, Karamea, Waitomo, and Te Anau.
Moa bones and eggshell fragments sometimes occur in active coastal sand dunes, where they may erode from
Densely intermingled moa bones have been encountered in swamps throughout New Zealand. The most well-known example is at Pyramid Valley in north Canterbury,[70] where bones from at least 183 individual moa have been excavated, mostly by Roger Duff of Canterbury Museum.[71] 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.[72]
Many New Zealand and international museums hold moa bone collections.
Feathers and soft tissues
Several examples of moa remains have been found with soft tissues (
- Dried muscle on bones of a female Dinornis robustus found at Tiger Hill in the Manuherikia River Valley by gold miners in 1864[74] (currently held by Yorkshire Museum)
- 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 1870Otago Museum)
- An articulated foot of a male D. giganteus with skin and foot pads preserved, found in a crevice on the Knobby Range in 1874Otago Museum)
- The type specimen of Megalapteryx didinus found near Queenstown in 1878[74] (currently held by Natural History Museum, London; see photograph of foot on this page)
- The lower leg of Pachyornis elephantopus, with skin and muscle, from the Hector Range in 1884;Cambridge University)
- The complete feathered leg of a M. didinus from Otago Museum)
- The head of a M. didinus found near Cromwell sometime before 1949Museum of New Zealand).
Two specimens are known from outside the Central Otago region:
- A complete foot of M. didinus found in a cave on Museum of New Zealand)
- A skeleton of Anomalopteryx didiformis with muscle, skin, and feather bases collected from a cave near Te Anau in 1980.[80]
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
Moa feathers are up to 23 cm (9 in) long, and a range of colours has been reported, including reddish-brown, white, yellowish, and purplish.[11] 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.[81]
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 as leading candidates for de-extinction.[82] Preliminary work involving the extraction of DNA has been undertaken by Japanese geneticist Ankoh Yasuyuki Shirota.[83][84]
Interest in the moa's potential for revival was further stirred in mid-2014 when New Zealand Member of Parliament Trevor Mallard suggested that bringing back some smaller species of moa within 50 years was a viable idea.[85] The idea was ridiculed by many, but gained support from some natural history experts.[86]
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.
Allen Curnow's poem, "The Skeleton of the Great Moa in the Canterbury Museum, Christchurch" was published in 1943.[87][88]
See also
- List of New Zealand species extinct in the Holocene
- Moa-nalo, several flightless ducks from the Hawaiian Islands that grew to be as large as geese.
- Elephant birds, flightless ratites up to over 3 metres tall that once lived on the island of Madagascar.
General:
- Late Quaternary prehistoric birds
- Island gigantism
- Megafauna
Footnotes
- ^ Brands, S. (2008)
- ^ a b Stephenson, Brent (2009)
- ^ Brodkob, Pierce (1963). "Catalogue of fossil birds 1. Archaeopterygiformes through Ardeiformes". Biological Sciences, Bulletin of the Florida State Museum. 7 (4): 180–293. Retrieved 30 December 2015.
- ^ a b OSNZ (2009)
- ^ a b Davies, S.J.J.F. (2003)
- ^ "Little bush moa | New Zealand Birds Online". nzbirdsonline.org.nz. Retrieved 24 July 2020.
- ^ . Retrieved 22 December 2014.
- ISSN 0906-7590.
- ^ a b Phillips, et al. (2010)
- ^ "Story: Moa". govt.nz. Retrieved 15 January 2022.
- ^ a b c d e f g h i j k Worthy & Holdaway (2002)
- ^ Schoon, Theo. "Cave drawing of a moa". Te Ara Encyclopedia of New Zealand. Te Ara.
- ^ "Te Manunui Rock Art Site". Heritage New Zealand.
- PMID 21596537.
- ^ S2CID 206555952. Archived from the original(PDF) on 30 May 2019.
- PMID 24825849.
- ^ a b c Turvey et al. (2005)
- ^ a b c Huynen, L.J., et al. (2003)
- ^ a b c d Bunce, M., et al. (2003)
- PMID 19923428.
- doi:10.3853/j.0067-1975.62.2010.1535. Archived from the original(PDF) on 11 April 2019.
