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to

Palaecol Research

New Zealand was the last significant land mass colonised by humans.

Its biological processes functioned without human intervention

until the 14th century CE.

The record of those pristine systems is the best in the world:

it is a key window into the functioning of natural ecosystems.

Palaecol Research is dedicated to fundamental research on New Zealand’s extra-ordinary palaeobiological resources and in applying the results to underpin the restoration of ecosystems and biodiversity.

Our philosophy is "Understanding the causes is the best way of finding remedies".

To support the research, Palaecol Research publishes, via its imprint Turnagra Press, books on New Zealand birds, palaeobiology, restoration ecology, general natural history, and other topics.

Palaecol Research is based in Christchurch, New Zealand.

Contact us at: turnagra@gmail.com

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Pyramid Valley, North Canterbury, South Island, is one of the major late Holocene fossil sites in New Zealand. Its sediments contain not only one of the most diverse fossil faunas but also a high resolution record of the climate and environment of the past 5000 years

Palaecol Research Ltd

Learning from the past to build a sustainable future

Palaecol Research Ltd is a private company based in Christchurch, New Zealand, and engaged in research on past ecosystems and ecological restoration consultancy.

Its imprint is Turnagra Press, which publishes books on New Zealand birds, past and present, history, and educational material.

Palaecol Research is dedicated to doing fundamental research on New Zealand’s extra-ordinary palaeobiological resources and in applying the results to underpin the restoration of ecosystems and biodiversity on the principle that knowing what causes a problem is the best way of finding a remedy.

New Zealand was the last significant land mass colonised by humans: its biological processes functioned without human intervention until the 14th century. The record of those pristine systems is the best in the world and a key window into the natural functioning of ecosystems.

DIRECTOR - Dr Richard N Holdaway

The excavation at Pyramid Valley, January-February 2008). The lake was dry, as it was during the first major excavations in 1949. The excavation pit (2 x 1.8 m) is beneath the tent at middle left. Pyramid Valley is a Queen Elizabeth II Covenant and … — The excavation at Pyramid Valley, January-February 2008). The lake was dry, as it was during the first major excavations in 1949. The excavation pit (2 x 1.8 m) is beneath the tent at middle left. Pyramid Valley is a Queen Elizabeth II Covenant and the excavation was conducted under permit from the QE II National Trust and the owners, Mike & Jan Hodgen and family. The view is to the northwest.

In addition to directing the research programmes of Palaecol Research Ltd and running its publication arm Turnagra Press, Dr Holdaway retains a close association with the University of Canterbury, Christchurch, New Zealand, where he is an Joint Adjunct Professor in the School of Biological Sciences and the School of Earth and Environment.

He has international collaborative research programmes in isotope palaeoecology and palaeobiology, extinction biology, restoration ecology, and animal migration. He has published in avian systematics and has a special interest is in applying palaeoecological information to current conservation problems.

Dr Holdaway is a past President of the New Zealand Ornithological Society (Inc.) and a past editor of their journal Notornis. He received the D.L. Serventy Medal of the Royal Australasian Ornithologists' Union (Birdlife Australia) in 2003 and chaired the D.L. Serventy Medal Sub-Committee of Birdlife Australia.

Dr Richard N Holdaway

Publications to 18 September 2023

Refereed papers

91 Holdaway, R.N. 2023. Empirical comparisons between the past 5000 years of European and Eastern Mediterranean history and precipitation as recorded by ice accumulation in the GISP2 (Greenland) ice core. Oxford Open Climate Change 3(1), kgad007. https://doi.org/10.1093/oxfclm/kgad007.

90 Holdaway, R.N.; Kennedy, B.; Duffy, B.G.; Xu, J.; Oppenheimer, C. 2023. Re-assessment of tree-ring radiocarbon age series for the “Millennium Eruption” of Changbaishan / Paektu volcano in relation to the precise date of 946 CE. Journal of volcanology and geothermal research 437: 107787. https://doi.org/10.1016/j.volgeores.2023.107787.

89 Holdaway, R.N. 2022. Moa, climate, and eruptions: Radiocarbon ages on habitat-specific moa show that their distributions were controlled by volcanic eruptions as well as climate. Notornis 69(4): 228–242.

88 Holdaway, R.N.; Allentoft, M.E. 2022. A basic statistical approach to determining adult sex ratios of moa (Aves: Dinornithiformes) from sample series, with potential regional and depositional biases. Notornis 69(3): 158-173.

87 Westbury, M.V.; De Cahsan, B.; Shepherd, L.D.; Holdaway, R.N.; Duchene, D.A.; Lorenzen, E.D. 2022. Genomic insights into the evolutionary relationships and demographic history of kiwi. PLoS ONE 17 (10): e0266430. https://doi.org/10.1371/journal.pnoe.0266430.

86 Holdaway, R.N. 2022. Radiocarbon ages for two of the three South Island takahe Porphyrio hochstetteri (Aves: Rallidae) from Pyramid Valley, North Canterbury, New Zealand. Notornis 69(2): 112-115.

85 Holdaway, R.N. 2022. Extreme female-biased sexual size dimorphism in Euryapteryx moa (Aves: Dinornithiformes: Emeidae). Notornis 69(1): 58-65.

84 Johnston, A.G.; Duffy, B.G.; Holdaway, R.N. 2022. When the lonely goose? Implications of a revised history of Pyramid Valley lake, South Island, New Zealand, and its surrounding vegetation for a radiocarbon age for the only South Island goose (Cnemiornis calcitrans) recovered from the deposit. Notornis 69(1): 18-35.

83 Holdaway, R.N. 2021. Mind the gap: potential implications of the chronology of the South Island adzebill (Aptornis defossor) (Aves: Aptornithidae) at Pyramid Valley, North Canterbury, New Zealand. Notornis 68(4):.

82 Holdaway, R.N. 2021. Two new radiocarbon ages for Haast’s eagle (Hieraaetus moorei) (Aves: Accipitridae). Notornis 68(4): 278-282.

81 Holdaway, R.N. 2021. Radiocarbon ages for kakapo (Strigops habroptilus) (Strigopidae; Strigopinae) from the Pyramid Valley lake bed deposit, north-eastern South Island, New Zealand. Notornis 68 (3): 234-238.

80 Holdaway, R.N.; Sagar, P.M.; Briskie, J.V. 2021. Individual long-distance migrant Chrysococcyx cuckoos repeat carbon and nitrogen stable isotope ratios after moulting in non-breeding range on successive migrations. Notornis, 68(1), 65-71.

79 Holdaway, R.N.; Rowe, R.J. 2020. Palaeoecological reconstructions depend on accurate species identification: examples from South Island, New Zealand, Pachyornis (Aves: Dinornithiformes). Notornis 67 (2): 494-502.

78 Holdaway, R.N.; Duffy, B.; Kennedy, B. 2019. Reply to ‘Wiggle-match radiocarbon dating of the Taupo eruption’. Nature communications 10: 4668.

77 Holdaway, R.N.; Duffy, B.; Kennedy, B. 2018. Magmatic carbon bias in 14C dating of major eruptions: evidence from the Taupo eruption. Nature communications 9: 4110.

76 Horton, T.W.; Hauser, N.; Zerbini, A.; Francis, M.P.; Domeier, M.L.; Andriolo, A.; Costa, D.P.; Robinson, P.W. Duffy, C.A.J.; Nasby-Lucas, N.; Holdaway, R.N.; Clapham, P.J. 2017. Route fidelity during marine megafauna migration. Frontiers in marine science.. doi.org/10.3389/fmars.2017.00422.

75 Hawke, D.J.; Cranney, O.R.; Horton, T.W.; Bury, S.J.; Brown, J.C.S.; Holdaway, R.N. 2017. Foliar and soil N and δ15N as restoration metrics at Pūtaringamotu Riccarton Bush, Christchurch city. Journal of the Royal Society of New Zealand 47: 319-335.

74 Allentoft, M.E.; Heller, R.; Holdaway, R.N.; Bunce, M. 2015. Ancient DNA microsatellite analyses of the extinct New Zealand giant moa (Dinornis robustus) identify relatives within a single fossil site. Heredity 115: 481-487.

73 Hale, M.; Harrow, G.; Bradfield, P.; Cubrinovska, I.; Holdaway, R.N. 2015. Genetic similarity of Hutton's shearwaters (Puffinus huttoni) from two relict breeding populations. Notornis 62: 130-134.

72 Holdaway, R.N.; Allentoft, M.E.; Jacomb, C.; Oskam, C.L.; Beavan, N.R.; Bunce, M. 2014. An extremely low density human population exterminated New Zealand moa. Nature communications 5:5436. doi: 10.1038/ncomms6436.

71 Allentoft, M.E.; Heller, R.; Oskam, C.L.; Lorenzen, E.D.; Hale, M.L.; Gilbert, M.T.P.; Jacomb, C.; Holdaway, R.N.; Bunce, M. 2014. Extinct New Zealand megafauna were not in decline before human colonization. Proceedings of the National Academy of Sciences, USA. Published on-line 17 March 2014. doi/10.1073/pnas.1314972111.

70 Brassey, C.A.; Holdaway, R.N.; Packham, A.G.; Anné, J.; Manning, P.L.; Sellers, W.I. 2013. More than one way of being a moa: Differences in leg bone robustness map divergent evolutionary trajectories in Dinornithidae and Emeidae (Dinornithiformes). PLoS One 8 (12): 1-10. doi: 10.1371/journal.pone.0082668.

69 Holdaway, R.N.; Williams, M.J.; Hawke, D.J. 2013. A comparison of the pre-human and present isotopic niches of brown teal (Anas chlorotis): implications for conservation. Notornis 60(3): 233-244.

68 Williams, M.J.; Holdaway, R.N.; Rogers, K.M. 2012. Feeding environments of New Zealand's extinct merganser revealed by stable isotope analyses. Wildfowl 62: 190-203.

67 Nobes, D.C.; Johnston, A.G.; Holdaway, R.N.; Horton, T.W. 2012. Ground penetrating radar imaging of the late Quaternary Pyramid Valley lake deposit, North Canterbury, New Zealand: correlation with paleoclimatic records and the stratigraphic sequence. Proceedings of the 14th International Conference on Ground Penetrating Radar June 4-8, 2012, Shanghai, China: 595-598.

66 Oskam, C.L.; Allentoft, M.E.; Walter, R.; Scofield, R.P.; Haile, J.; Holdaway, R.N.; Bunce, M.; Jacomb, C. 2012. Ancient DNA analyses of early archaeological sites in New Zealand reveal extreme exploitation of moa (Aves: Dinornithiformes) at all life stages. Quaternary Science Reviews 52: 41-48. http://dx.doi.org/10.1016/j.quascirev.2012.07.007.

65 Allentoft, M.E.; Collins, M.; Harker, D.; Haile, J.; Oskam, C.L.; Hale, M.L.; Campos, P.F.; Samaniego, J.A.;, Gilbert, M.T.P.; Willerslev, E.;, Zhang, G.; Scofield, R.P.; Holdaway, R.N.; Bunce, M. (2012) The half-life of DNA in bone: measuring decay kinetics in 158 dated fossils. Proceedings of the Royal Society B: Biological Sciences 279(1748): 4724-4733.http://dx.doi.org/10.1098/rspb.2012.1745.

64 Holdaway, R.N.; Hawke, D.J.; Bunce, M.; Allentoft, M.E. 2011. Identification of an optimal sampling position for stable isotopic analysis of bone collagen of extinct moa (Aves: Emeidae). Notornis 58: 1-7.

63 Allentoft, M.E.; Scofield, R.P.; Oskam, C.M.; Hale, M.L.; Holdaway, R.N.; Bunce, M. 2011. A molecular characterization of a newly discovered megafaunal fossil site in North Canterbury, South Island, New Zealand. Journal of the Royal Society of New Zealand. First, 2011, 1-16. http://dx.doi.org/10.1080/030306758.2011.574821

62 Oskam, C.M.; Jacomb, C.; Allentoft, M.E.; Walter, R.; Scofield, R.P.; Haile, J.; Holdaway, R.N.; Bunce, M. 2011. Molecular and morphological analyses of avian eggshell excavated from a late thirteenth century earth oven. Journal of archaeological science 38 (10): 2589-2595.

