Megalapteryx didinus

Life-forms

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Archosauromorpha
Cladus: Crurotarsi
Divisio: Archosauria
Cladus: Avemetatarsalia
Cladus: Ornithodira
Subtaxon: Dinosauromorpha
Cladus: Dinosauriformes
Cladus: Dracohors
Cladus: Dinosauria
Cladus: Saurischia
Cladus: Eusaurischia
Subordo: Theropoda
Cladus: Neotheropoda
Cladus: Averostra
Cladus: Tetanurae
Cladus: Avetheropoda
Cladus: Coelurosauria
Cladus: Tyrannoraptora
Cladus: Maniraptoromorpha
Cladus: Maniraptoriformes
Cladus: Maniraptora
Cladus: Pennaraptora
Cladus: Paraves
Cladus: Eumaniraptora
Cladus: Avialae
Infraclassis: Aves
Cladus: Avebrevicauda
Cladus: Pygostylia
Cladus: Ornithothoraces
Cladus: Ornithuromorpha
Cladus: Carinatae
Parvclassis: Neornithes
Cohors: Palaeognathae
Ordo: †Dinornithiformes

Familia: †Megalapterygidae
Genus: †Megalapteryx
Species: †Megalapteryx didinus
Name

Megalapteryx didinus (Owen, 1883)
Synonyms

Anomalopteryx didina Lydekker, 1891: 277
Dinornis didinus Owen, 1883: 257 (nomen nudum 1882) (protonym)
Megalapteryx benhami Archey 1941: 35 [syn. Worthy (1988: 107)]
Megalapteryx didinus Oliver, 1930
Megalapteryx hamiltoni Rothschild, 1907: 197
Megalapteryx hectori Haast, 1886: 162 (nomen nudum 1884, 1886)
Megalapteryx tenuipes Lydekker, 1891: 251
Palaeocasuarius elegans Forbes, 1892: 189
Palaeocasuarius haasti Forbes, 1892: 189
Palaeocasuarius velox Forbes, 1892: 189

References
Primary references

Owen, R. 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): 257–261, plates LIX–LXI. Communicated 20 June 1882. BHL DOI: 10.1111/j.1096-3642.1883.tb00360.x Reference page.
von Haast, J. 1886. On Megalapteryx hectori, a new gigantic species of apterygian bird. Transactions of the Zoological Society of London 12(5): 161–169, plate XXX. BHL DOI: 10.1111/j.1096-3642.1886.tb00010.x Reference page.
Lydekker, R. 1891. Catalogue of the fossil birds in the British Museum (Natural History). British Museum (Natural History): London. BHL DOI: 10.5962/bhl.title.8301 Reference page.
Forbes, H.O. 1892. Preliminary notice of additions to the extinct Avifauna of New Zealand. (Abstract.) Transactions and proceedings of the New Zealand Institute 24: 185–189. BHL Reference page.
Rothschild, L.W. 1907. Extinct birds. Hutchinson & Co.: London. BHL Reference page.
Archey, G. 1941. The moa: a study of the Dinornithiformes. Bulletin of the Auckland Institute and Museum 1. BHL Reference page.

Additional references

Worthy, T.H. 1988. A re-examination of the moa genus Megalapteryx. Notornis 35: 99–108. PDF ResearchGate Reference page.
Worthy, T.H. & Scofield, R.P. 2012. Twenty-first century advances in knowledge of the biology of moa (Aves: Dinornithiformes): a new morphological analysis and moa diagnoses revised. New Zealand Journal of Zoology 39(2): 87–153. DOI: 10.1080/03014223.2012.665060 Open access Reference page.
Gill, B.J., Bell, B.D., Chambers, G.K., Medway, D.G., Palma, R.L., Scofield, R.P., Tennyson, A.J.D., & Worthy, T.H. 2010. Checklist of the Birds of New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency, Antarctica. 4th ed. Te Tapa Press, Wellington, New Zealand: 500 pp. ISBN 978-1-877385-59-9. ResearchGate Reference page.

Vernacular names
English: Upland Moa
日本語: オオツバサモア

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

Megalapteryidae

Megalapteryx didinus
Dinornithidae
Dinornis

D. robustus

D. novaezealandiae
Emeidae
Pachyornis

P. australis

P. elephantopus

P. geranoides

Anomalopteryx didiformis

Emeus crassus

Euryapteryx curtus
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:

British Museum A16, found at Queenstown in 1876, is the type of the species.
Otago Museum C.68.2A, leg with much muscle tissue, skin and feathers from the Old Man Range
Museum of New Zealand Te Papa Tongarewa NMNZ S.000400, a skeleton with tissue on neck and head from the Cromwell area.[20]
Museum of New Zealand Te Papa Tongarewa NMNZ S.023080, a foot with some muscle and sinews, found on 7 January 1987 at Mount Owen. This was dated to be about 3,300–3,400 years old.[21]
Museum of New Zealand Te Papa Tongarewa NMNZ S.027950, feathers found in 1949 at Takahe Valley, Fiordland, New Zealand.[22]
Canterbury Museum NZ 1725, Remains of one partial egg which have been found at the Rakaia River in 1971 are tentatively attributed to this species. The radiocarbon date of approximately AD 1300–1400 is in line with this. Unusually, the eggshell is dark olive green, but even if the egg is of M. didinus, the shell colour may have varied between individual eggs.[23]
Museum of New Zealand Te Papa Tongarewa NMNZ S.023700, complete skeleton found by Trevor Worthy in March 1987 at Honeycomb Hill Cave, Oparara Valley[24]
Otago Museum AV10049, skeleton and partial egg found in 2002 at Serpentine Range, Humboldt Mountains.[25]

References

"Megalapteryx didinus. NZTCS". nztcs.org.nz. Retrieved 29 April 2023.
Brands, Sheila J. (1989). "The Taxonomicon". Zwaag, Netherlands: Universal Taxonomic Services. Retrieved 21 January 2010.
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.
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.
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.
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.
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.
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.
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.
Flannery, Tim, "A Gap in Nature: Discovering the World's Extinct Animals", October 2001, "[1]"
Museum of New Zealand, "Upland Moa", 1998, http://collections.tepapa.govt.nz/theme.aspx?irn=1348
TerraNature, "Flightless Birds: Moa", http://terranature.org/moa.htm
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.
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.
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.
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.
Gill, B. J. (2006). "A Catalogue of Moa Eggs (Aves: Dinornithiformes)". Records of the Auckland Museum. 43: 55–80. ISSN 1174-9202. JSTOR 42905885. Wikidata Q58623352.
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
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.
Museum of New Zealand(a). "Megalapteryx didinus". Collections Online. Museum of New Zealand Te Papa Tongarewa. Retrieved 18 July 2010.
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.
Museum of New Zealand(b). "Megalapteryx didinus". Collections Online. Museum of New Zealand Te Papa Tongarewa. Retrieved 18 July 2010.
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.
Museum of New Zealand(c). "Megalapteryx didinus". Collections Online. Museum of New Zealand Te Papa Tongarewa. Retrieved 18 July 2010.
"THE HUNT IS ON: Upland Moa Recovery Project".