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==Taxonomy==
==Taxonomy==
The upland moa was named as ''Dinornis didinus'' in 1883 by [[Richard Owen]] from mummified material found in 1878 by H. L. Squires in [[Queenstown, New Zealand]] and sent to the British Museum. The [[holotype]] specimen consists of a mummified head and partial neck, and two mummified legs and feet which preserve the feathers.<ref>{{cite journal |last1=Owen |first1=Richard |title=On Dinornis (Part XXIV): containing a Description of the Head and Feet, with their dried Integuments, of an Individual of the species Dinornis didinus, Owen |journal=Transactions of the Zoological Society of London |date=1883 |volume=11 |issue=8 |pages=257-261 |url=https://archive.org/details/transactionsofzo11zool/page/257/mode/1up |access-date=7 July 2023 |publisher=London : Zoological Society of London}}</ref>
The upland moa was named as ''Dinornis didinus'' in 1883 by [[Richard Owen]] from mummified material found in 1878 by H. L. Squires in [[Queenstown, New Zealand]] and sent to the British Museum. The [[holotype]] specimen consists of a mummified head and partial neck, and two mummified legs and feet which preserve the feathers.<ref>{{cite journal |last1=Owen |first1=Richard |title=On Dinornis (Part XXIV): containing a Description of the Head and Feet, with their dried Integuments, of an Individual of the species Dinornis didinus, Owen |journal=Transactions of the Zoological Society of London |date=1883 |volume=11 |issue=8 |pages=257–261 |url=https://archive.org/details/transactionsofzo11zool/page/257/mode/1up |access-date=7 July 2023 |publisher=London : Zoological Society of London}}</ref>