61 Horton, T.W.; Holdaway, R.N.; Zerbini, A.N.; Hauser, N.; Garrigue, C.; Andriolo, A.; Clapham, P.J. 2011. Straight as an arrow: humpback whales swim constant course tracks during long-distance migration. Biology letters. Published online 20 April 2011. doi:10.1098/rsbl.2011.0279.

60 Allentoft, M.E.; Oskam, C.; Houston, J.; Hale, M.L.; Gilbert, M.T.P.; Rasmussen, M.; Spencer, P.; Jacomb, C.; Willerslev, E.; Holdaway, RN.; Bunce, M. 2011. Profiling the Dead: Generating microsatellite data from fossil bones of extinct megafauna - protocols, problems, and prospects. PLoS ONE 6 (1): e16670.

59 Holdaway, R.N.; Thorneycroft, J.M.; McClelland, P.; Bunce, M. 2010. Former presence of a parakeet (Cyanoramphus sp.) on Campbell Island, New Zealand subantarctic, with notes on the island's fossil sites and fossil record. Notornis 57: 8-18.

58 Allen, M.S.; Holdaway, R.N. 2010. Archaeological avifauna of Harataonga, Great Barrier Island, New Zealand: implications for avian palaeontology, Maori prehistory, and archaeofaunal recovery techniques. Journal of the Royal Society of New Zealand 40: 11-25.

57 Oskam, C.L.; Haile, J.; McLay, E.; Rigby, P.; Allentoft, M.E.; Olsen, M.E.; Bengtsson, C.; Miller, G.H.; Schwenninger, J-L.; Jacomb, C.; Walter, R.; Baynes, A.; Dortch, J.; Parker-Pearson, M.; Gilbert, M.T.P.; Holdaway, R.N.; Willerslev, E.; Bunce, M. 2010. Fossil avian eggshell preserves ancient DNA. 2010. Proceedings of the Royal Society B. Published online 10 March 2010 doi: 10.1098/rspb.2009.2019.

56 Allentoft, M.E.; Bunce, M.; Scofield, R.P.; Hale, M.L.; Holdaway, R.N. 2010. Highly skewed sex ratios and biased fossil deposition of moa: ancient DNA provides new insight on New Zealand's extinct megafauna. Quaternary science reviews 29: 753-762. doi:10.1016/j.quascirev.2009.11.022.

55 Steeves, T.E.; Holdaway, R.N.; Hale, M.L.; McLay, E.; McAllan, I.A.W.; Christian, M.; Hauber, M.; Bunce, M. 2010. Merging ancient and modern DNA: extinct seabird taxon rediscovered in the North Tasman Sea. Biology letters 6: 94-97. Published online before print August 12, 2009, doi:10.1098/rsbl.2009.0478

54 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. Proceedings of the National Academy of Sciences, USA 106: 20646-20651. Published online before print November 18, 2009, doi:10.1073/pnas.0906660106

53 Hawke, D.J.; Holdaway, R.N. 2009. Nutrient sources for forest birds captured within an undisturbed petrel colony, and management implications. Emu 109: 163-169.

52 Thomsen P.F.; Elias, S.; Gilbert, M.T.P.; Haile, J.; Munch, K.; Kuzmina, S.; Froese, D.G.; Sher, A.; Holdaway, R.N.; Willerslev, E. 2009. Non-destructive sampling of ancient insect DNA. PLoS ONE 4(4): e5048. doi:10.1371/journal.pone.0005048

51 Allentoft, M.E.; Schuster, S.C.; Holdaway, R.N.; Hale, J.; McLay, M.; Oskam, C.M.; Gilbert, M.T.P.; Spencer, P.B.S.; Willerslev, E.; Bunce, M. 2009. Identification of microsatellites from an extinct moa species using high throughput (454) sequence data. BioTechniques 46: 195-200.

50 Haile, J.; Holdaway, R.N.; Oliver, K.; Bunce, M.; Gilbert, M.T.P.; Nielsen, R.; Munch, K.; Ho, S.Y.W.; Shapiro, B.; Willerslev, E. 2007. Ancient DNA chronology within sediment deposits: are palaeobiological reconstructions possible and is DNA leaching a factor. Molecular biology and evolution 24: 982-989.

49 Holdaway, R.N.; Hawke, D.J.; Hyatt, O.; Wood, G.C. 2007. Stable isotopic (δ15N, δ13C) analysis of wood in trees growing in past and present colonies of burrow-nesting seabirds in New Zealand. I. δ15N in two species of conifer (Podocarpaceae) from a mainland colony of Westland petrels (Procellaria westlandica), Punakaiki, South Island. Journal of the Royal Society of New Zealand 37: 75-84.

48 Bunce, M.; Holdaway, R.N. 2006. New Zealand’s extinct giant eagle. Bulletin of the British Ornithologists’ Club 126A: 4-7.

47 Harrow, G.; Hawke, D.J.; Holdaway, R.N. 2006. Surface soil chemistry at an alpine procellariid breeding colony in New Zealand, and comparison with a lowland site. New Zealand journal of zoology 33: 165-174.

46 Hawke, D.J.; Holdaway, R.N. 2005. Avian assimilation and dispersal of carbon and nitrogen brought ashore by breeding Westland petrels Procellaria westlandica: a stable isotope study. Journal of zoology, London 266: 419-426.

45 Winn, J.M.; Holdaway, R.N. 2005. Egg predation by South Island kaka (Nestor meridionalis). Notornis 52(2): 106-108.

44 Turvey, S.T.; Green, Owen, R.; Holdaway, R.N. 2005. Cortical growth marks reveal extended juvenile development in New Zealand moa. Nature 435: 940-943.

43 Turvey, S.T.; Holdaway, R.N. 2005. Postnatal ontogeny, population structure and extinction of the giant moa Dinornis. Journal of morphology 265: 70-86.

42 Bunce, M.; Szulkin, M.; Lerner, H.R.L.; Barnes, I.; Shapiro, B.; Cooper, A.; Holdaway, R.N. 2005. Ancient DNA provides new insights into the evolutionary history of New Zealand’s extinct giant eagle. Public Library of Science, Biology 3: 44-46.

41 Harding, J.S.; Hawke, D.J.; Holdaway, R.N.; Winterbourn, M.J. 2004. Incorporation into stream food webs of marine-derived nutrients from petrel breeding colonies. Freshwater biology 49: 576-586.

40 Holdaway, R.N.; Jones, M.D.; Beavan Athfield, N.R. 2003. Establishment and extinction of a population of South Georgian diving petrel (Pelecanoides georgicus) at Mason Bay, Stewart Island, New Zealand, during the late Holocene. Journal of the Royal Society of New Zealand 33(3): 601-622.

39 Holdaway, R.N.; Jones, M.D.; Beavan Athfield, N.R. 2002. Late Holocene extinction of the New Zealand owlet-nightjar Aegotheles novaezealandiae (Scarlett, 1968). Journal of the Royal Society of New Zealand 32(4): 653-667.

38 Holdaway, R.N.; Jones, M.D.; Beavan Athfield, N.R. 2002. Late Holocene extinction of Chenonetta finschi van Beneden, 1875, an endemic, almost flightless, New Zealand duck. Journal of the Royal Society of New Zealand 32(4): 629-651.

37 Holdaway, R.N.; Roberts, R.G.; Beavan Athfield, N.R.; Olley, J.M.; Worthy, T.H. 2002. Optical dating of quartz sediments and accelerator mass spectrometry 14C dating of bone gelatin and moa eggshell: a comparison of age estimates for non-archaeological deposits in New Zealand. Journal of the Royal Society of New Zealand 32(3): 463-505.

36 Worthy, T.H.; Holdaway, R.N.; Alloway, B.V.; Jones, J.; Winn, J.; Turner, D. 2002. A rich Pleistocene-Holocene avifaunal sequence from Te Waka #1: terrestrial fossil vertebrate faunas from inland Hawke’s Bay, North Island, New Zealand. Part 2. Tuhinga 13: 1-38.

35 Holdaway, R.N.; Anderson, A.J. 2001. The avifauna of the Emily Bay archaeological site, Norfolk Island, South-west Pacific: a preliminary account. Records of the Australian Museum special issue. The Prehistoric Archaeology of Norfolk Island, Southwest Pacific. Editors A. Anderson and P. White. Records of the Australian Museum, Supplement: 87-102.

34 Holdaway, R.N.; Worthy, T.H. 2001. Publication of supplementary data for 14C AMS ages. Journal of the Royal Society of New Zealand 31(2): 453-456.

33 Holdaway, R.N.; Worthy, T.H.; Tennyson, A.J.D. 2001. A working list of breeding bird species of the New Zealand region at first human contact. New Zealand journal of zoology 28(2): 119-187.

32 Worthy, T.H.; Holdaway, R.N. 2000. Terrestrial fossil vertebrate faunas from inland Hawke’s Bay, North Island, New Zealand. Part 1. Records of the Canterbury Museum 14: 89-154.

31 Holdaway, R.N.; Jacomb, C. 2000. Rapid extinction of the moas (Aves: Dinornithiformes): model, test, and implications. Science 287(5461, 24 March): 2250-2254.

30 Worthy, T.H.; Holdaway, R.N. 1999. Introduction for Hartree, W.H., The nesting behaviour of moas. Notornis 46(4): 457-460.

29 Holdaway, R.N.; Beavan, N.R. 1999. Reliable 14C AMS dates on bird and Pacific rat Rattus exulans bone gelatin, from a CaCO3-rich deposit. Journal of the Royal Society of New Zealand 29(3): 185-211.

28 Holdaway, R.N. 1999. A spatio-temporal model for the invasion of the New Zealand archipelago by the Pacific rat Rattus exulans. Journal of the Royal Society of New Zealand 29(2): 91-105.

27 Holdaway, R.N. 1999. Introduced predators and avian extinction in New Zealand. pp. 189-238 In: MacPhee, R.D.E. (ed.). Extinctions in near time: causes, contexts, and consequences. Advances in vertebrate paleobiology. New York, Kluwer Academic/Plenum Publishers. 394 p.

26 Hawke, D.J.; Holdaway, R.N.; Causer, J.E.; Ogden, S. 1999. Soil indicators of pre-European seabird breeding in New Zealand at sites identified by predator deposits. Australian journal of soil research 37: 103-113.

25 Holdaway, R.N.; Anderson, A.J. 1998. 14C AMS dates on Rattus exulans bones from natural and archaeological contexts on Norfolk Island South-west Pacific. Archaeology in New Zealand 41(3): 195-198.

24 Worthy, T.H.; Holdaway, R.N.; Sorenson, M.D.; Cooper, A.C. 1997. Description of the first complete skeleton of the extinct New Zealand goose Cnemiornis calcitrans (Aves: Anatidae) and a reassessment of the relationships of Cnemiornis. Journal of zoology, London 243: 695-723.

23 Holdaway, R.N.; Worthy, T.H. 1997. A reappraisal of the Late Quaternary fossil vertebrates of Pyramid Valley Swamp, North Canterbury, New Zealand. New Zealand journal of zoology 24: 69-121.

22 Holdaway, R.N. 1996. Arrival of rats in New Zealand. Nature 384 (no. 6606, 21 November): 225-226.

21 Worthy, T.H.; Holdaway, R.N. 1996. Quaternaryfossilfaunas, overlapping taphonomies, and palaeofaunal reconstruction in North Canterbury, South Island, New Zealand. Journal of the Royal Society of New Zealand 26(3): 275-361.

20 Holdaway, R.N.; Worthy, T.H. 1996. Diet and biology of the laughing owl Sceloglaux albifacies (Aves: Strigidae) on Takaka Hill, Nelson, New Zealand. Journal of zoology, London 239: 545-572.