- S2CID 83768608.
- PMID 15928096.
- ^ Worthy (1987)
- ^ Worthy, et al. (1988)
- ^ doi:10.3853/j.0067-1975.62.2010.1546. Archived from the original(PDF) on 11 April 2019.
- PMID 19923428.
- PMID 21596537.
- PMID 21596537.
- PMID 24832669.
- ^ Worthy, Trevor (1998)a
- ^ Worthy, Trevor (1998)b
- ^ Worthy, Trevor & Holdaway, Richard (1993)
- ^ Worthy, Trevor & Holdaway, Richard (1994)
- ^ Worthy, Trevor & Holdaway, Richard (1995)
- ^ Worthy, Trevor & Holdaway, Richard (1996)
- W & T Avery Ltd.Retrieved 3 February 2015.
- ^ Teviotdale D. (1932). "The material culture of the Moa-hunters in Murihiku – 2. Evidence of Zoology". The Journal of the Polynesian Society. 41 (162): 81–120. Retrieved 3 February 2015.
- ^ Burrows, et al. (1981)
- ^ a b c Wood (2007)
- ^ Horrocks, et al. (2004)
- ^ OCLC 83611783.
- S2CID 33405428.
- S2CID 1515413. Archived from the original(PDF) on 28 January 2019.
- ^ Hartree (1999)
- ^ a b Wood, J.R. (2008)
- ^ Gill, B.J. (2007)
- ^ a b Huynen, Leon; Gill, Brian J.; Millar, Craig D.; and Lambert, David M. (2010)
- ^ Yong, Ed. (2010)
- S2CID 245807815.
- ISBN 978-1107402171.
- .
- ^ Holdaway & Jacomb (2000)
- ^ a b c d Fuller, Errol (1987)
- ^ "Alice McKenzie and the Moa". Radio New Zealand.
- ^ "Alice Mackenzie describes seeing a moa and talks about her book, Pioneers of Martins Bay".
- ^ a b c Anderson (1989)
- ^ Anderson, Atholl (1990). Prodigious Birds: Moas and Moa-Hunting in New Zealand. Cambridge University Press.
- ^ Purcell, Rosamond (1999)
- ^ Gould, C. (1886)
- ^ Heuvelmans, B (1959)
- ^ a b c Laing, Doug (2008)
- ^ Nickell, Joe (26 May 2017). "The New Zealand Moa: From Extinct Bird to Cryptid". Skeptical Briefs. 27 (1). Center for Inquiry: 8–9.
- ^ Nickell, Joe (26 May 2017). "The New Zealand Moa: From Extinct Bird to Cryptid". Skeptical Inquirer. Retrieved 12 May 2019.
- ^ Polack, J.S. (1838)
- ^ Hill, H. (1913)
- ^ Dieffenbach, E. (1843)
- ^ Taonga, New Zealand Ministry for Culture and Heritage Te Manatu. "4. – Moa – Te Ara: The Encyclopedia of New Zealand". teara.govt.nz. Retrieved 4 May 2019.
- OCLC 819110163.
- ^ Holdaway, Richard & Worthy, Trevor (1997)
- ^ Davidson, Janet. "Roger Shepherd Duff". Dictionary of New Zealand Biography. Ministry for Culture and Heritage.
- ^ Wood, J.R., et al. (2008)
- ^ "Digitising moa". Retrieved 2 February 2022.
- ^ a b Owen, R. (1879)
- ^ Hutton, F.W. & Coughtrey, M. (1875)
- ^ a b Buller, W.L. (1888)
- ^ Hamilton, A. (1894)
- ^ Vickers-Rich, et al. (1995)
- ^ Worthy, Trevor (1989)
- ^ Forrest, R.M. (1987)
- PMID 19570784.
- ^ Le Roux, M., "Scientists plan to resurrect a range of extinct animals using DNA and cloning", Courier Mail, 23 April 2013. Retrieved 25 July 2014.
- ^ Young, E (1997). "Moa genes could rise from the dead". New Scientist. 153 (2063).
- ^ "Life in the Old Moa Yet", New Zealand Science Monthly, February 1997. Retrieved 25 July 2014.
