The effect of climate and environmental change on the megafaunal moa of New Zealand in the absence of humans

Quaternary Science Reviews

Volumn 50, Issue , 2012, Pages 141-153

  Rawlence, Nicolas J ^a,b   Metcalf, Jessica L ^a,c   Wood, Jamie R ^d   Worthy, Trevor H ^a   Austin, Jeremy J ^a,e   Cooper, Alan ^a  

^a UNIVERSITY OF ADELAIDE   (Australia)

^b UNIVERSITY OF WAIKATO   (New Zealand)

^c UNIVERSITY OF COLORADO   (United States)

^d LANDCARE RESEARCH   (New Zealand)

^e MUSEUM VICTORIA   (Australia)

Author keywords

Ancient DNA; Climate change; Dinornithiformes; Extinction; Habitat tracking; Isotopes; Megafauna; Moa; New Zealand; Population demographics

Indexed keywords

DINORNITHIFORMES; MEGAFAUNA; MOA; NEW ZEALAND; POPULATION DEMOGRAPHICS;

BIRDS; CLIMATE CHANGE; ECOSYSTEMS; ISOTOPES; LIGHT EXTINCTION; POPULATION STATISTICS;

POPULATION DISTRIBUTION;

BIRD; CLIMATE EFFECT; CLIMATE VARIATION; DEMOGRAPHIC TREND; ENVIRONMENTAL CHANGE; HABITAT MANAGEMENT; HUMAN SETTLEMENT; ISOTOPIC ANALYSIS; MITOCHONDRIAL DNA; PHYLOGEOGRAPHY; POPULATION SIZE; QUATERNARY; RADIOCARBON DATING; TRACKWAY;

NEW ZEALAND;

AVES; DINORNITHIFORMES; PACHYORNIS; PACHYORNIS AUSTRALIS; PACHYORNIS ELEPHANTOPUS; PALAEOGNATHAE;

EID: 84865424871     PISSN: 02773791     EISSN: None     Source Type: Journal    
DOI: 10.1016/j.quascirev.2012.07.004     Document Type: Article

References (92)