19 Worthy, T.H.; Holdaway, R.N. 1996. Taphonomy of two New Zealand microvertebrate deposits, Takaka Hill, Nelson, New Zealand, and identification of the avian predator responsible. Historical biology 12: 1-24.

18 Holdaway, R.N. 1995. A fossil record of the Black Stilt Himantopus novaezelandiae Gould, 1841. New Zealand natural sciences 22: 69-74.

17 Worthy, T.H.; Holdaway, R.N. 1995. Quaternary fossil faunas from caves on Mt Cookson, North Canterbury, South Island, New Zealand. Journal of the Royal Society of New Zealand 25(3): 333-370.

16 McGlone, M.S.; Anderson, A.J. Holdaway, R.N. 1994. An ecological approach to the Polynesian settlement of New Zealand. pp. 136-163 In: Sutton, D.G. (ed.) The origins of the first New Zealanders. Auckland, Auckland University Press. 269 p.

15 Holdaway, R.N.; Worthy, T.H. 1994. A new fossil species of shearwater Puffinus from the Late Quaternary of the South Island, New Zealand, and notes on the biogeography of the Puffinus gavia superspecies. Emu 94: 201-215.

14 Holdaway, R.N. 1994. An exploratory phylogenetic analysis of the genera of the Accipitridae, with notes on the biogeography of the family. pp. 601-649 In: Meyburg, B-U.; Chancellor, R.D. (ed.) Raptor conservation today. London & Berlin, WWGBP/The Pica Press.

13 Worthy, T.H.; Holdaway, R.N. 1994. Quaternary fossil faunas from caves in Takaka Valley and on Takaka Hill, northwest Nelson, South Island, New Zealand. Journal of the Royal Society of New Zealand 24(3): 297-391.

12 Worthy, T.H.; Holdaway, R.N. 1994. Scraps from an owl’s table— predatoractivityas a significant taphonomic process newly recognised from New Zealand Quaternary deposits. Alcheringa 18: 229-245.

11 Holdaway, R.N. 1993. First North Island fossil record of Kea, and morphological and morphometric comparison of Kea and Kaka. Notornis 40(2): 95-108.

10 Worthy, T.H.; Holdaway, R.N. 1993. Quaternary fossil faunas from caves in the Punakaiki area, West Coast, South Island, New Zealand. Journal of the Royal Society of New Zealand 23(3): 147-254.

9 Holdaway, R.N. 1991. Sibley et al.’s classification of living birds applied to the New Zealand list. Notornis 38(2): 152-162.

8 Holdaway, R.N. 1990. Seeing -ii to -i; proposed changes to the specific names of some New Zealand birds. New Zealand natural sciences 17: 85-88.

7 Holdaway, R.N. 1990. Harpagornis assimilis Haast, 1874, a synonym of Harpagornis moorei Haast, 1872 (Aves: Accipitridae). New Zealand natural sciences 17: 39-47.

6 Holdaway, R.N. 1990. Changes in the diversity of New Zealand forest birds. New Zealand journal of zoology 17(3): 309-321.

5 Duncan, K.W.; Holdaway, R.N. 1989. Footprint pressures and locomotion of moas and ungulates and their effects on the New Zealand indigenous biota through trampling. New Zealand journal of ecology 12 (supplement): 97-101.

4 Holdaway, R.N. 1989. New Zealand’s pre-human avifauna and its vulnerability. New Zealand journal of ecology 12 (supplement): 11-25.

3 Holdaway, R.N. 1988. The New Zealand passerine list: What if Sibley & Ahlquist are right? Notornis 35(1): 63-70.

2 Cunningham, J.B.; Holdaway, R.N. 1986. Morphology and head colour in the Yellowhead. Notornis 33: 33-36.

1 Holdaway, R.N. 1980. Royal Spoonbills nesting near Blenheim. Notornis 27(2): 168-169.

Reviews

2 Holdaway, R.N. 2020. Review of Grusin, Richard (ed.), After extinction. Center for 21st Century Studies. University of Minnesota Press, Minneapolis. 2018. The quarterly review of biology 95 (1): 92-93.

1 Holdaway, R.N. 2018. Review of Heads, Michael, Biogeography and evolution in New Zealand. CRC Biogeography Series. CRC Press, Boca Raton (Florida). The quarterly review of biology 93: 268.

BOOKs

1 Holdaway, R.N.; Morris, R.B. 2015. Pyramid Valley and beyond: Discovering the prehistoric birdlife of North Canterbury, New Zealand. Christchurch, Turnagra Press, 44 p.

2 Christian, M.L.; Holdaway, R.N.; Smith, J.L.; Coyne, P.D. 2012. A comparative atlas of bird distribution in the Norfolk Island Group, Southwest Pacific Ocean, 1978-2005. The Authors & Norfolk Island Fauna and Flora Society, Norfolk Island. 96 p.

3 Worthy, T.H.; Holdaway, R.N. 2002. The lost world of the moa: terrestrial animals of prehistoric New Zealand. Bloomington, Indiana University Press and Canterbury University Press. 718 p.

PREPRINT

Holdaway, R.N. 2021. Palaeobiological evidence for Southern Hemisphere Younger Dryas and volcanogenic cold periods. Climate of the Past Discussion [Preprint]. doi: https://doi.org/10.5194/cp-2021-154,2021

In Press

Submitted - In review

1 Holdaway, R.N. Empirical comparisons between the past 5000 years of European and eastern Mediterranean history and precipitation as recorded by ice accumulation in the GISP2 (Greenland) ice core. Oxford Open Climate Change.

BOOK SECTIONS

7 Holdaway, R.N. Zoology of the Voyage of HMS Erebus and Terror. In: Treasures of the Library. Christchurch, Canterbury University Press.

6 King, C.M.; Roberts, C.D.; Bell, B.D.; Fordyce, R.E.; Nicoll, R.S.; Worthy, T. H.; Pauling, C.D.; Hitchmough, R.A.; Keyes, I.W.; Baker, A.N.; Stewart, A.L.; Hiller, N.; McDowall, R.M.; Holdaway, R.N.; McPhee, R.P.; Schwarzhans, W.W.; Tennyson A.J.D.; Rust, S.; Macadie, I. 2009. Phylum Chordata: Lancelets, Fishes, Amphibians, Reptiles, Birds, Mammals. pp. 431-551 In: Gordon, D.P. (ed.) New Zealand inventory of biodiversity. Vol. 1. Kingdom Animalia: Radiata, Lophotrophozoa, Deuterostomia. Christchurch, Canterbury University Press.

5 Holdaway, R.N.; Worthy, T.H. 2008. Late Quaternary avifauna. pp. 445-492 In: Winterbourn, M.J.; Knox, G.A.; Burrows, C.J.; Marsden, I.D. (ed.) The natural history of Canterbury, 3rd ed. Christchurch, Canterbury University Press.

4 Holdaway, R.N.; Worthy, T.H. 2006. Evolution of New Zealand and its vertebrates. pp. 111-128 In: Merrick, J.R.; Archer, M.; Hickey, G.M.; Lee, M.S.Y. (ed.) Evolution and biogeography of Australasian vertebrates. Sydney, Auscipub. 942 p.

3 Holdaway, R.N. 2005. Birds. pp. 128-160 In: Jones, M.B.; Marsden, I. (ed.) Life in the estuary: illustrated guide and ecology. Christchurch, University of Canterbury Press. 179 p.

2 Holdaway, R.N. 1988. Birds. pp. 97-99 In: Stevens, G.R.; McGlone, M.S.; McCulloch, B. Prehistoric New Zealand . Auckland, Heinemann Reed. 128 p.

1 Holdaway, R.N. 1983. Birds. pp. 103-148 In: Jones, M.B. Animals of the estuary shore: illustrated guide and ecology. Christchurch, University of Canterbury Press. 162 p.

EDITED VOLUMES

1 Holdaway, R.N. (Editor) 1994. Chatham Islands ornithology. Notornis 41 (supplement). 208 p.

CONFERENCE ABSTRACTS and posters

26 Holdaway, R.N.; Kennedy, B.; Duffy, B.G. Poster - Potential implications for Taupo eruption wiggle match dates from off edifice radiocarbon series bracketing a distal lobe of the ignimbrite. International Association of Volcanology and Chemistry of the Earth’s Interior (IAVCEI) conference, Rotorua, New Zealand, January-February 2023

25 Holdaway, R.N. Poster - Moa as monitors of volcanogenic environment change. Ornithological Society of New Zealand Conference, Christchurch, June 2022

24 Holdaway, R.N. The Manawatū-Tararua Highway fossil avifauna: a unique window into the late Pleistocene of North Island, New Zealand. Ornithological Society of New Zealand Conference, Christchurch, June 2022

23 Holdaway, R.N. Poster - Hidden diversity in the New Zealand avifauna. Ornithological Society of New Zealand Conference, Wellington, June 2019

22 Holdaway, R.N.; Sagar, P.M.; Briskie, J.V. First tracking of shining cuckoo migration using geolocators and investigation of their non-breeding habitat use by stable isotope analysis.Ornithological Society of New Zealand Conference, Wellington, June 2019

21 Holdaway, R.N.; Christian, M. Migration of wedge-tailed shearwaters. Ornithological Society of New Zealand Conference, Blenheim

20 Holdaway, R.N. Speciation in oceanic seabirds. Ornithological Society of New Zealand Conference, Nelson

19 Holdaway, R.N.; Jacomb, C.; Walter, R.; Brooks, E.; Oskam, C.; Allentoft, M.; Bunce, M. 2012. How long did people and megafauna co-exist in New Zealand? Re-dating the occupation of Wairau Bar and the extinction of moa (Aves: Dinornithiformes). 21st International Radiocarbon Conference, Paris.

18 Horton, T.W.; Holdaway, R.N.; Zerbini, A.; Andriolo, A.; Clapham, P.J. 2010. Solar-magnetic orientation during leatherback turtle (Dermochelys coriacea), great white shark (Carcharodon carcharias), arctic tern (Sterna paradisaea), and humpback whale (Megaptera novaeangliae) long-distance migration. Poster B33F-0447: American Geophysical Union Fall Meeting, San Francisco, 13-17 December, 2010.

17 Holdaway, R.N.; Horton, T.W.; Rowe, R.J.; Sagar, P.M.; Clapham, P. 2009. Allopatric speciation in pelagic vertebrates: Becoming isolated in a continuum. Evolution 2009: Darwin Bicentennial Conference, Melbourne, Australia.

16 Holdaway, R.N.; Hawke, D.J.; Christian, M. 2007. The present obscuring the past: seabirds and island conservation biology. Quaternary international 167-168 supplement: 174. XVII INQUA Congress, Cairns, Australia.

15 Wilson, F.K.; Steeves, T.; Hale, M.; Bunce, M.; Christian, M.; Holdaway, R.N. 2007. Brown-eyed boobies: from extinct to extant to endangered? International Waterbirds Conference, Barcelona.

14 Sauvageot, R.; Holdaway, R.N.; McFarlane, D. 2005. Fluorine relative dating and taphonomy of moa bone from Pyramid Valley, New Zealand. Poster: Geological Society of America Annual Meeting, Salt Lake City, October 16-19, 2005. GSA Abstracts with Programs 37, no. 7.

13 Holdaway, R.N.; Beavan Athfield, N.R.; Roberts, R.G. 2003. OSL and 14C ages on Pacific rat bones from New Zealand. 18th International Radiocarbon Conference, Wellington, 1-5 September: 114.

12 Holdaway, R.N. 2003. Causes and effects of Late Holocene vertebrate extinctions in New Zealand. XVI INQUA Congress, Reno, USA, July 23-30: 88.

11 Holdaway, R.N.; Hawke, D.J.; Worthy, T.H. 2002. Palaebiological perspectives on Westland birds. Notornis 49(3): 192.

10 Holdaway, R.; Holdaway, R. 2002. ARPASAT – Archaeological and Palaeontological surveys using satellite technology. 29th International Symposium on Remote Sensing of Environment, Buenos Aires, 8-12 April 2002.