- ^ O'Brien, T. Mallard: Bring the moa back to life within 50 years", 3news, 1 July 2014. Retrieved 25 July 2014.
- ^ Tohill, M.-J., "Expert supports Moa revival idea", stuff.co.nz, 9 July 2014. Retrieved 25 July 2014.
- ^ "A poem a day: The Skeleton of the Great Moa in the Canterbury Museum, Christchurch – Allen Curnow". 25 April 2011. Archived from the original on 1 February 2020.
- ^ Curnow, Allen (1944). Sailing or Drowning. Wellington: Progressive Publishing Society.
Notes
- ^ The word moa is from the Māori language and is both singular and plural. Usage in New Zealand English and in the scientific literature in recent years has been changing to reflect this.
- Saint Bathans Fauna.
References
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- Hartree, W.H. (1999). "A preliminary report on the nesting habits of moas in the East Coast of the North Island" (PDF). Notornis. 46 (4): 457–460.
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- Horrocks, M.; et al. (2004). "Plant remains in coprolites: diet of a subalpine moa (Dinornithiformes) from southern New Zealand". Emu. 104 (2): 149–156. S2CID 86345660.
- Hutton, F.W.; Coughtrey, M. (1874). "Notice of the Earnscleugh Cave". Transactions and Proceedings of the New Zealand Institute. 7: 138–144.
- Huynen, Leon; Gill, Brian J.; Millar, Craig D.; Lambert, David M. (30 August 2010). "Ancient DNA Reveals Extreme Egg Morphology and Nesting Behavior in New Zealand's Extinct Moa". Proceedings of the National Academy of Sciences. 107 (30): 16201–16206. PMID 20805485.
- Huynen, Leon J.; Millar, Craig D.; Scofield, R.P.; Lambert, David M. (2003). "Nuclear DNA sequences detect species limits in ancient moa". Nature. 425 (6954): 175–178. S2CID 4413995.
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- Purcell, Rosamond (1999). Swift as a Shadow. Mariner Books. p. 32. ISBN 978-0-395-89228-2.
- Stephenson, Brent (5 January 2009). "New Zealand Recognised Bird Names (NZRBN) database". New Zealand: Ornithological Society of New Zealand. Archived from the original on 25 April 2015. Retrieved 10 May 2010.
- Turvey, Samuel T.; Green, Owen R.; S2CID 4308841.
- Vickers-Rich, P; Trusler, P; Rowley, MJ; Cooper, A; Chambers, GK; Bock, WJ; Millener, PR; Worthy, Trevor; Yaldwyn, JC (1995). "Morphology, myology, collagen and DNA of a mummified moa, Megalapteryx didinus (Aves: Dinornithiformes) from New Zealand" (PDF). Tuhinga: Records of the Museum of New Zealand te Papa Tongarewa. 4: 1–26. Archived from the original (PDF) on 22 May 2010.
- Wood, J.R. (2007). "Moa gizzard content analyses: further information on the diet of Dinornis robustus and Emeus crassus, and the first evidence for the diet of Pachyornis elephantopus (Aves: Dinornithiformes)". Records of the Canterbury Museum. 21: 27–39.
- Wood, J.R. (2008). "Moa (Aves: Dinornithiformes) nesting material from rockshelters in the semi-arid interior of South Island, New Zealand". Journal of the Royal Society of New Zealand. 38 (3): 115–129. S2CID 129645654.
- Wood, J.R.; hdl:2440/62495.
- Worthy, Trevor (1989). "Mummified moa remains from Mt. Owen, northwest Nelson" (PDF). Notornis. 36: 36–38.
- ISBN 978-0-253-34034-4.
- Worthy, Trevor (March 2009). "A moa sighting?". Te Ara – the Encyclopedia of New Zealand. Retrieved 14 February 2011.
- Yong, Ed (10 March 2010). "DNA from the Largest Bird Ever Sequenced from Fossil Eggshells". Discover Magazine. Archived from the original on 22 September 2020. Retrieved 14 February 2011.
External links
- TerraNature list of New Zealand's extinct birds
- TerraNature page on Moa
- Tree of Life classification and references
- Moa article in Te Ara – the Encyclopedia of New Zealand
- 3D model of a moa skull