In 2005, a genetic study suggested that ''M. benhami'', which had previously been considered a [[junior synonym]] of ''M. didinus'', may have been a valid species after all.<ref>{{Cite journal | doi = 10.1073/pnas.0409435102| title = Reconstructing the tempo and mode of evolution in an extinct clade of birds with ancient DNA: The giant moas of New Zealand| journal = Proceedings of the National Academy of Sciences| volume = 102| issue = 23| pages = 8257–62| year = 2005| last1 = Baker | first1 = A. J.| last2 = Huynen | first2 = L. J.| last3 = Haddrath | first3 = O.| last4 = Millar | first4 = C. D.| last5 = Lambert | first5 = D. M. | pmid=15928096 | pmc=1149408| bibcode = 2005PNAS..102.8257B| doi-access = free}}</ref><ref>{{cite encyclopedia |last=Davies |first=S. J. J. F.|editor=Hutchins, Michael |encyclopedia=Grzimek's Animal Life Encyclopedia |title=Moas |edition=2 |year=2003 |publisher=Gale Group |volume=8 Birds I Tinamous and Ratites to Hoatzins |location=Farmington Hills, MI|isbn=978-0-7876-5784-0 |pages=95–98}}</ref>
In 2005, a genetic study suggested that ''M. benhami'', which had previously been considered a [[junior synonym]] of ''M. didinus'', may have been a valid species after all.<ref>{{Cite journal | doi = 10.1073/pnas.0409435102| title = Reconstructing the tempo and mode of evolution in an extinct clade of birds with ancient DNA: The giant moas of New Zealand| journal = Proceedings of the National Academy of Sciences| volume = 102| issue = 23| pages = 8257–62| year = 2005| last1 = Baker | first1 = A. J.| last2 = Huynen | first2 = L. J.| last3 = Haddrath | first3 = O.| last4 = Millar | first4 = C. D.| last5 = Lambert | first5 = D. M. | pmid=15928096 | pmc=1149408| bibcode = 2005PNAS..102.8257B| doi-access = free}}</ref><ref>{{cite encyclopedia |last=Davies |first=S. J. J. F.|editor=Hutchins, Michael |encyclopedia=Grzimek's Animal Life Encyclopedia |title=Moas |edition=2 |year=2003 |publisher=Gale Group |volume=8 Birds I Tinamous and Ratites to Hoatzins |location=Farmington Hills, MI|isbn=978-0-7876-5784-0 |pages=95–98}}</ref>
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==Behavior and ecology==
==Behavior and ecology==
[[File:Moaupland.jpg|upright|thumb|Mummified head head of the holotype, as illustrated in Owen's 1883 article]]
[[File:Moaupland.jpg|upright|thumb|Mummified head head of the holotype, as illustrated in Owen's 1883 article]]
The upland moa was herbivorous, its diet extrapolated from fossilised stomach contents, droppings, and the structure of its beak and [[crop (anatomy)|crop]]. It ate leaves and small twigs, using its beak to "shear&nbsp;... with scissor-like moves".<ref name="Te Papa" /> However, bio-mechanical testing of its beak and head has shown it was best suited to feed by pulling backwards, possibly while twisting its head to the side.<ref>{{cite journal |last1=Attard |first1=Marie R. G. |last2=Wilson |first2=Laura A. B. |last3=Worthy |first3=Trevor H. |last4=Scofield |first4=Paul |last5=Johnston |first5=Peter |last6=Parr |first6=William C. H. |last7=Wroe |first7=Stephen |title=Moa diet fits the bill: virtual reconstruction incorporating mummified remains and prediction of biomechanical performance in avian giants |journal=Proceedings of the Royal Society B: Biological Sciences |date=2016 |volume=283 |issue=1822 |pages=20152043 |doi=10.1098/rspb.2015.2043 |url=https://www.researchgate.net/publication/290395211_Moa_diet_fits_the_bill_Virtual_reconstruction_incorporating_mummified_remains_and_prediction_of_biomechanical_performance_in_avian_giants |access-date=21 July 2023|pmc=4721086 }}</ref> Its food required grinding before it could be digested, as indicated by its large crop.<ref name= "Te Papa"/> Studies of the upland moa's [[coprolite]]s provided evidence that branchlets of trees such as ''[[Nothofagus]]'', various lake-edge herbs, [[tussock (grass)|tussock]], and the nectar-rich flowers made up part of its diet.<ref>{{cite journal |last1=Horrocks |first1=Mark |last2=D'Costa |first2=Donna |last3=Wallace |first3=Rod |last4=Gardner |first4=Rhys |last5=Kondo |first5=Renzo |title=Plant remains in coprolites: diet of a subalpine moa (Dinornithiformes) from southern New Zealand |journal=Emu - Austral Ornithology |date=2004 |volume=104 |issue=2 |pages=149–156 |doi=10.1071/MU03019}}</ref><ref name="Wood et al 2012">{{cite journal |last1=Wood |first1=Jamie R. |last2=Wilmshurst |first2=Janet M. |last3=Wagstaff |first3=Steven J. |last4=Worthy |first4=Trevor H. |last5=Rawlence |first5=Nicolas J. |last6=Cooper |first6=Alan |title=High-Resolution Coproecology: Using Coprolites to Reconstruct the Habits and Habitats of New Zealand’s Extinct Upland Moa (Megalapteryx didinus) |journal=PLoS ONE |date=2012 |volume=7 |issue=6 |pages=e40025 |doi=10.1371/journal.pone.0040025 |url=https://www.researchgate.net/publication/228331799_High-Resolution_Coproecology_Using_Coprolites_to_Reconstruct_the_Habits_and_Habitats_of_New_Zealand%27s_Extinct_Upland_Moa_Megalapteryx_didinus |access-date=21 July 2023|pmc=3386916 }}</ref> Based on the pollen present in the coprolites, it is suggested they had a similar lifestyle to the living [[takahē]], feeding in high alpine areas during the summer and descending to lower altitude forests in the winter. They may also have played an important role in the seed dispersal of alpine plants.<ref name="Wood et al 2012"/>
The upland moa was herbivorous, its diet extrapolated from fossilised stomach contents, droppings, and the structure of its beak and [[crop (anatomy)|crop]]. It ate leaves and small twigs, using its beak to "shear&nbsp;... with scissor-like moves".<ref name="Te Papa" /> However, bio-mechanical testing of its beak and head has shown it was best suited to feed by pulling backwards, possibly while twisting its head to the side.<ref>{{cite journal |last1=Attard |first1=Marie R. G. |last2=Wilson |first2=Laura A. B. |last3=Worthy |first3=Trevor H. |last4=Scofield |first4=Paul |last5=Johnston |first5=Peter |last6=Parr |first6=William C. H. |last7=Wroe |first7=Stephen |title=Moa diet fits the bill: virtual reconstruction incorporating mummified remains and prediction of biomechanical performance in avian giants |journal=Proceedings of the Royal Society B: Biological Sciences |date=2016 |volume=283 |issue=1822 |pages=20152043 |doi=10.1098/rspb.2015.2043 |pmid=26763698 |url=https://www.researchgate.net/publication/290395211 |access-date=21 July 2023|pmc=4721086 }}</ref> Its food required grinding before it could be digested, as indicated by its large crop.<ref name= "Te Papa"/> Studies of the upland moa's [[coprolite]]s provided evidence that branchlets of trees such as ''[[Nothofagus]]'', various lake-edge herbs, [[tussock (grass)|tussock]], and the nectar-rich flowers made up part of its diet.<ref>{{cite journal |last1=Horrocks |first1=Mark |last2=D'Costa |first2=Donna |last3=Wallace |first3=Rod |last4=Gardner |first4=Rhys |last5=Kondo |first5=Renzo |title=Plant remains in coprolites: diet of a subalpine moa (Dinornithiformes) from southern New Zealand |journal=Emu - Austral Ornithology |date=2004 |volume=104 |issue=2 |pages=149–156 |doi=10.1071/MU03019|s2cid=86345660 }}</ref><ref name="Wood et al 2012">{{cite journal |last1=Wood |first1=Jamie R. |last2=Wilmshurst |first2=Janet M. |last3=Wagstaff |first3=Steven J. |last4=Worthy |first4=Trevor H. |last5=Rawlence |first5=Nicolas J. |last6=Cooper |first6=Alan |title=High-Resolution Coproecology: Using Coprolites to Reconstruct the Habits and Habitats of New Zealand's Extinct Upland Moa (Megalapteryx didinus) |journal=PLOS ONE |date=2012 |volume=7 |issue=6 |pages=e40025 |doi=10.1371/journal.pone.0040025 |pmid=22768206 |url=https://www.researchgate.net/publication/228331799 |access-date=21 July 2023|pmc=3386916 |bibcode=2012PLoSO...740025W |doi-access=free }}</ref> Based on the pollen present in the coprolites, it is suggested they had a similar lifestyle to the living [[takahē]], feeding in high alpine areas during the summer and descending to lower altitude forests in the winter. They may also have played an important role in the seed dispersal of alpine plants.<ref name="Wood et al 2012"/>