1. Allentoft M.E., Rawlence N.J. Moa's ark or volant ghosts of Gondwana? Insights from nineteen years of ancient DNA research on the extinct moa (Aves: Dinornithiformes) of New Zealand. Ann. Anat. 2012, 194:36-51.
2. Allentoft M.E., Bunce M., Scofield R.P., Hale M.L., Holdaway R.N. Highly skewed sex ratios and biased fossil deposition of moa: ancient DNA provides new insight on New Zealand's extinct megafauna. Quat. Sci. Rev. 2010, 29:753-762.
3. Allentoft M.E., Oskam C., Houston J., Hale M.L., Gilbert M.T.P., Rasmussen M., Spencer P., Jacomb C., Willerslev E., Holdaway R.N., Bunce M. Profiling the dead: generating microsatellite data from fossil bones of extinct megafauna - protocols, problems, and prospects. PLoS One 2011, 6:e16670.
4. Allentoft M.E., Schuster S., Holdaway R.N., Hale M.L., McLay E., Oskam C., Gilbert M.T.P., Spencer P., Willerslev E., Bunce M. Identification of microsatellites from an extinct moa species using high-throughput (454) sequence data. Biotechniques 2009, 42:195-200.
5. Allentoft, M.E., Scofield, R.P., Oskam, C.L., Hale, M.L., Holdaway, R.N., Bunce, M. A molecular characterization of a newly discovered megafaunal fossil site in North Canterbury, South Island, New Zealand. J. Roy. Soc. New Zeal., . http://10.1080/03036758.2011.574821.
6. Alloway B.V., Lowe D.J., Barrell D.J.A., Newnham R.M., Almond P.C., Augustinus P.C., Bertler N.A.N., Carter L., Litchfield N.J., McGlone M.S., Shulmeister J., Vandergoes M.J., Williams P.W., INTIMATE members Towards a climate event stratigraphy for New Zealand over the past 30,000 years (NZ-INTIMATE project). J. Quat. Sci. 2007, 22:9-35.
7. Amundson R., Austin A.T., Schuur E.A.G., Yoo K., Matzek V., Kendall C., Uebersax A., Brenner D., Baisden W.T. Global patterns of the isotopic composition of soil and plant nitrogen. Global Biogeochem. Cy. 2003, 17:1031-1040.
8. Anderson C.N.K., Ramakrishnan U., Chan Y.L., Hadly E.A. Serial SimCoal: a population genetics model for data from multiple populations and points in time. Bioinformatics 2006, 21:1733-1734.
9. Anderson A. Prodigious Birds 1989, Cambridge University Press, Cambridge.
10. Baker A.J., Huynen L.J., Haddrath O., Miller C.D., Lambert D.M. Reconstructing the tempo and mode of evolution in an extinct clade of birds with ancient DNA: the giant moas of New Zealand. P. Natl. Acad. Sci. U. S. A. 2005, 102:8257-8262.
11. Barnosky A.D., Koch P.L., Feranec R.S., Wing S.L., Shabel A.B. Assessing the causes of Late Pleistocene extinctions on the continents. Science 2004, 306:70-75.
12. Barrell D.J.A., Alloway B.V., Shulmeister J., Newnham R.M. Towards a Climate Event Stratigraphy for New Zealand over the Past 30,000 years 2005, Institute of Geological and Nuclear Sciences, Science Report 2005/07 and Poster.
13. Beaumont M., Zhang W., Balding D. Approximate Bayesian computation in population genetics. Genetics 2002, 162:2025.
14. Bourdon E., Castanet J., Ricqles A., Scofield P., Tennyson A., Lamrous H., Cubo J. Bone growth marks reveal protracted growth in New Zealand kiwi (Aves, Apterygidae). Biol. Lett. 2010, 5:639-642.
15. Brock F., Higham T.F.G., Ditchfield P., Bronk Ramsey C. Current pretreatment methods for AMS radiocarbon dating at the Oxford Radiocarbon Accelerator Unit (ORAU). Radiocarbon 2010, 52:103-112.
16. Bronk Ramsey C., Higham T.F.G., Bowles A., Hedges R. Improvements to the pretreatment of bone at Oxford. Radiocarbon 2004, 46:155-163.
17. Bronk Ramsey C., Higham T.F.G., Leach P. Towards high precision AMS: progress and limitations. Radiocarbon 2004, 46:7-24.
18. Bunce M., Worthy T.H., Ford T., Hoppitt W., Willerslev E., Drummond A., Cooper A. Extreme reversed sexual size dimorphism in the extinct New Zealand moa Dinornis. Nature 2003, 425:172-175.
19. 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. The evolutionary history of the extinct ratite moa and New Zealand Neogene paleogeography. P. Natl. Acad. Sci. U. S. A. 2009, 106:20646-20651.