9 Worthy, T.H.; Holdaway, R.N. 2001. On petrels, fallout from catastrophic eruptions, and ice ages - an unparalleled record from Te Waka in inland Hawke's Bay. Notornis 48(3): 183.

8 Ragano Beavan, N.; McFadgen, B.; Sparks, R.; Holdaway, R.N. 1999. Reliability of 14C AMS dating of small vertebrate bone: implications for the timing of Holocene extinctions by the introduction of Rattus exulans to New Zealand. Proceedings of the Xth Archaeozoological Conference, Victoria, Canada.

7 Hawke, D.; Holdaway, R.N.; Causer, J.; Ogden, S. 1999. Chemical tracers of former mainland seabird breeding colonies. Notornis 46(3): 407.

6 Holdaway, R.N. 1999. The late Holocene avifauna of Canterbury. Notornis 46(3): 406-407.

5 Holdaway, R.N.; Roberts, R.G.; Worthy, T.H. 1999. OSL and radiocarbon dating of late Holocene faunal sites in New Zealand. Records of the Western Australian Museum supplement no. 57: 403.

4 Holdaway, R.N. 1999. Times to extinction and the New Zealand extinction chronology. Records of the Western Australian Museum supplement no. 57: 403.

3 Brathwaite, D.H.; Holdaway, R.N. 1987. A preliminary reconstruction of Harpagornis moorei: top predator in the moa’s world. New Zealand journal of ecology 10: 162.

2 Duncan, K.W.; Holdaway, R.N. 1987. Footprint pressures of moas and ungulates. New Zealand journal of ecology 10: 161.

1 Holdaway, R.N.; Brathwaite, D.H. 1987. New Zealand’s prehuman avian ecology: a scenario (without moas). New Zealand journal of ecology 10: 161.

letters

5 Holdaway, R.N. 2023. Egg predation by South Island kaka. Notornis 70 (2): 96.

4 Holdaway, R.N. 2021. Shore plover relationships. Notornis 68 (2): 182.

3 Holdaway, R.N. 2010. Comment on Rhodes et al. (2009) The roles of predation, microclimate and cavity abundance in the evolution of New Zealand’s tree-cavity nesting avifauna. Notornis 56: 190-200. Notornis 57 (2): 110-111.

2 Scofield, P.; Worthy, T.; Holdaway, R.N.; Bunce, M.; Cooper, A.; Tennyson, A. 2005. Recent claims for more moa and huge errors in museum identifications — cutting through the spin. Te Ara - Journal of Museums Aotearoa 30(2): 29-31.

1 Holdaway, R.N. 1999. Errors and omissions in a list. Notornis 46: 320.

popular publications

17 Holdaway, R.N. 2019. Caveat collectors: Birds, extinction and Bill Hammond. Christchurch Art Gallery Bulletin B.198: 26-33.

16 Holdaway, R.N. 2017. Haast’s eagle. pp. 22-30 In: Easther, E. (ed.) Bird words: New Zealand writers on birds. Vintage (Penguin Random House), Auckland, New Zealand. Revisd and reprinted from New Zealand geographic4: 56-64.

15 Holdaway, R.N.; Christian, M. 2010. Stopping the fourth wave: Conservation and restoration of the Norfolk Island ecosystem. pp. 30-35 In: Kirkwood, J.; O'Connor, J. (Comp.). The state of Australia's birds 2010: Islands and birds. Wingspan 20 (4) Supplement.

14 Holdaway, R.N. 2006. Two roads to Paradise, Lost. New Zealand geographic 80: 20-22.

13 Holdaway, R.N. 2005. Campbell’s lost parrot and other cautionary tales. New Zealand geographic 73: 8-10.

12 Hawke, D.; Holdaway, R.N. 2003. Mainland petrel breeding as a driver of terrestrial ecosystem processes. ConScience (Conservation science newsletter) 47: 6-7.

11 Holdaway, R.N. 2002. [Review] “Feral future” (Tim Low, Viking 1999) and “The new nature: winners and losers in wild Australia” (Tim Low, Viking 2002). New Zealand geographic 59: 11-13.

10 Holdaway, R.N. 2002. [Review] “Eternal frontier” (Tim Flannery, Atlantic Monthly Press, New York). New Zealand geographic 56: 110-111.

9 Holdaway, R.N. 1999. On extinction. New Zealand geographic 44: 9-11.

8 Holdaway, R.N. 1998. Grounded! Why do some birds walk? New Zealand geographic 37: 102-121.

7 Worthy, T.H.; Holdaway, R.N. 1996. Laughter in the night. New Zealand geographic 32: 86-100.

6 Holdaway, R.N. 1996. Wide-eyed and wonderful. New Zealand geographic 32: 99.

5 Holdaway, R.N. 1996. The owls of New Zealand. New Zealand geographic 32: 96-97.

4 Holdaway, R.N. 1996. Stephens Island - a chance lost. New Zealand geographic 32: 81-82.

3 Holdaway, R.N. 1996. [Review] “The Song of the Dodo” (David Quammen, Hutchinson, 1996). New Zealand geographic 31: 6, 18-19.

2 Holdaway, R.N.; Worthy, T.H. 1991. Lost in time. New Zealand geographic 12: 51-68.

1 Holdaway, R.N. 1989. Terror of the forests. New Zealand geographic 4: 56-64.

TECHNICAL REPORTS

1 Holdaway, R.N. 2010. The identity, geological age, and significance of moa remains recovered from the

route of the East Taupo Arterial - November 2009. Report to the Taupo District Council, June 2010.

outreach

U3A lectures

School visits

GROUNDED! WHY DO SOME BIRD WALK?

Ah, to be as free as a bird! To be able to take off and soar away, leaving all cares behind. To rise up on wings of eagles; to glide across the oceans like an albatross.

For as long as humans have dreamed, they have dreamt of emulating birds: of sprouting wings and taking flight. Indeed, a bird in flight seems to us the very quintessence of unfettered liberty. Jonathan Livingston liberty.

Birds have used their ability well. With mastery of the air, they have colonised more parts of the glade than any other group of vertebrates. For 65 million years they have ruled the daytime skies, and they contested them with pterosaurs for millions of years before that.

Given their success as a group, it seems puzzling that many birds have lost the ability to fly — handed in their wings, so to speak. Many of their names are familiar to us, for New Zealand happens to be a world centre for flightlessness.

We are internationally known, of course, by our snuffling, shuffling national symbol, the kiwi. And crowds flock to the natural history sections of our museums to view our “giraffes on two legs” — the several species of extinct moa. Visitors to semi-remote South Island destinations such as the caves on Takaka Hill or the seal colony at Cape Foulwind have to run the gauntlet of weka eager for food or for anything bright and shiny. Fly they cannot, but they can run fast enough to beat the careless tourist for his biscuit or hop into a car when the door is left ajar, and they can jump high enough to snatch car keys out of the hand of the unwary.

The list grows longer: ostriches have left the zoo to go down to the farm, where there have been joined lately by emus. Penguins are so well-known that they strut products on TV. And hardly a week goes by that one or other of our most endangered birds — kakapo, takahe and, increasingly, kiwi themselves — doesn’t get a walk-on part to report some success or catastrophe in the fight to save our heritage.

Step back in time and a catalogue of flightless birds from these islands increases yet again: three flightless wrens (the only flightless songbirds in the world[i]), two flightless geese, a duck or two, seven rails and two adzebills. All as flightless as the dinosaurs that birds evolved from, and all now as extinct, as the dodo.

If flight is the defining characteristic of a bird, why would these and many other species choose to give it up?

There is no single reason, but the answers all have much to do with how much it costs to fly. We all know that flight is expensive; your airline ticket makes that clear. And the costs are the same for birds. Just as an aeroplane needs an appropriate structure, fuel and maintenance, so birds need to grow the wings and muscles to get them off the ground, and then maintain the structures and generate the energy to use them.

Birds pay that price to secure the advantages of more efficient mobility, for to fly a certain distance requires less energy than to walk or run the same course. This is simply a reflection of the relative cost of the different types of locomotion and energy terms. When one then considers that a blackbird can fly directly across a valley, where as a rat must pick its way down one side and up the other, around obstacles and up and down every small undulation — probably covering at least three times the distance as a blackbird — flight becomes an even more attractive proposition. For aquatic birds, swimming uses even less energy than flight, for bodies are naturally supported in water without expending energy. Standing upright on land or hovering in air both require energy before you even start to move forward.

Speed, too, is relevant to energy efficiency, and birds fare well here also. In general, higher speeds result in less overall energy expenditure to cover a given distance. Running a kilometre uses less energy than walking it. At the end of the run you may feel more exhausted, but this is because your muscles have demanded energy more rapidly than your heart and lungs are accustomed to supplying it, and parts of your person have gone into a temporary energy overdraft. Flight, by its very nature, is generally a very rapid form of locomotion. If a bird flies too slowly, then, like an aircraft, it will stall and plummet.

High-speed locomotion and relatively low energy demands allow birds to operate over much larger areas than comparably sized mammals or lizards. Birds are able to take advantage of highly dispersed food supplies that would be out of reach for earthbound animals. Improved mobility is useful even for short-range work. For a bird, the fruit or nectar in the next tree is not a long climb or dangerous jump away; it is just a short flight. It is far easier for a starling to move to a fresh field of grass grubs than it is for a hedgehog to find a new source of snails.

And birds can move with the seasons. Some migrate thousands of kilometres to take advantage of rich seasonal food supplies. Even in New Zealand, far from the great flyways of North America and the streams of hawks and stalks crossing the Straits of Gibraltar and the Bosporus, the migrations of birds have caught the imagination from prehistoric times: godwits congregating in the Far North before leaving for Siberia, streams of sooty shearwaters passing Otago Peninsula and shining cuckoos announcing their arrival from the Solomon Islands in spring.

With migration comes colonisation. Even tiny islands far from any other land usually have a bird fauna, while almost the only mammals to have colonised distant islands are bats. Flightless mammals, large or small, make bad overseas travellers. Though theoretically they could float on rafts of vegetation across large distances of ocean, they rarely do. Flight has given birds an edge in populating the planet, and this is reflected in the number of species alive today. Birds outnumber mammals in number of species by nearly three to one, and the ratio would have been far greater had not many birds been exterminated in the recent past by foreign predators introduced to their domain by humans.

Which brings up a key point about the costs and benefits of flying. There are good reasons (which we will presently explore) for a species to abandon flight, but for most birds there is an even better reason not to: to avoid being eaten. Being able to get around to new places or exploit distant resources are good reasons for birds to keep flying, but the crucial advantage is in escaping from danger. To get away and survive is to live and breed. Flying allows birds to live in environments filled with sharp claws and hungry mouths. That imperative can only be set aside when birds find themselves in places without predators. Those places have classically been islands.

Most oceanic islands had no resident mammals and therefore no predators on the ground which hunted by smell, and few that could hunt in the dark, under such circumstances, birds could “afford” to become flightless. On continents, the domain of mammals, the advantage of being able to fly away from a predator remained so important that there are almost no flightless birds from those parts of the globe. The few that do live on continents — ostriches and their relatives — are big, and their ancestors may have given up flight before there were any large predatory mammals to contend with.

On islands, where there were no claws or teeth to avoid, the way was open for some birds to save the energy normally spent on building and maintaining wings, of course, that meant living on the ground, and it also usually meant occupying a new ecological niche (the biological “slot” into which a species fits in an ecosystem). Often that niche was one that would have been occupied by a mammal, had it been around: for example, the niche for an insect-eating nocturnal animal, or a leaf-browsing daytime herbivore.

The kiwi is described as an “honorary mammal” by virtue of its adaptation to a niche normally occupied by a mammal. Kiwi seem to do in New Zealand what anteaters and insectivores to elsewhere. Indeed, they are very like to two-legged tenrecs (the fascinating Madagascan parallels to hedgehogs or giant shrews), or selenodonts (their counterparts in the Caribbean), or Australian echidnas, sniffing through the leaf litter at night in search of worms and grubs — prey which they detect by sense of smell[ii], a faculty essential to mammals but largely absent in birds. Even kiwi body temperatures are more like those of mammals and birds. Flighted birds normally have a high metabolic rate (because of their high levels of activity) and, related to this, maintain body temperature of 38°-43.8°C, compared with mammals that 36°-38°C. Kiwi temperatures range from 37° to 38.6°.