This moa usually laid only 1 to 2 blue-green coloured eggs at once,<ref name="Te Papa"/><ref>{{cite journal |last1=Igic |first1=Branislav |last2=Greenwood |first2=David R. |last3=Palmer |first3=David J. |last4=Cassey |first4=Phillip |last5=Gill |first5=Brian J. |last6=Grim |first6=Tomas |last7=Brennan |first7=Patricia L. R. |last8=Bassett |first8=Suzanne M. |last9=Battley |first9=Phil F. |last10=Hauber |first10=Mark E. |title=Detecting pigments from colourful eggshells of extinct birds |journal=Chemoecology |date=2010 |volume=20 |issue=1 |pages=43–48 |doi=10.1007/s00049-009-0038-2 |url=https://www.researchgate.net/publication/226960070_Detecting_pigments_from_colourful_eggshells_of_extinct_birds}}</ref> and was likely the only type of moa to lay eggs that were not white in colour.<ref>{{Cite journal| issn = 1174-9202| volume = 43| pages = 55–80| last = Gill| first = B. J.| title = A Catalogue of Moa Eggs (Aves: Dinornithiformes)| journal = [[Auckland War Memorial Museum|Records of the Auckland Museum]]| date = 2006 |url=https://www.researchgate.net/publication/273695552_A_catalogue_of_moa_eggs_Aves_Dinornithiformes |access-date=21 July 2023}}</ref> Like the emu and ostrich, male moa cared for the young.<ref name="Flannery"/> The upland moa's only predator before the arrival of humans in New Zealand was the [[Haast's eagle]].<ref name= "Te Papa"/>
This moa usually laid only 1 to 2 blue-green coloured eggs at once,<ref name="Te Papa"/><ref>{{cite journal |last1=Igic |first1=Branislav |last2=Greenwood |first2=David R. |last3=Palmer |first3=David J. |last4=Cassey |first4=Phillip |last5=Gill |first5=Brian J. |last6=Grim |first6=Tomas |last7=Brennan |first7=Patricia L. R. |last8=Bassett |first8=Suzanne M. |last9=Battley |first9=Phil F. |last10=Hauber |first10=Mark E. |title=Detecting pigments from colourful eggshells of extinct birds |journal=Chemoecology |date=2010 |volume=20 |issue=1 |pages=43–48 |doi=10.1007/s00049-009-0038-2 |s2cid=10956718 |url=https://www.researchgate.net/publication/226960070}}</ref> and was likely the only type of moa to lay eggs that were not white in colour.<ref>{{Cite journal| issn = 1174-9202| volume = 43| pages = 55–80| last = Gill| first = B. J.| title = A Catalogue of Moa Eggs (Aves: Dinornithiformes)| journal = [[Auckland War Memorial Museum|Records of the Auckland Museum]]| date = 2006 |url=https://www.researchgate.net/publication/273695552 |access-date=21 July 2023}}</ref> Like the emu and ostrich, male moa cared for the young.<ref name="Flannery"/> The upland moa's only predator before the arrival of humans in New Zealand was the [[Haast's eagle]].<ref name= "Te Papa"/>