20. Burge P.I., Schulmeister J. An MIS 5a to MIS 4 (or early MIS 3) environmental and climatic reconstruction from the northwest South Island, New Zealand, using beetle fossils. J. Quat. Sci. 2007, 22:501-516.
21. Burnham K.P., Anderson D.R. Model Selection and Inference: A Practical Information - Theoretic Approach 2002, Springer-Verlag, New York. second ed.
22. Burrows C.J., Greenland D.E. An analysis of the evidence for climate change in New Zealand in the last thousand years: evidence from diverse natural phenomena and from instrumental records. J. Roy. Soc. New Zeal. 1979, 9:321-373.
23. Clark G.R., Petchey P., McGlone M.S., Bristow P. Faunal and floral remains from Earnscleugh Cave, Central Otago, New Zealand. J. Roy. Soc. New Zeal. 1996, 26:363-380.
24. Cooper A., Poinar H.N. Ancient DNA: do it right or not at all. Science 2000, 289:1139.
25. Cooper A., Fox-Lalueza C., Anderson A., Rambaut A., Austin J., Ward R. Complete mitochondrial genome sequences of two extinct moas clarify ratite evolution. Nature 2001, 409:704-707.
26. The Natural History of Southern New Zealand 2004, Otago University Press, Dunedin. J. Darby, R.E. Fordyce, A. Mark, K. Probert, C. Townsend (Eds.).
27. Drummond A.J., Rambaut A. BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evol. Biol. 2007, 7:214.
28. Drummond A.J., Rambaut A., Shapiro B., Pybus O.G. Bayesian coalescent inference of past population dynamics from molecular sequences. Mol. Biol. Evol. 2005, 22:1185-1192.
29. Field J.H., Fillios M., Wroe S. Chronological overlap between humans and megafauna in Sahul (Pleistocene Australia - New Guinea): a review of the evidence. Earth Sci. Rev. 2008, 89:97-115.
30. Gemmell N.J., Schwartz M.K., Robertson B.C. Moa were many. P. Roy. Soc. Lond. B Bio. 2004, 271:S430-S432.
31. 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. Checklist of the Birds of New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency, Antarctica 2010, Ornithological Society of New Zealand and Te Papa Press, Wellington. fourth ed.
32. Golledge N.R., Mackintosh A.M., Anderson B.M., Buckley K.M., Doughty A.M., Barrell D.J.A., Denton G.H., Vandergoes M.J., Andersen B.G., Schaefer J.M. Last Glacial Maximum climate in New Zealand inferred from a modelled Southern Alps icefield. Quat. Sci. Rev. 2012, 46:30-45.
33. Hajdas I., Lowe D.J., Newnham R.M., Bonani G. Timing of the late-glacial climate reversal in the Southern Hemisphere using high resolution radiocarbon chronology for Kaipo bog, New Zealand. Quat. Res. 2006, 65:340-345.
34. Ho S.Y.W., Larson G. Molecular clocks: when times are a-changin'. Trends Genet. 2006, 22:79-83.
35. Ho S.Y.W., Shapiro B. Skyline-plot methods for estimating demographic history from nucleotide sequences. Mol. Ecol. Res. 2011, 11:423-434.
36. Ho S.Y.W., Phillips M.J., Cooper A., Drummond A.J. Time dependency of molecular rate estimates and systematic overestimation of recent divergence times. Mol. Biol. Evol. 2005, 22:1561-1568.
37. Huynen L., Millar C.D., Scofield R.P., Lambert D.M. Nuclear DNA sequences detect species limits in ancient moa. Nature 2003, 425:175-178.
38. Kaplan M.R., Schaefer J.M., Denton G.H., Barrell D.J.A., Chinn T.J.H., Putnam A.E., Anderson B.J., Finkel R.C., Schwartz R., Doughty A.M. Glacier retreat in New Zealand during the Younger Dryas stadial. Nature 2010, 467:194-197.
39. Kumar S., Tamura K., Nei M. MEGA3: integrated software for molecular evolutionary genetics analysis and sequence alignment. Brief. Bioinform. 2004, 5:150-163.
40. Lambert D.M., Baker A., Huynen L., Haddrath O., Herbert P.D.N., Miller C.D. Is a large-scale DNA-based inventory of ancient life possible?. J. Hered. 2005, 96:279-284.
41. Leonard J.A., Wayne R.A., Cooper A. Population genetics of ice age brown bears. P. Natl. Acad. Sci. U. S. A. 2000, 97:1651-1654.