The kiwi evolved without the stoats and dogs that threaten their existence today. The 400 North Island brown kiwi killed by a single dog over a six-week period in 1987 might have had a chance of their wings had been more than rudiments hidden in the body feathers. The selective advantage of being able to fly even of a short distance to escape a predator is so strong that it takes to complete absence of ground predators for many generations before selection for other trays can be in the process leading to flightlessness.

Eat plants, grow large — and walk

Like all warm-blooded creatures, birds walk a metabolic tightrope, balancing the amount of energy they can get from their food with the amount they need in order to maintain body temperature, to grow, move and reproduce. A bird puts a lot of energy into building and maintaining the muscles, bones and feathers of its wings, and into fuel for its muscles when it flies. If a bird do not have to fly, the materials for construction and repair and the huge fuel-consumption bel — as much is 15-20 times the energy consumed when resting — could be channelled into growing rapidly and maturing earlier and into extra breeding effort. Take away ground predators and flightless species would have a competitive edge.

But other factors determine whether a species can make the transition. Obviously, even where there are no predatory mammals, not all birds are flightless. New Zealand has been called the land of flightless birds, but most of our species can fly. Animals develop as a coherent entity, not as separate parts. If deleting or slowing the growth of one part of the animal affects the development of other parts, the embryo will die. To become flightless, a bird must be able to develop properly in every way except for its wings. Yet the wings and their supporting breastbone are so important that in most birds they develop early and rapidly.

A few groups which live on the ground and whose young leave the nest early develop their legs faster than their wings. It is in these groups that flightlessness is most easily achieved. The best examples are rails and waterfowl, and New Zealand was particularly rich in flightless rails and ducks. Ten of our 13 rails were flightless, that only two of the 10, the weka and the South Island takahe, survive.

One way for groups with normal patterns of development to become flightless is simply to become too large or heavy for the wings to function. For energetic and aerodynamic reasons, flying birds can exist in only a limited range of sizes. Some very large birds have gone beyond those limits because their ecology and diet demand large size. When circumstances permit, such species, including the largest swans and bustards (very large running birds of the steppes of Asia and Africa), can avoid predation by their sheer size and aggressiveness, or by running or swimming away. They retained their full-size wings but find it hard to develop sufficient power to fly. In these species, the largest males fly very rarely, if at all.

In one species of Patagonian steamer duck, the young birds can fly, but as they grow, males gradually become too heavy for the wings to support them. Then they can escape only by ploughing across the water like miniature paddle-steamers — hence the name given by 19th-century sailors. They retain wings, but flight imposes to greater demand on the energy supply or the ability of their wing area to support the bird’s weight. In these ducks, gradual loss of flight occurs within the lifetime of an individual.

Another species of steamer duck has arrested the development of its wings to the point that it can never fly, even when young and light.

The evolution of large size is controlled to some extent by the absence or relaxation of predation pressure, but is mainly driven by diet, specifically a vegetarian diet. New Zealand, Hawaii and other oceanic islands are natural laboratories for observing the evolution of organisms isolated from the intense competition of life on continent. Not only did these islands lack mammalian predators, but herbivorous mammals as well, no large antelope or buffalo chomped on grasses, shrubs and tree leaves. No rodents chewed on fallen seeds and nuts, all that potential food was just sitting there unmolested. The opportunities for herbivores were wide open for any birds that could exploit them, and species from several groups did just that, nowhere more successfully than in New Zealand, where kakapo, rails, extinct geese and moa all feasted on plants.

The problem is, leaf material contains relatively little nourishment. Herbivores need to process a lot of plant matter to extract the energy and nutrients they need. Any means of lowering energy demands and improving digestion imparts great evolutionary advantages to the species able to do it. Discarding wings and breast muscles reduces energy requirements. Being big allows an animal to have a longer gut, which is more efficient at processing plant tissue because the material takes longer to pass through and so has more time to ferment and be digested.

Even so, much of what is eaten has no nutritional value and is voided. Our own takahe, a solid but not huge bird which dines on plant matter, produces a one-centimetre-diameter “cable” of fibrous green droppings eight metres long each day!

Larger animals also use less energy per gram of body weight than do small animals, both for staying warm and moving about, gram for gram, an elephant needs much less food than a mouse does, and an ostrich consumes relatively meagre rations compared with a hummingbird. The basic resting metabolic rate (effectively energy used while sleeping) for a 17 kg cassowary is 29 kcal/kg/day; for a 450-gram pigeon it is 235 kcal/kg/day, and for a tiny four-gram hummingbird it is a staggering 1410 kcal/kg/day! These rainbow-coloured midgets are in constant danger of burn-out, and must eat their own weight and food each day. The average human resting metabolic rate is 12.5 kcal/kg/day.

Given these figures, it might appear that life favours the large, but it’s not that simple. The daily food intake of one elephant would sustain a plague of mice, and this means that elephants will always be far less numerous than mice. When really hard times come, the elephant may starve, and so may most of the mice, but it is more likely that a few mice will still be able to eke out an existence, poised to reinfest the Serengeti when the conditions become favourable again. Fortunately for the safari industry, individual elephants don’t starve as quickly as mice do, and many survive long droughts.

For herbivorous birds then, there are advantages in being large: they save energy and they can recover more energy from their food. The price to be paid is a reduction in their ability to escape from predators and to move to new feeding grounds. Where there are no predators and the year-round supply of food within easy walking distance, these liabilities are easily ignored, and if the species can follow the evolutionary path to flightlessness, it will.

The dodo is perhaps the classic flightless vegetarian. An overgrown, docile pigeon, it did not long survive the arrival of hungry humans on Mauritius. Although the birds were not considered particularly good eating, sailors weren’t choosy, and the last birds were taken in 1662. We even have a written record, “When we held one by the leg he let out a cry; others came running forward to help the prisoner and were themselves caught.” So exited the dodo, and very nearly all traces of its existence. The last known dodo skin was consumed in a fire at Oxford early last century[iii], only the head of the leg being saved from the flames.

Unlike New Zealand’s moa, there are woefully few dodo bones in collections. Surprisingly, a large proportion are in New Zealand, brought here by the first experts on New Zealand’s own flightless birds. These bones are helping today’s scientists unravel the further mysteries of extinct birds in the Pacific.

The Hawaiian counterparts of our moa were the moa-nalo, flightless ducks as large as geese but which evolved from smaller ducks which flew. Many ducks feed on land when they can, as anyone who has seen the flocks feeding on acorns in Hagley Park will know. But the Hagley Park ducks have to beware of dogs and children; the ancestors of the moa-nalo did not. The ducks evolved into new species on the different islands, each cropping the forest-floor plants in a different way and rubbing shoulders with a range of flightless geese, of which the endangered Hawaiian goose is the last survivor — it can fly.

Large vegetarian waterfowl also shared New Zealand with the moa. Both main islands had their own flightless geese. Close relatives of the living Australian Cape Barren goose, the two species of Cnemiornis seem to have been confined to small areas of grass beside rivers and lakes. They are rare in fossil deposits but are known to have been eaten by the earliest New Zealanders.

It is worth noting that not all plant foods are low in energy. Flowers, seeds and fruit, for example, give high rewards, and those who eat them need not be large but must have mobility. Species that live on fallen fruit on the forest floor must move around to take advantage of different fruiting seasons. Seed-eaters must follow the seasonal supply of seeding grasses or depend on seeds shed at other seasons. The unpredictability of supply at any one point places a premium on mobility, and for birds that usually means flight. Only the cassowaries among the living flightless birds depend on fallen fruit. Their considerable size allows them to range far enough to eke out a living on the floor of the rainforest. Some of New Zealand’s forest moa probably depended on fruiting of the matai and miro trees.

Swimming to safety on virtual islands

Apart from islands, the other great refuges from predators are lakes and the sea. Here each bird can be its own island, well away from land-based enemies, and many of the most successful groups of flightless birds are aquatic.

Even flighted species, such as most ducks, geese and swans, use their water refuges for protection while they moulted their flight feathers (usually all at once) and are unable to fly for some weeks each year. Species that mould a few feathers at the time must cope with the reduced efficiency of the imperfect wing.

That water for many ducks as important for protection rather than food is clear when waterfowl are isolated on predator-free islands. In pre-human (and pre-rat) New Zealand, several ducks moved out of the streams, ponds and lagoons and onto the land. Until the rats came, Finsch’s duck, the brown teal and the blue duck wandered in the forests and shrubland, well away from running or standing water. Fortunately for the survival of the brown teal, it also lived in watery habitats, but Finsch’s duck was wedded to the land. Fossils show that its wings began to shrink after the last Ice Age, as taller vegetation once more covered the land. Having lost its powers of flight, it had no refuges when the Pacific rat arrived, and died out about 500 years ago[iv].

On outlying islands, other small ducks closely related to the brown teal responded to the absence of predators and an environment dominated by strong westerly winds (posing a threat to flying land birds of blowing them irrevocably out to sea) by becoming completely flightless. The Auckland Island teal lives along the shoreline and in the dense grasslands, feeding on insects and crustaceans and kelp piles and litter. Its vulnerability to predators was starkly revealed when cats and pigs were introduced to the main Auckland Island in the 19th century. Very soon, the little ducks were confined to small islets offshore which the predators have never reached.

The same fate befell the Campbell Island teal. It was thought to be extinct until a few were discovered living on steep and exposed Dent Island of Campbell Island’s west coast. There, the ducks move about under the thick grassland and scrub and have never seen a pond or stream. Under the thick vegetation, they are safe from their only predator, the brown skua. On Campbell Island itself, the introduction of Norway rats in the 19th century doomed the ducks to the early extinction.

Thousands of kilometres to the east of Campbell Island, the steamer ducks of Patagonia live on a continent, but with rocky stacks offshore for safe breeding, and they have been able to reduce their investment in flight as a result. In the remote Galápagos Islands, a flightless cormorant now lives now only on the western islands of the group with a cold waters around Barbara and Isabella islands are rich enough and fish to make it unnecessary to go far to find food, the cormorant used to be found in other islands, but the cats and pigs introduced in the past 200 years found them easy prey. The Galápagos species is now the only flightless cormorant. One hundred and fifty years ago the spectacled cormorant lived among the rocks and islets of Bering Sea, again protected by its harsh environment, and near rich fishing grounds. People found them an easy source of bait for their own fishing, and they were gone so quickly that only six skins remain.

“Flight” of the watery kind

The oceans have also provided protection to some groups of birds that traded flight in the thin air for flight in water. The world’s 13 species of penguins are the remaining members of a group that seems to have evolved along the southern shores of the Pacific, before the Drake Passage opened and Antarctica became an island continent.

Fossil evidence shows that New Zealand was a centre for the evolution and radiation of penguins. Some of the early forms were giants, fully two metres high, but all had anatomical adaptations that make them amongst the most efficient swimmers of all. Their bones are dense, unlike those of birds that fly in air, their bodies are covered in a layer of small feathers that streamline their form, and the wings have become flippers.

The wing bones themselves are flat and broad, and there are many more and shorter feathers than on a normal bird wing. By flapping these wings in a medium that is 900 times denser than air, some penguins can dive to more than 300 metres; others migrate thousands of kilometres.

Although the perfection of their adaptation to the aquatic environment has made them both awkward and vulnerable on land, their breeding grounds are usually on islands or stretches of coastline where there are few or no mammalian predators to take their toll. Now, some species are in trouble. Yellow-eyed penguins once nested as far north as Cook Strait in Nelson, but hungry humans trapped the northern populations to extinction. Dogs, cats and people disturb and maim the survivors on the Otago Peninsula. The rare crested penguin, now almost confined to, and called after, Fiordland, was also much more widely distributed before people arrived.

Birds of the southern latitudes, penguins reach their northern limits at the Galápagos Islands, where the tiny endangered Galápagos penguin just crosses the Equator.