==Extinction==
==Extinction==

Revision as of 21:20, 31 October 2023

Upland moa
Temporal range: Pleistocene-Holocene
Mounted skeleton from Honeycomb Hill Cave, collection of Te Papa

Extinct (c. 1500) (NZ TCS)[1]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Infraclass: Palaeognathae
Clade: Notopalaeognathae
Order: Dinornithiformes
Family: Megalapterygidae
Bunce et al., 2009
Genus: Megalapteryx
Haast 1886[2]
Species:
M. didinus
Binomial name
Megalapteryx didinus
(Owen, 1883)[2][3]
Synonyms
list
  • Palaeocasuarius Forbes 1892 ex Rothschild 1907
  • Dinornis didinus Owen 1882 ex Owen 1883
  • Anomalopteryx didina (Owen 1883) Lydekker 1891
  • Megalapteryx hectori Haast 1884 ex Haast 1886
  • Megalapteryx tenuipes Lydekker, 1891
  • Megalapteryx hamiltoni Rothschild 1907
  • Palaeocasuarius velox Forbes 1892 ex Rothschild 1907
  • Palaeocasuarius elegans Rothschild 1907
  • Palaeocasuarius haasti Forbes 1892 ex Rothschild 1907
  • Megalapteryx benhami Archey 1941

The upland moa (Megalapteryx didinus) is an extinct species of moa that was endemic to New Zealand. It is a ratite, a grouping of flightless birds with no keel on the sternum. It was the last moa species to become extinct, vanishing around 1500 CE, and was predominantly found in alpine and sub-alpine environments. [4]

Taxonomy

The upland moa was named as Dinornis didinus in 1883 by Richard Owen from mummified material found in 1878 by H. L. Squires in Queenstown, New Zealand and sent to the British Museum. The holotype specimen consists of a mummified head and partial neck, and two mummified legs and feet which preserve the feathers.[5]

In 2005, a genetic study suggested that M. benhami, which had previously been considered a junior synonym of M. didinus, may have been a valid species after all.[6][7]

The cladogram below follows a 2009 analysis by Bunce et al.:[8]

Dinornithiformes

Description

Restoration from 1907

At less than 1 metre (3.3 ft) tall and about 17 to 34 kilograms (37 to 75 lb) in weight, the upland moa was among the smallest of the moa species.[9] Unlike other moas, it had feathers covering all of its body but the beak and the soles of its feet, an adaptation to its cold environment.[10] Scientists believed in the past that the upland moa held its neck and head upright; however, it actually carried itself in a stooped posture with its head level to its back. This would have helped it travel through the abundant vegetation in its habitat, whereas an extended neck would have been more suited to open spaces.[11] It had no wings or tail.[12]

Distribution and habitat

The upland moa lived only on New Zealand's South Island, in mountains and sub-alpine regions. They travelled to elevations as high as 2000 m (7000 ft).[11]