42. Lorenzen E.D., Nogues-Bravo D., Orlando L., Weinstock J., Binladen J., Marske K.A., Ugan A., Borregaard M.K., Gilbert M.T.P., Nielsen R., Ho S.Y.W., Goebel T., Graf K.E., Byers D., Stenderup J.T., Rasmussen M., Campos P.F., Leonard J.A., Koepfli K.P., Froese D., Zazula G., Stafford T.W., Sorensen K.A., Batra P., Haywood A.M., Singarayer J.S., Valdes P.J., Boeskorov G., Burns J.A., Davydov S.P., Haile J., Jenkins D.L., Kosintsev P., Kuznetsova T., Lai X., Martin L.D., McDonald H.G., Mol D., Meldgaard M., Munch K., Stephan E., Sablin M., Sommer M., Sipko T., Scott E., Suchard M.A., Tikhonov A., Willerslev R., Wayne R.K., Cooper A., Hofreiter M., Sher A., Shapiro B., Rahbek C., Willerslev E. Species-specific responses of Late Quaternary megafauna to climate and humans. Nature 2011, 479:359-364.
43. Lorrey A.M., Vandergoes M., Almond P., Renwick J., Stephens T., Bostock H., Mackintosh A., Newnham R., Williams P.W., Ackerley D., Neil H., Fowler A.M. Palaeocirculation across New Zealand during the last glacial maximum at ∼21 ka. Quat. Sci. Rev. 2012, 36:189-213.
44. Macken A.C., Prideaux G.J., Reed E.H. Variation and pattern in the responses of mammal faunas to Late Pleistocene climatic change in southeastern South Australia. J. Quat. Sci. 2012, 27:415-424.
45. Martin P.S. Prehistoric overkill: the global model. Quaternary Extinctions: A Prehistoric Revolution 1984, 354-403. Arizona University Press, Tuscon. P.S. Martin, R.G. Klein (Eds.).
46. McGlone M.S. New Zealand. Vegetation History 1988, 557-599. Kluwer Academic Publishers, Norwell. B. Huntley, T. Webb (Eds.).
47. McGlone M.S., Mark A.F., Bell D. Late Pleistocene and Holocene vegetation history, Central Otago, South Island, New Zealand. J. Roy. Soc. New Zeal. 1995, 25:1-22.
48. McGlone M.S., Newnham R.M., Moar N.T. The vegetation cover of New Zealand during the Last Glacial Maximum: do pollen records under-represent woody vegetation?. Terra Australis 32 2010, ANU E-Press, Australian National University, Canberra. S.G. Haberle, J. Stevenson, M. Prebble (Eds.).
49. McWethy D.B., Whitlock C., Wilmshurst J.M., McGlone M.S., Fromont M., Li X., Krall A.D., Hobbs W.O., Fritz S.C., Cook E.R. Rapid landscape transformation in South Island, New Zealand, following initial Polynesian settlement. P. Natl. Acad. Sci. U. S. A. 2010, 107:21343-21348.
50. Moar N.T. Late Quaternary vegetation. The Natural History of Canterbury 2008, Canterbury University Press, Christchurch. M. Winterbourn, G. Knox, C. Burrows, I. Marsden (Eds.).
51. Moar N.T., Suggate R.P. Pollen analysis of Late Otiran and Aranuian sediments at Blue Spur Road (S51), North Westland. New Zeal. J. Geol. Geop. 1973, 16:333-344.
52. Moar N.T., Suggate R.P. Contributions to the Quaternary history of New Zealand flora 8: interglacial and glacial vegetation in the Westport District South Island. New Zeal. J. Bot. 1979, 17:361-387.
53. Newnham R.M., Vandergoes M.J., Hendy C.H., Lowe D.J., Preusser F. A terrestrial palynological record for the last two glacial cycles from southwestern New Zealand. Quat. Sci. Rev. 2007, 26:517-535.
54. Nogues-Bravo D., Rodriguez J., Hortal J., Batra P., Araujo M.B. Climate change, humans, and the extinction of the woolly mammoth. PLoS Biol. 2008, 6:e79.
55. Nystrom V., Humphrey J., Skoglund P., Mckeown N.J., Vartanyan S., Shaw P.W., Linden K., Jakobsson M., Barnes I., Angerbjorn A., Lister A., Dalen L. Microsatellite genotyping reveals end-Pleistocene decline in mammoth autosomal genetic variation. Mol. Ecol. 2012, 21:3391-3402.
56. Orlando L., Metcalf J., Alberdi M.T., Telles-Antunes M., Bonhean D., Otte M., Martin F., Eisenmann V., Mashkour M., Morello F., Prado J.L., Salas-Gismondi R., Shockey B.J., Wrinn P.J., Vasil-ev S.K., Ovodov N.D., Cherry M.I., Hopwood B., Male D., Austin J.J., Hanni C., Cooper A. Revising the recent evolutionary history of equids using ancient DNA. P. Natl. Acad. Sci. U. S. A. 2009, 106:21754-21759.