Further north, other groups of seabirds have abandoned the air for life beneath the ocean wave, notably auks. Auks are among the most abundant of Northern seabirds. In colonies of many thousands, puffins, guillemots, razorbills and many others line the cliffs of the northern oceans in the brief summer. All these birds catch their prey by diving and swimming beneath the surface with half-folded wings. Their wings are compromises: short and narrow, they are still able to carry the birds in the air, but they also compact enough to work under water.

At least one species in the north Atlantic, the great auk, went for optimisation of the underwater effort, and the wings became too short, stiff and paddle-like for flight. This bird is one of the most famous flightless species of them all, not least because we know when the last two were killed (in 1844 on an island off the coast of Iceland).

Thousands of years ago, great auks habited the coasts of France and Spain. Until the Middle Ages there were still birds in Britain and Norway exploiting huge shoals of fish within swimming range of the cliffs and stacks where they bred. Other auks depended on food supplies further offshore, or move between feeding grounds with the seasons.

By the 18th century, the great auk had been decimated by persistent raids on its breeding grounds. Human exploitation of the bird for use as food and fish bait as well as for its feathers doomed the only flightless bird native to Europe in recent times. The great auk was the bird first called pingouin, and it was the ecological equivalent of the penguin in the North Atlantic. It even looked like a real penguin, and when first west-European mariners visited the Cape of Good Hope in Patagonia they thought they were seeing the same animal the name stuck, even after the great auk was harried to extinction.

On the other side of America, a few million years ago, another group of auks went wholeheartedly into flightlessness. All known species of mancalline auks, fossil birds from California, were flightless. In the warm seas, they took the niche now occupied by the smaller penguins in colder southern waters. In the North Pacific at the same time, the counterparts of the giant penguins were the huge plotopterids. Some of these birds were over two metres long. Distantly related to pelicans, in their day plotopterids ranged from Japan to California. As with the giant penguins, the demise of the huge plotopterids may have coincided with the rise of the seals as major inshore predators of fish and squid.

Penguins and auks can at least run, hop and toboggan on land and snow. Grebes are so well adapted to water that their short legs cannot support them on land, and they flop onto nests of floating vegetation to brood their eggs. The young are carried on the parents’ backs, and they never move on dry land at all.

It is not surprising that at least some grebes have given up flying as well. For these, the islands of flightlessness have been lakes, in Guatemala, Peru and Madagascar. For the giant pied-billed grebe of Lake Managua, being isolated on one lake has already proved fatal. Its habitat was altered by the introduction of exotic predatory fish (for human sport), and its close relative (and probable ancestor) the pied-billed grebe has colonised the lake as well. A combination of predation on the young and hybridisation pushed the giant pied-billed grebe into oblivion in the 1980s.

The fate of two other flightless grebes, each in their own evolutionary prison, hangs in the balance.

Flightless grebes may be almost predictable, but ibises seem unlikely candidates for flightlessness. Long-legged, long-billed and swift on the wing, ibises live in wetlands in the warmer parts of the world. They can fly strongly: glossy ibises regularly cross the Tasman to New Zealand.

Several millennia ago, some ibises reached Jamaica. Others made the epic trip to Hawaii. In both places their descendants stayed and moved into the forests. In Hawaii, they became so much ground birds that they have been compared with kiwi. The convergence in habits and structure was amazing. They lived on the forest floor, eating snails and other small invertebrates. When people arrived, they burnt the forests, and the tasty flightless birds could not survive the dual impact of habitat destruction and the cooking pot.

Why are there no flightless eagles?

Vegetarians and insect-eaters are surrounded by their food, and their food is slow-moving or stationary. Predatory birds have to hunt, and the food usually tries to get away. So it is almost impossible for a bird to be a predator of rapid-moving prey and not to fly. Even in the sea, penguins and auks “fly” through the water.

There are so few flightless predatory birds because it is just too expensive for relatively small, warm-blooded animals to walk and run after living prey, and dead animals are spread too thinly to search for on foot. So there are no flightless eagles, hawks or vultures. But there were flightless owls.

Before humans arrived on Cuba, the island was home to a huge owl which had such a flat breastbone that it almost certainly did not have big enough muscles to allow flight. Hunting by night, the bird could catch enough large rodents and insectivorous mammals — even young ground sloths — to make a living. It shared the island with an immense eagle, almost as large as the moa-eating Haast’s eagle in New Zealand.

The New Zealand adzebills were the exception that proves the “no flightless predator” rule. The strange distant relatives of the cranes[v] took to eating the one supply of animal protein in New Zealand that was not too fast to catch: reptiles. With their strong, strangely-shaped beaks they could hold struggling tuatara or big lizards and even dig them out of the ground. Some may even have caught and eaten the petrels the bred extensively on the mainland until rats, pigs, cats and humans arrived.

Like most specialised predators, adzebills were few in number; there’s not much room at the top of the food chain.

Insect-eaters, seed-eaters and omnivores are better candidates for flightlessness and than carnivores are. Their food is usually common and readily available, and escape without flying is easy for birds small enough to live in very dense vegetation or among rocks. Small, secretive and able to live in very dense vegetation, more rails have become flightless than any other bird group.

Another reason for the abundance of flightless rails is that, although rails are strong flyers, they fly buoyantly on their broad wings and are easily blown off course. This combination of attributes is probably led to their colonisation of most oceanic islands — even small specks of land far from any continent. But having flown there, it is the individuals that are not blown away that survive and breed. There is a paradox here: strong flyers most often reach places where it is easiest to become flightless.

Flightlessness and extinction

The apparent safety of predator-free islands has turned out to be an illusion for many flightless birds. People have colonised, or at least visit, nearly every speck of land on Earth. Visitation itself is not usually a problem, but people have a taking habit of taking camp followers with them. These include rats, cats, dogs and pigs. Alone or together, they have proved deadly to flightless birds.

Together with the destruction of habitat by burning and logging, and with hunting for food or sport, predatory mammals have been responsible for the extinction of most of the species of flightless bird alive in the past 10,000 years. New Zealand is a classic case. Of about 40 flightless species alive in New Zealand 1000 years ago, at least 29 of become extinct, 25 of those before Europeans arrived.

On islands everywhere, flightless birds have succumbed to the onslaught. It has been calculated that, in the South Pacific alone, about 4000 species of bird, perhaps more than 1000 of them flightless, have become extinct in the past 4000 years.

Their refuges were safe so long as their isolation remained and although some birds can become flightless in remarkably few generations, they can never regain the power of flight quickly enough once the cat is out of the bag. Losses have accumulated at least 100,000 times more rapidly in the last few millennia than they do through the normal background rate of extinction.

Ultimately, the ease and speed with which flightless birds disappear when their isolation is ended show the true value of flight. There are always environmental and physiological pressures toward saving energy. For most species, those pressures are balanced by the realities of survival and environments filled with hungry mouths and where food is hard to come by. Species that relax their guard, in the water or on islands, and choose to walk or swim, must always be at the mercy of the next invader.

Paradoxically, flightlessness is, or was, not rare, because there are many islands, and many species have reached them and stayed and changed. On those islands the pressure of competing with mammals for aeons were stripped away.

For most species, the way of life adopted by the ancestors could not be altered, and they still fly. For others — those that were able — release from a world of mammals meant that the very organs of success on the continents — wings — were now superfluous. There were new realms to conquer and nothing to drive them into the air. The wings that were so efficient in giving them a competitive edge on the mainland could be dispensed with. If man had not intruded, the number of flightless birds taking the place of stay-at-home mammals might well have equalled that of flying species. Such sheer abundance is a challenging thought for those who see birds as the epitome of freedom: the rulers of the sky.

Copyright Richard N. Holdaway

This article was first published in New Zealand geographic No. 37, January-March 1998

[i] Apart from a lark in the Canary Islands.

[ii] Evidence suggests that vibration rather than smell is the key to obtaining their food.

[iii] In the early 19th century.

[iv] A few may have survived into the 18th century.

[v] The group is thought now to be closer to rails.

Norfolk Island

Adult Tasman booby (Sula tasmani), Norfolk Island. The Tasman booby is one of the largest subtropical and tropical sulids and also one of the rarest. Most readily distinguished from others in the "masked booby" group by its brown eyes (yellow in the others), it breeds today only at Norfolk Island and in the Kermadec Islands. A colony at Lord Howe Island was exterminated early in the European period (immediately post-1788).

Juvenile Tasman booby (Sula tasmani) in flight over Point Howe, northwestern Norfolk Island. In brown boobies (Sula leucogaster) the brown extends farther down the breast: the brown is also smaller than the Tasman.

Juvenile Tasman booby (Sula tasmani) at its nest at Point Howe, northwestern Norfolk Island. Note the brown eye characteristic of this species. Tasman boobies are common as fossils in the dunes at Cemetery Bay on the southeastern coast, and the species was described from fossils from there and Lord Howe Island and recognised only recently to be be the bird still living at Norfolk Island.

Female greater frigatebird (Fregata minor) over the northwestern corner (Point Howe) of Norfolk Island. Up to 20 birds (almost all females) now visit the island.

Female greater frigatebird (Fregata minor) in flight over Norfolk Island. The species has become a frequent visitor in the past decade or two, but males are still very rare at the island.

A red-tailed tropicbird (Phaethon rubricauda) in display flight, Phillip Island, Norfolk Island group.

The wedge-tailed or Pacific shearwaters (Puffinus pacificus) of Norfolk Island and the Kermadecs are the largest of the species. Significant differences in size and breeding season among the different breeding populations suggest that several cryptic species are included under the name. On Norfolk Island, wedge-tailed shearwaters, known locally as "ghost birds" from their calls at the nesting burrows, are the commonest breeding seabird despite significant reductions in their numbers caused by cat predation.

A good fossil record shows that the black-winged petrel (Pterodroma nigripennis) was not part of the original seabird fauna of Norfolk Island. Its place was taken by a bird, now extinct, very like Pycroft's petrel (Pterodroma pycrofti) of northern New Zealand. In the second half of the 20th century, black-winged petrels started to colonise new areas across the Southwest Pacific, including Norfolk Island, where it is now a common breeder on Phillip Island and increasing on the main island when protected from cat predation.

A juvenile black-winged petrel (Pterodroma nigripennis) just out of the nest burrow (note the "collar" of down). It was found in the middle of Burnt Pine, the main settlement, and was released the following evening.

The white tern (Gygis alba) breeds in the remaining Norfolk Island forest and on isolated trees. Its "nest" is usually just an irregularity in an horizontal branch. In this seemingly precarious position, the adults successfully incubate the single egg and the chick is fed on small fish until it is able to fly. The egg and chick are vulnerable to introduced rats and cats, and now to predation by the self-introduced white-faced heron.

Norfolk Island has its own endemic subspecies of the widespread sacred kingfisher (Todirhamphus sanctus norfolcensis). It is still common and readily observed hunting from power lines and high branches.

The endemic Norfolk Island subspecies (Ninox novaeseelandiae undulata) of the widespread boobook or morepork owl went extinct as a pure population between 1978 and 2005. The last surviving female was joined by two males of the New Zealand subspecies, and the hybrid population, of which this bird is an example, is now widespread in the remaining forest.

The endemic Norfolk Island subspecies (Ninox novaeseelandiae undulata) of the widespread boobook or morepork owl went extinct as a pure population between 1978 and 2005. The last surviving female was joined by two males of the New Zealand subspecies. These are two birds of the hybrid population which is now widespread in the remaining forest.

Norfolk Island has its own fan-tailed flycatcher, the grey fantail (Rhipidura pelzelni). It is still common over most of the island, in forest and gardens.

The Norfolk Island whistler (Pachycephalaxanthoprocta) is one of the few endemic birds that has coped with predation by cats and rats introduced by Polynesians and Europeans over the past 1000 years.

The Norfolk Island whistler (Pachycephala xanthopygia) is closely related to the golden whistler but differs in that the male is not brightly coloured. The reason for the species adjective - the yellow around the vent - is obvious here.