Behavior and ecology

Mummified head head of the holotype, as illustrated in Owen's 1883 article

The upland moa was herbivorous, its diet extrapolated from fossilised stomach contents, droppings, and the structure of its beak and crop. It ate leaves and small twigs, using its beak to "shear ... with scissor-like moves".[11] However, bio-mechanical testing of its beak and head has shown it was best suited to feed by pulling backwards, possibly while twisting its head to the side.[13] Its food required grinding before it could be digested, as indicated by its large crop.[11] Studies of the upland moa's coprolites provided evidence that branchlets of trees such as Nothofagus, various lake-edge herbs, tussock, and the nectar-rich flowers made up part of its diet.[14][15] Based on the pollen present in the coprolites, it is suggested they had a similar lifestyle to the living takahē, feeding in high alpine areas during the summer and descending to lower altitude forests in the winter. They may also have played an important role in the seed dispersal of alpine plants.[15]

This moa usually laid only 1 to 2 blue-green coloured eggs at once,[11][16] and was likely the only type of moa to lay eggs that were not white in colour.[17] Like the emu and ostrich, male moa cared for the young.[10] The upland moa's only predator before the arrival of humans in New Zealand was the Haast's eagle.[11]

Extinction

Humans first came in contact with the upland moa around 1250 to 1300 AD, when the Māori people arrived in New Zealand from Polynesia. Moa, a docile animal, were an easy source of food for the Māori (who called them "moa pukepuke")[1] and were eventually hunted to extinction in 1445.[11][18]

Discoveries

Preserved foot of the holotype specimen

The species has the best-preserved mummified remains of any moa species. [19] Several specimens with soft tissue and feather remains are known:

References

  1. ^ a b "Megalapteryx didinus. NZTCS". nztcs.org.nz. Retrieved 29 April 2023.
  2. ^ a b Brands, Sheila J. (1989). "The Taxonomicon". Zwaag, Netherlands: Universal Taxonomic Services. Retrieved 21 January 2010.
  3. ^ Checklist Committee Ornithological Society of New Zealand (2010). "Checklist-of-Birds of New Zealand, Norfolk and Macquarie Islands and the Ross Dependency Antarctica" (PDF). Te Papa Press. Retrieved 4 January 2016.
  4. ^ Tennyson, Alan J. D. (2006). Extinct birds of New Zealand. Paul Martinson. Wellington, N.Z.: Te Papa Press. ISBN 978-0-909010-21-8. OCLC 80016906.
  5. ^ Owen, Richard (1883). "On Dinornis (Part XXIV): containing a Description of the Head and Feet, with their dried Integuments, of an Individual of the species Dinornis didinus, Owen". Transactions of the Zoological Society of London. 11 (8). London : Zoological Society of London: 257–261. Retrieved 7 July 2023.
  6. ^ Baker, A. J.; Huynen, L. J.; Haddrath, O.; Millar, C. D.; Lambert, D. M. (2005). "Reconstructing the tempo and mode of evolution in an extinct clade of birds with ancient DNA: The giant moas of New Zealand". Proceedings of the National Academy of Sciences. 102 (23): 8257–62. Bibcode:2005PNAS..102.8257B. doi:10.1073/pnas.0409435102. PMC 1149408. PMID 15928096.
  7. ^ Davies, S. J. J. F. (2003). "Moas". In Hutchins, Michael (ed.). Grzimek's Animal Life Encyclopedia. Vol. 8 Birds I Tinamous and Ratites to Hoatzins (2 ed.). Farmington Hills, MI: Gale Group. pp. 95–98. ISBN 978-0-7876-5784-0.
  8. ^ Bunce, M.; Worthy, T. H.; Phillips, M. J.; Holdaway, R. N.; Willerslev, E.; Haile, J.; Shapiro, B.; Scofield, R. P.; Drummond, A.; Kamp, P. J. J.; Cooper, A. (2009). "The evolutionary history of the extinct ratite moa and New Zealand Neogene paleogeography" (PDF). Proceedings of the National Academy of Sciences. 106 (49): 20646–20651. Bibcode:2009PNAS..10620646B. doi:10.1073/pnas.0906660106. PMC 2791642. PMID 19923428.
  9. ^ Vickers-Rich, P.; Trusler, P.; Rowley, M. J.; Cooper, A. (1 January 1995). "Morphology, myology, collagen and DNA of a mummified upland moa, Megalapteryx didinus (Aves: dinorthiformes) from New Zealand". Tuhinga: The Records of the Museum of New Zealand te Papa Tongarewa. 4: 1–26.
  10. ^ a b Flannery, Tim, "A Gap in Nature: Discovering the World's Extinct Animals", October 2001, "[1]"
  11. ^ a b c d e f g Museum of New Zealand, "Upland Moa", 1998, http://collections.tepapa.govt.nz/theme.aspx?irn=1348
  12. ^ TerraNature, "Flightless Birds: Moa", http://terranature.org/moa.htm
  13. ^ Attard, Marie R. G.; Wilson, Laura A. B.; Worthy, Trevor H.; Scofield, Paul; Johnston, Peter; Parr, William C. H.; Wroe, Stephen (2016). "Moa diet fits the bill: virtual reconstruction incorporating mummified remains and prediction of biomechanical performance in avian giants". Proceedings of the Royal Society B: Biological Sciences. 283 (1822): 20152043. doi:10.1098/rspb.2015.2043. PMC 4721086. PMID 26763698. Retrieved 21 July 2023.
  14. ^ Horrocks, Mark; D'Costa, Donna; Wallace, Rod; Gardner, Rhys; Kondo, Renzo (2004). "Plant remains in coprolites: diet of a subalpine moa (Dinornithiformes) from southern New Zealand". Emu - Austral Ornithology. 104 (2): 149–156. doi:10.1071/MU03019. S2CID 86345660.
  15. ^ a b Wood, Jamie R.; Wilmshurst, Janet M.; Wagstaff, Steven J.; Worthy, Trevor H.; Rawlence, Nicolas J.; Cooper, Alan (2012). "High-Resolution Coproecology: Using Coprolites to Reconstruct the Habits and Habitats of New Zealand's Extinct Upland Moa (Megalapteryx didinus)". PLOS ONE. 7 (6): e40025. Bibcode:2012PLoSO...740025W. doi:10.1371/journal.pone.0040025. PMC 3386916. PMID 22768206. Retrieved 21 July 2023.
  16. ^ Igic, Branislav; Greenwood, David R.; Palmer, David J.; Cassey, Phillip; Gill, Brian J.; Grim, Tomas; Brennan, Patricia L. R.; Bassett, Suzanne M.; Battley, Phil F.; Hauber, Mark E. (2010). "Detecting pigments from colourful eggshells of extinct birds". Chemoecology. 20 (1): 43–48. doi:10.1007/s00049-009-0038-2. S2CID 10956718.
  17. ^ Gill, B. J. (2006). "A Catalogue of Moa Eggs (Aves: Dinornithiformes)". Records of the Auckland Museum. 43: 55–80. ISSN 1174-9202. Retrieved 21 July 2023.
  18. ^ Worthy, Trevor H.'Moa – Moa and people', Te Ara – the Encyclopedia of New Zealand, updated 13-Jul-12 URL: http://www.TeAra.govt.nz/en/moa/page-4
  19. ^ Tennyson, Alan J. D. (2006). Extinct birds of New Zealand. Paul Martinson. Wellington, N.Z.: Te Papa Press. p. 20. ISBN 978-0-909010-21-8. OCLC 80016906.
  20. ^ Museum of New Zealand(a). "Megalapteryx didinus". Collections Online. Museum of New Zealand Te Papa Tongarewa. Retrieved 18 July 2010.
  21. ^ Worthy, Trevor H. (1989). "Mummified moa remains from Mt Owen, northwest Nelson" (PDF). Notornis. 36 (1): 36–38. Archived from the original (PDF) on 27 October 2007. Retrieved 19 August 2006.
  22. ^ Museum of New Zealand(b). "Megalapteryx didinus". Collections Online. Museum of New Zealand Te Papa Tongarewa. Retrieved 18 July 2010.
  23. ^ McCulloch, Beverley (1992). "Unique, dark olive-green moa eggshell from Redcliffe Hill, Rakaia Gorge, Canterbury" (PDF). Notornis. 39 (1): 63–65. Archived from the original (PDF) on 18 October 2008. Retrieved 20 August 2006.
  24. ^ Museum of New Zealand(c). "Megalapteryx didinus". Collections Online. Museum of New Zealand Te Papa Tongarewa. Retrieved 18 July 2010.
  25. ^ "THE HUNT IS ON: Upland Moa Recovery Project".