57. Presscott G.W., Williams D.R., Balmford A., Green R.E., Manica A. Quantitative global analysis of the role of climate and people in explaining late Quaternary megafaunal extinctions. P. Natl. Acad. Sci. U. S. A. 2012, 109:4527-4531.
58. Prideaux G. Systematics and Evolution of the Stenurine Kangaroos 2004, University of California Publications, Berkeley.
59. Prideaux G.J., Roberts R.G., Megirian D., Westaway K.E., Hellstrom J.C., Olley J.M. Mammalian responses to Pleistocene climate change in southeastern Australia. Geology 2007, 35:33-36.
60. Prost S., Smirnov N., Fedorov V.B., Sommer R.S., Stiller M., Nagel D., Knapp M., Hofreiter M. Influence of climate warming on Arctic mammals? New insights from ancient DNA studies of the Collared Lemming Dicrostonyx torquatus. PLoS One 2010, 5:e10447.
61. Ramakrishnan U., Hadly E.A. Using phylochronology to reveal population histories: review and synthesis of 29 ancient DNA studies. Mol. Ecol. 2009, 18:1310-1330.
62. Rawlence, N.J., Cooper, A. Radiocarbon dating of the youngest moa (Aves: Dinornithiformes) from a natural site in New Zealand. J. Roy. Soc. New Zeal., . doi:10.1080/03036758.2012.658817.
63. Rawlence N.J., Scofield R.P., Wood J.R., Wilmshurst J.M., Moar N.T., Worthy T.H. New palaeontological data from the excavation of the Late Glacial Glencrieff miring bone deposit, north Canterbury, South Island, New Zealand. J. Roy. Soc. New Zeal. 2011, 41:217-236.
64. Rawlence N.J., Wood J.R., Armstrong K., Cooper A. DNA content and distribution in ancient feathers and potential to reconstruct the plumage of extinct avian taxa. P. Roy. Soc. Lond. B Bio. 2009, 276:3395-3402.
65. Reimer P.J., Baillie M.G.L., Bard E., Bayliss A., Beck J.W., Bertrand C.J.H., Blackwell P.G., Buck C.E., Burr G.S., Cutler K.B., Damon P.E., Edwards R.L., Fairbanks R.G., Friedrich M., Guilderson T.P., Hogg A.G., Hughen K.A., Kromer B., McCormac G., Manning S., Bronk Ramsey C., Reimer R.W., Remmele S., Southon J.R., Stuiver M., Talamo S., Taylor F.W., van der Plicht J., Weyhenmeyer C.E. INTCAL04 terrestrial radiocarbon age calibration, 0-26 Cal Kya BP. Radiocarbon 2004, 46:1029-1058.
66. Reimer P.J., Baillie M.G.L., Bard E., Bayliss A., Beck J.W., Blackwell P.G., Bronk Ramsey C., Buck C.E., Burr G.S., Edwards R.L., Friedrich M., Grootes P.M., Guilderson T.P., Hajdas I., Heaton T.J., Hogg A.G., Hughen K.A., Kaiser K.F., Kromer B., McCormac F.G., Manning S.W., Reimer R.W., Richards D.A., Southon J.R., Talamo S., Turney C.S.M., van der Plicht J., Weyhenmeyer C.E. INTCAL09 and MARINE09 radiocarbon age calibration curves, 0-50,000 years cal BP. Radiocarbon 2009, 51:1111-1150.
67. Ripple W.J., van Valkenburgh B. Linking top-down forces in Pleistocene megafaunal extinctions. BioScience 2010, 60:516-526.
68. Roberts R.G., Flannery T.F., Ayliffe L.K., Yoshida H., Olley J.M., Prideaux G.J., Laslett G.M., Baynes A., Smith M.A., Jones R., Smith B.L. New ages for the last Australian megafauna: continent-wide extinction about 46,000 years ago. Science 2001, 292:1888-1892.
69. Rule S., Brook B.W., Haberle S.G., Turney C.S.M., Kershaw A.P., Johnson C.N. The aftermath of megafaunal extinction: ecosystem transformation in Pleistocene Australia. Science 2012, 335:1483-1486.
70. Shulmeister J., McLea W.L., Singer C., McKay R.M., Hosie C. Late Quaternary pollen records from the Lower Cobb Valley and adjacent areas, north-west Nelson, New Zealand. New Zeal. J. Bot. 2003, 41:503-533.
71. Tennyson, A., Martinson, P., 2007 (reprint). Extinct Birds of New Zealand. Te Papa Press, Wellington.
72. Turvey S.T., Green O.R., Holdaway R.N. Cortical growth marks reveal extended juvenile development in New Zealand moa. Nature 2005, 435:940-943.