The Norfolk Island environment

Geology

Norfolk (3455 ha), Nepean (10 ha), and Philip (190 ha) islands are the much-restricted post-glacial sub-aerial sections of a much larger (320,000 ha, c 85 times the present extent) flat-topped edifice on the Norfolk Island Ridge (Jones & McDougall 1973). The area of sub aerial exposure has repeatedly expanded and contracted in concert with sea level changes associated with the Quaternary glacial events, and presumably the terrestrial biota will have gone through repeated cycles of restrictions in area (“bottleneck events”, with each amelioration) and expansions (with enhanced chance of over-water colonisation, at each glacial low sea level stand). Jones & McDougall (1973) emphasise the apparent tectonic stability of the Norfolk Island Ridge, on which Norfolk Island and its outliers is situated), with no evidence for significant uplift or subsidence, so that sea level changes associated with ice volume fluctuations have been the only determinants of land area apart from the relatively much smaller effects of erosion of the presently emergent islands.

The rocks are primarily oceanic basalts of Pliocene to early Pleistocene age (Jones & McDougall 1973); some late Pleistocene superficial calcarenites are exposed at the southern end of Norfolk Island and form most of Nepean Island. The youngest basalts on Norfolk Island are about 2.3 million years (ma) old (Jones & McDougall 1973; McDougall 1973), which is midway between the ages of Oahu (2.6-3.0 ma) and Molokai (1.8 – 2.0 ma) islands in the Hawaiian chain (Vitousek 2004).

Norfolk Island soils and evidence for marine nutrient subsidies

The soils of Norfolk Island are developed for the most part at the surface of a weathered layer of basalt that varies between 12 and 40 metres in depth (Jones & McDougall 1973). They are at the older end of the age series (0.3 ka to 4.1 ma) discussed by Vitousek (2004), and hence likely to exhibit similar characteristics in terms of available nutrients, i.e. very low levels of phosphorus contributed by the parent rock. On Norfolk Island itself, the weathering blanket and deep soils preclude exposure of the basement rock except for the coastal cliffs and tiny areas near the summit of the island. The soil profile data presented by Stephens & Hutton (1954) show lower levels of, for example, phosphorus (P), with depth. This suggests that there is little contribution from the parent rock. The low relief of much of the island means that erosion cannot supply new weathering surfaces.

Although their discussion of the development of the Norfolk Island soils was based on a greatly expanded time frame than the one now accepted for the island, Hutton & Stephens (1956) and Stephens & Hutton (1954) provide a valuable map and baseline data for the nutrient levels in the different soil systems as they existed over 50 years ago. Most of the island is covered by their Rooty Hill clay, which had total P content of 800 mg/kg, which is well below the values measured for Norfolk Island basalts of 1620-1750 mg/kg Jones & McDougall 1973).

Stephens & Hutton (1954) identified the most fertile soil (their “Selwyn clay”) as being associated with the presence of burrow-nesting seabirds (their “sooty petrels”; actually the wedge-tailed or Pacific shearwater, Puffinus pacificus) along the cliff tops on the western side of Norfolk Island. The Selwyn clay had up to 4300 mg/kg total P. Two samples of Steel’s Point clay (from near Steel Point and from the plateau at the north-western corner of the island) reached 3000-3200 mg/kg total P. These may well have been from areas that also had nesting shearwaters in the early 1950s, as they are present along the cliff edges in that area today. Hutton & Stephens (1956) note that the burrowing by the shearwaters “may be responsible for the remarkably uniform and dark colour of the whole profile of this soil”. This would result from the ploughing in of organic matter by the birds (Hawke 2005).

Re-interpretation of the calcareous rocks at Kingston and on Nepean Island (Jones & McDougall 1973) has shown that they are the product of sub aerial dune formation during recent low sea stands and are not part of a former makatea. The best explanation of the source of the enrichment in Ca2+ and Mg2+ would seem to be the former presence of large breeding populations of burrow-nesting and other seabirds across much of the island (Holdaway & Anderson 2001). It is likely, too, that the differences between the “mature” Middlegate clay, with its high rutile and haematite levels, and the “more juvenile” Palm Glen clay may be the result of contributions by burrow-nesting seabirds on the slopes of Mt Pitt and Mt Bates, where there were large populations until the 1790s (Hoare 1987), and probably significant numbers until the mid-1940s (Holdaway et al. unpubl data), and a lack of such effects in the more subdued terrain of the southern tableland. The big unknown is the P levels in the “skeletal” soils on the main ridges of Mts Pitt and Bates, where historically seabirds were known to nest, and where stable isotopic data (see below) suggest that marine nutrients were significant in the ecosystem (Holdaway et al. unpubl. data). Anecdotal information supports the general impression of the trends in fertility.

The highest levels of P in surface samples examined by Stephens & Hutton (1954) were associated with the lowest N:P ratios, showing that the N level, reflecting the relatively constant levels of N in Norfolk Island soils at that time, and the concentration of P in coastal soils, associated with residual breeding colonies of shearwaters.

The recent history of nutrient sources for vegetation on Norfolk Island is being examined in a study using stable isotopes in the wood of large specimens of Norfolk Island pine (Araucaria heterophylla), obtained from stumps (2) or the severed butt (1) of trees on the northeastern, southwestern, and western sides of the Mt Pitt massif. This work is in progress.

Work in New Zealand (Harding et al. 2004; Harrow et al. 2006; Hawke & Holdaway 2005, 2009; Holdaway et al. 2007) has shown that the nitrogen applied by seabirds is more labile and lost more rapidly than the phosphorus, but that the phosphorus also can have a relatively short life in soils that have rapid drainage. Plant growth and hence productivity is limited more by low levels of phosphorus than of nitrogen, and phosphorus enhances growth and production. That is so with natural, bird-derived applications of phosphorus as well as for artificial applications of the element. As noted below (Summary Point 3), the present value of the phosphorus emplaced annually in the Mt Pitt area by Providence petrels at the start of the First Settlement was likely to be up to A$250,000. The effects of such a natural enhancement of the terrestrial nutrient flux must have been profound.

Summary

1. The terrestrial biota of Norfolk Island developed in the presence of significant transfers of nutrients from the oceanic to the terrestrial food chain by large populations of burrow-nesting and arboreal sea bird

2. The decimation of the sea bird populations in general, and their complete removal from the high parts of Norfolk Island, has all-but eliminated the flow of nutrients to the terrestrial ecosystems

3. The seabirds supplied plant-available nutrients at a rate that cannot be maintained from other natural sources; in particular, the intensely weathered basalt rocks of Norfolk Island have never been able to match the supply brought in by seabirds and cannot do so now, given the age of the rock

4. The invertebrate and remnant vertebrate faunas of Norfolk Island are being affected and damaged by the changes in the vegetation, and by low levels of nutrients in the soils and vegetation. The faunas will decline further and the vegetation will probably change to a completely different composition if the supply of seabird-vectored nutrients is not restored

5. Application of artificial fertilisers will not be a long-term solution, because it cannot mimic the mode of application or ploughing of soils to depths of > 50 cm

6. Hence, burrow-nesting petrels will be essential to the survival of the Norfolk Island terrestrial ecosystems

7. The species of burrow-nesting petrels cannot survive in the presence of mammalian predators such as cats and the species of rats at present on Norfolk Island

8. To restore and maintain the terrestrial ecosystems of Norfolk Island, the mammalian predators must be removed – not controlled – and kept off the island

9. Technologies exist that allow the restoration of viable breeding populations of burrow-nesting petrels to their former breeding ranges

10. The alternative to rodent eradication and restoration of seabird population is the continued deterioration and eventual depauperation of the fauna and flora of Norfolk Island, even with increased levels of intervention, with a reduction in tourism and tourist income and ongoing costs of amelioration

Extra notes for Norfolk Island section summary

References

Bancroft, W.J.; Garkaklis, M.J.; Roberts, J.D. 2005a. Burrow building in seabird colonies: a soil-forming process in island ecosystems. Pedobiologia 49: 149-165.

Bell, M.; Bell, B.D.; Bell, E.A. 2004. Translocation of fluttering shearwater (Puffinus gavia) chicks to create a new colony. Notornis 52(1): 11-15.

Croll, D.A.; Maron, J.L.; Estes, J.A.; Danner, E.M.; Byrd, G.V. 2005. Introduced predators transform Subarctic islands from grassland to tundra. Science 307: 1959-1961.

Fairbanks, R.G. 1989. A 17,000 year glacio-eustatic sea level record: influence of glacial melting rates on the Younger Dryas event and deep ocean circulation. Nature 342: 637-642.

Harding, J.S.; Hawke, D.J.; Holdaway, R.N.; Winterbourn, M.J. 2004. Incorporation into stream food webs of marine-derived nutrients from petrel breeding colonies. Freshwater biology 49: 576-586.

Harrow, G.; Hawke, D.J.; Holdaway, R.N. 2006. Surface soil chemistry at an alpine procellariid breeding colony in New Zealand, and comparison with a lowland site. New Zealand journal of zoology 33: 165-174.

Hawke, D.J.; Holdaway, R.N. 2009. Nutrient sources for forest birds captured within an undisturbed petrel colony, and management implications. Emu 109: 163-169.

Harding, J. S.; Hawke, D.J.; Holdaway, R.N.; Winterbourn, M.J. 2004. Incorporation into stream food webs of marine-derived nutrients from petrel breeding colonies. Freshwater biology 49: 576-586.

Hawke, D.J. 2005. Soil P in a forested seabird colony: inventories, parent material contributions, and N:P stoichiometry. Australian journal of soil research 43: 957-962

Hawke, D.J. ; Holdaway, R.N. 2005. Avian assimilation and dispersal of carbon and nitrogen brought ashore by breeding Westland petrels Procellaria westlandica: a stable isotope study. Journal ofzoology, London 266: 419-426.

Hoare, M. 1987. Norfolk Island: an outline of its history 1774-1987. St Lucia, University of Queensland Press.

Holdaway, R.N. 1999. Introduced predators and avifaunal extinction in New Zealand. pp. 189-238 In: MacPhee, R.D.E. (ed.) Extinctions in near time: causes, contexts, and consequences. New York: Kluwer Academic/Plenum Press.

Holdaway, R.N.; Anderson, A.J. 2001. Avifauna from the Emily Bay settlement site, Norfolk Island: A preliminary account. Records of the Australian Museum supplement 27: 85-100.

Holdaway, R.N.; Hawke, D.J.; Hyatt, O.; Wood, G.C. 2007. Stable isotopic (δ15N, δ 13C) analysis of wood in trees growing in past and present colonies of burrow-nesting seabirds in New Zealand. I. δ15N in two species of conifer (Podocarpaceae) from a mainland colony of Westland petrels (Procellaria westlandica), Punakaiki, South Island. Journal of the Royal Society of New Zealand 37: 75-84.

Hutton, J.T.; Stephens, C.G. 1956. The palaeopedology of Norfolk Island. Journal of soil science 7: 255-267.

Jones, J.G.; McDougall, I. 1973. Geological history of Norfolk and Philip Islands, Southwest Pacific Ocean. Journal of the Geological Society of Australia 20: 239-254.

Main, W. de L.; McKnight, D.G. 1981. Norfolk Island bathymetry, 1:200,000. Island Chart Series. Wellington, New Zealand Oceanographic Institute.

McDougall, I.; Jones, J.G. 1973. [Petrology of the Norfolk Island basalts]. Journal of the Geological Society of Australia 20.

Miskelly, C.; Timlin, G.; Cotter, R. 2004. Common diving petrels (Pelecanoides urinatrix) recolonise Mana Island. Notornis 51(4): 245-246.

Olson, S.L.; Wingate, D.B.; Hearty, P.J.; Grady, F.V. 2005. Prodromus of vertebrate paleontology and geochronology of Bermuda. Monografies de la Societat d’Història Natural de les Balears 12: 219-232.

Orth, K. 1980. A study of the geology, palaeontology, and biogeography of Nepean Island and the Kingston area of Norfolk Island, Southwest Pacific. Geology 301 project, Earth Sciences Department, Monash University, Clayton, Victoria.

Stephens, C.G.; Hutton, J.T. 1954. A soil and land-use study of the Australian territory of Norfolk Island, South Pacific Ocean. Soils and land use series no. 12. C.S.I.R.O., Melbourne.

Taylor, G.A. 2000. Action plan for seabird conservation in New Zealand. Part A: Threatened seabirds. Threatened species occasional publication 16; Part B: Non-Threatened seabirds. Threatened species occasional publication 17. Wellington: New Zealand Department of Conservation.

Turner, J.S.; Smithers, C.N.; Hooglund, R.D. 1968. The conservation of Norfolk Island. Special publication no. 1, Australian Conservation Foundation, Melbourne.

Vitousek, P.M. 2004. Nutrient cycling and limitation: Hawai‘i as a model system. Princeton University Press, Oxford and Princeton.

Veevers, J.J. 1976. The modern coastal sedimentary rock complex of Norfolk and Nepean Islands. In: Abell, R.S. A groundwater investigation on Norfolk Island. B.M.R. geology and geophysics record 1976/62.

Walker, J.; Thompson, C.H.; Jehne, W. 1983. Soil weathering stage, vegetation succession, and canopy dieback. Pacific science 37: 471-481.

Walker, L.R.; Aplet, G.H. 1994. Growth and fertilization responses of Hawaiian tree ferns. Biotropica 26: 378-383.

More information on Norfolk Island birds can be found in:

Norfolk Island... the Birds

by Margaret L. Christian. 2005. Green Eyes Publications, Norfolk Island. ISBN 0-9758212-0-2

Available from Margaret Christian, P.O. Box 999, Norfolk Island, South West Pacific

A comparative atlas of bird distribution in the Norfolk Island Group South West Pacific Ocean 1978-2005

by Margaret L. Christian, Richard N. Holdaway, John L. Smith, Peter D. Coyne. 2012.

The Flora & Fauna Society of Norfolk Island, South West Pacific. ISBN 0-9758212-1-0

Available from Margaret Christian, P.O. Box 999, Norfolk Island, South West Pacific

A review of Norfolk Island birds: past and present

by R. Schodde, P. Fullagar, N. Hermes. 1983. Australian National Parks and Wildlife Service Special Publication 8.

Australian National Parks and Wildlife Service, Commonwealth of Australia.

Images of Norfolk Island birds

Female greater frigatebird (Fregata minor) over the northwestern corner (Point Howe) of Norfolk Island. Up to 20 birds (almost all females) now visit the island.

Female greater frigatebird (Fregata minor) in flight over Norfolk Island. The species has become a frequent visitor in the past decade or two, but males are still very rare at the island.

A red-tailed tropicbird (Phaethon rubricauda) in display flight, Phillip Island, Norfolk Island group.

The endemic Norfolk Island subspecies (Ninox novaeseelandiae undulata) of the widespread boobook or morepork owl went extinct as a pure population between 1978 and 2005. The last surviving female was joined by two males of the New Zealand subspecies. These are two birds of the hybrid population which is now widespread in the remaining forest.

Norfolk Island has its own fan-tailed flycatcher, the grey fantail (Rhipidura pelzelni). It is still common over most of the island, in forest and gardens.

TURNAGRA PRESS

Contact Turnagra Press at: turnagra@gmail.com

Images from the 2008 Excavation at Pyramid Valley, featured in Pyramid Valley and Beyond

Palaecol Research also publicises natural history books for the South West Pacific...

Published in 2012 by the Flora & Fauna Society of Norfolk Island.Available from Margaret Christian, P.O. Box 999, Norfolk Island, South West Pacific. — Published in 2012 by the Flora & Fauna Society of Norfolk Island.
Available from Margaret Christian, P.O. Box 999, Norfolk Island, South West Pacific.

Sample maps for a Norfolk Island endemic species

Norfolk Island... The Birds

is also available from Margaret Christian, P.O. Box 999, Norfolk Island, South West Pacific

Consultancy

Page under development...

Does your institution protect your data?

Do your colleagues play by the rules?

In these days of competitive science, data are your most valuable assets.

Their security must be assured. But does your institution really protect your IP?

And do your peers respect the norms of scientific publication?

Our recent experience suggests that bad things can happen, as we describe below.

In a nutshell, a competing group had had a manuscript declined because it lacked data supporting their assertions. To fill the gap, they then accessed data from the embargoed MSc thesis of a graduate student in our group and used those data as the basis for a revised manuscript. The revised version was then accepted and published, denying our group priority in a novel and important revision of current understanding of an extinct ecosystem.

The data were obtained from the thesis Abstract that had, with the knowledge of the student or their supervisors, been made available on the university's library's web page. The thesis was, however, clearly labelled as being EMBARGOED.

Copies of correspondence are provided below in support of the chronology of events, statements made, and conclusions drawn here.

UNAUTHORISED PUBLICATION OF DATA FROM AN 'EMBARGOED' MSc thesis

In July 2014, an MSc thesis reporting stable isotopic analyses of vegetation from present and past New Zealand forests was lodged in the Library of the University of Canterbury, Christchurch, New Zealand. The work was done in the University's Department of Geological Sciences and School of Biological Sciences. The thesis contained sensitive information pertaining to a wider study funded originally by the Marsden Fund of the Royal Society of New Zealand, so the the student and her supervisors placed the thesis under 'Embargo' for two years, until 31 July 2016, to permit unimpeded publication of the results.

Without reference to the student or her supervisors, however, the thesis Abstract was displayed on-line on the University of Canterbury Library website. A drop-down form was supplied whereby a researcher could request an electronic copy of the thesis, even though the thesis was clearly marked as 'EMBARGOED' until 31 July 2016.

Despite the thesis still under embargo, a paper published on 16 May 2016in the journal Quaternary Science Reviews included nine citations of the thesis (not of the Abstract). The paper, although headed as an Opinion, is set out in the form of a research paper.

Dietary interpretations for extinct megafauna using coprolites, intestinal contents and stable isotopes: Complimentary or contradictory?

Quaternary Science Reviews 142: 173-175

Nicolas J. Rawlence (Allan Wilson Centre, Department of Zoology, University of Otago, Dunedin, New Zealand

& Canterbury Museum, Christchurch, New Zealand)

Jamie R. Wood (Long Term Ecology Laboratory, Landcare Research, Lincoln, New Zealand)

Herve Bocherens (Fachbereich Geowissenschaften, Forschungsbereich Paläobiologie, Universität Tübingen, Tübingen, Germany & Senckenberg Center for Human Evolution and Palaeoenvironment (HEP), Universität Tübingen, Tübingen, Germany)

Karyne M. Rogers (National Isotope Centre, GNS Science, Lower Hutt, New Zealand)

The paper records its receipt on 10 April 2016 and that a revised version was received on 10 May. The paper was accepted on 11 May and published on-line on 16 May.

However, the paper does not record the fact that an initial version had been submitted to the journal on 12 February 2016. This initial version was declined with an option to re-submit, in accordance with the reviewers' recommendations. The reason for rejection was that the conclusions were not supported by data.

University of Canterbury records show that on 7 March 2016, someone from the University of Otago accessed the Abstract and attempted to download the thesis but received only a request form. The records then show that the lead author, Dr Rawlence, then applied via the form for release of the (clearly still embargoed) thesis. The thesis was never supplied, but the Abstract, as normal for a thesis, included the critical data on which the conclusions were based, was available on the Library web site.

The version submitted on 10 April (published on 16 May) included nine citations of the thesis (despite the journal having a citation category for Abstracts, which was the only part of the thesis accessed), and the interpretations are based on data from the thesis. Although the University of Otago has stated in correspondence that the thesis was cited only in generality, the paper includes data that are clearly from the thesis abstract, which much be construed as citation in detail, not in generality.

To summarise, the first version of the paper did not include data from the MSc thesis and was rejected because of lack of supporting data. The new version included data directly acquired from the embargoed MSc thesis and was then accepted and published.

There is nothing in the paper to reveal that the MSc research on whose results the paper's publication depended was done by a different laboratory, unrepresented in the authorship. The critical measurements cited in the paper were made in the Department of Geological Sciences, University of Canterbury, on samples collected in 2008 under a Marsden Fund contract with Palaecol Research Ltd.

The credit accruing to authors on publication is, of course, balanced by the collective responsibility of the authors for the content of the paper.

The student's supervisors brought the matter to the attention of Quaternary Science Reviews and the senior management team of the University of Canterbury.

On being apprised of the situation, the journal initiated an investigation and provided information on the earlier history of the paper. One of the reviewers had published several times with one of the paper's authors, but of course it was the reviewer's responsibility to declare such a potential conflict of interest. That they provided a review confirms that no such declaration was made. The journal, after the initial review, indicated that no further action would be taken until a report was received from the University of Canterbury. To our knowledge, no such report has been provided.

On being apprised of the publication of embargoed data, the University of Canterbury requested, and was provided, with full details - as they were known at the time - of the matter and with a course of action that should follow to rectify the situation so far as possible. The supervisors were assured that the University of Canterbury took the matter very seriously.

The University of Canterbury Vice Chancellor (Research) wrote to their opposite number at the University of Otago. The response presented the view that the citations were only in general and did not involve detail. The citation of numerical data in the paper refutes this position. The response also indicated that Dr Rawlence would apologise to the student. To date, the student has not received any communication, apology or otherwise, from Dr Rawlence or from any other of the authors. A letter addressed to the student has, however, reached one of the supervisors. This presents the view that Dr Rawlence was unaware that the thesis data were embargoed and suggested that its citation was a mistake for which he is apologetic. It is hard to reconcile these statements with the contents of the paper (where the data are critical to the argument) or access to the thesis Abstract, which was clearly marked as being Embargoed. Dr Rawlence states that the paper could have been published without reference to the thesis. This is in conflict with the actual history of the paper, which was rejected for absence of data in support of their conclusions, and accepted only when the MS was revised, based on the thesis data. The letter has not been passed on to the student as it is the authors' responsibility to make good their promise of an apology, not to expect others to do it for them.

So far, the University has taken the issue up with the University of Otago only, and none of the other authors or their institutions.

Our views of what would constitute a correct response by the authors were set out clearly in the first communication to the University of Canterbury and the same views have been reiterated at meetings and in correspondence.

To date, despite assurances that the matter is important, nothing further has been done except for the belated removal of access to Abstracts and other details of embargoed theses from the University's Library web site.

CORRESPONDENCE WITH

QUATERNARY SCIENCE REVIEWS

THE UNIVERSITY OF CANTERBURY

UC 1

UC 2

UC 3

UC 4

UC 5

UC 6

UC 7

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UC 10

RESPONSE BY UNIVERSITY OF OTAGO

RAWLENCE LETTER

In our view, the responses by Dr Rawlence and his co-authors, and by the University of Canterbury fall far short of what is appropriate and necessary. The use of the data cannot have been a mistake, as the nine citations in second version of the manuscript show. The assertion that the manuscript could have been published without the thesis data is obviously incorrect as the first version of the manuscript, which lacked those very data, was declined. The statement that the authors did not know that the thesis was embargoed cannot be reconciled with the "Embargoed" notice accompanying the thesis on the Library web site and the application for release of an embargoed thesis. Hence the view that the thesis abstract material was freely available cannot be sustained.

As noted in the first communication to the University of Canterbury, we see two problems: first, that the Abstract of an embargoed thesis was put on on-line at all; and second, that the authors of the paper in Quaternary Science Reviews used data from that thesis to support their own contentions, without reference to other owners of the intellectual property. It was clear from the nature of the material analysed and the isotopic measurements made, that others were involved in ownership of the data: for example, no MSc student runs their own stable isotopic laboratory. Two stable isotope specialists on the authorship might give the impression that the measurements were made in one or other of their laboratories, and some might even suppose a connection between the student and those laboratories: none such existed.

The appropriation of data violates the fundamental principles of scientific publication.

We have set out repeatedly what we believe to be appropriate actions by the individuals and institutions involved. None of those actions has yet been taken.