73. Vandergoes M.J., Newnham R.M., Hendy C.H., Lovell T.V., Preusser F., Fitzsimons S.J., Hogg A., Kasper H.U., Schluchter C. Regional isolation forcing of Late Quaternary climate change in the Southern Hemisphere. Nature 2005, 436:242-245.
74. Walker M., Johnsen S., Rasmussen S.O., Popp T., Steffensen J.-P., Gibbard P., Hoek W., Lowe J.J., Andrews J., Björck S., Cwynar L., Hughen K., Kershaw P., Kromer B., Litt T., Lowe D.J., Nakagawa T., Newnham R.M., Schwander J. Formal definition and dating of the GSSP (Global Stratotype Section and Point) for the base of the Holocene using the Greenland NGRIP ice core, and selected auxiliary records. J. Quat. Sci. 2009, 24:3-17.
75. Wardle P. Vegetation of New Zealand 1991, Cambridge University Press, Cambridge.
76. Weinstock J., Willerslev E., Sher A., Tong W., Ho S.Y.W., Rubenstein D., Storer J., Burns J., Martin L., Bravi C., Prieto A., Froese D., Scott E., Xulong L., Cooper A. Evolution, systematics, and phylogeography of Pleistocene horses in the new world: a molecular perspective. PLoS Biol. 2005, 3:e241.
77. 13C chronologies from South Island, New Zealand - did a global Younger Dryas really exist?. Earth Planet. Sc. Lett. 2005, 230:301-317.
78. Wilmshurst J.M., Anderson A.J., Higham T.F.G., Worthy T.H. Dating the late prehistoric dispersal of Polynesians to New Zealand using the commensal Pacific rat. P. Natl. Acad. Sci. U. S. A. 2008, 105:7676-7680.
79. Wood J.R. Moa gizzard content analysis: further information on the diet of Dinornis robustus and Emeus crassus, and the first evidence for the diet of Pachyornis elephantopus (Aves: Dinornithiformes). Rec. Cant. Mus. 2007, 21:27-39.
80. Wood J.R., Rawlence N.J., Rogers G.M., Austin J.J., Worthy T.H., Cooper A. Coprolite deposits reveal the diet and ecology of the extinct New Zealand megaherbivore moa (Aves: Dinornithiformes). Quat. Sci. Rev. 2008, 27:2593-2602.
81. Worthy T.H. Validation of Pachyornis australis Oliver (Aves: Dinornithiformes), a medium sized moa from the South Island New Zealand. New Zeal. J. Geol. Geop. 1989, 32:255-266.
82. Worthy T.H. Fossils of Honeycomb Hill 1993, Museum of New Zealand Te Papa Tongawera, Wellington.
83. Worthy T.H. Late Quaternary changes in the moa fauna on the West Coast of the South Island, New Zealand. Rec. South Aust. Mus. 1994, 27:125-134.
84. Worthy T.H. Fossil deposits in the Hodge Creek Cave System, on the northern foothills of Mt Arthur, Nelson, South Island, New Zealand. Notornis 1997, 44:111-124.
85. Worthy T.H. Quaternary fossil faunas of Otago, South Island, New Zealand. J. Roy. Soc. New Zeal. 1998, 28:421-521.
86. Worthy T.H. The Quaternary fossil avifauna of Southland, South Island, New Zealand. J. Roy. Soc. New Zeal. 1998, 28:537-589.
87. Worthy T.H. Changes induced in the New Zealand avifauna by climate during the last glacial-interglacial period. Proceedings of the 4th international meeting of the society of avian paleontology and evolution (Washington, D.C., June 1996). Sm. C. Paleob. 1999, 89:111-123.
88. Worthy T.H., Holdaway R.N. Quaternary fossil faunas from caves in Takaka Valley and on Takaka Hill, northwest Nelson, South Island, New Zealand. J. Roy. Soc. New Zeal. 1994, 24:297-391.
89. Worthy T.H., Holdaway R.N. Quaternary fossil faunas from caves on Mt Cookson, North Canterbury, New Zealand. J. Roy. Soc. New Zeal. 1995, 25:333-370.
90. Worthy T.H., Holdaway R.N. Quaternary fossil faunas, overlapping taphonomies, and palaeofaunal reconstruction in North Canterbury, South Island, New Zealand. J. Roy. Soc. New Zeal. 1996, 26:275-361.
91. Worthy T.H., Holdaway R.N. The Lost World of the Moa: Prehistoric Life of New Zealand 2002, Canterbury University Press, Christchurch.
92. Worthy T.H., Scofield R.P. 21st century advances in knowledge of the biology of moa (Aves: Dinornithiformes), diagnoses revised, and a new morphological analysis. New Zeal. J. Zool. 2012, 39:87-153.