Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 5374 No. 2: 16 Nov. 2023
Type: Article
Published: 2023-11-16
DOI: 10.11646/zootaxa.5374.2.7
Page range: 263-294
Abstract views: 698
PDF downloaded: 24

On the sand and among the crowds: a new species of Woodworthia gecko (Reptilia: Diplodactylidae) from Auckland, Aotearoa/ New Zealand

Bioresearches (Babbage Consultants Ltd.); Level 4; 68 Beach Road; Auckland 1010; New Zealand
School of Environmental and Animal Sciences; Te Pukenga - New Zealand Institute of Skills and Technology; Private Bag 92025; Auckland 1142; New Zealand
Independent herpetologist; Onehunga; Auckland
151 Woodburn Drive; Takapu Valley; Wellington 5028
Reptilia Taxonomy Te Oneone Rangatira/ Muriwai Beach duneland Woodhill Oaia Island conservation Woodworthia korowai sp. nov.

Abstract

Woodworthia is a diverse genus of diplodactylid geckos found in Aotearoa/ New Zealand, with 17 likely species. Despite this diversity, only two species have been formally described: Woodworthia maculata (Gray, 1845) and W. chrysosiretica (Robb, 1980). In this paper, we use an integrated taxonomic approach to describe a new species of Woodworthia gecko, Woodworthia korowai sp. nov., found along the western coastline of the Auckland Region, New Zealand. Although this species occurs in duneland habitat behind a popular beach near New Zealand’s most populated city, it was only recognised as a distinct taxon in 2016. We describe W. korowai sp. nov. based on a suite of morphological character states and substantial genetic divergence, based on the mitochondrial NADH dehydrogenase subunit 2 (ND2) gene, that distinguish it from W. maculata sensu stricto and all other known species of Woodworthia. Phylogenetic reconstruction and molecular dating place it sister to the W. maculata group, with an estimated time of divergence in the mid to late Pliocene. This gecko is one of the most geographically restricted of all Woodworthia geckos, occupying an area of less than 500 km2 within the Auckland Region. Its narrow range and coastal association make it susceptible to environmental and genetic stochasticity. Furthermore, the popularity and recreational usage of the dune system threaten its habitat. Therefore, we hope that this description will bring attention to the value of coastal environments and the unique and sensitive duneland of Te Korowai-o-Te-Tonga/ South Kaipara Peninsula and Te Oneone Rangatira/ Muriwai Beach in particular and encourage conservation efforts to protect this newly described species and its habitat.

 

References

  1. Barr, B.P., Chapple, D.G., Hitchmough, R.A., Patterson, G.B. & Board, N.T. (2021) A new species of Oligosoma (Squamata: Scincidae) from the northern North Island, New Zealand. Zootaxa, 5047 (4), 401–415. https://doi.org/10.11646/zootaxa.5047.4.1
  2. Bell, T. (2014) Standardized common names for New Zealand reptiles. BioGecko, 2, 8–11.
  3. Böhme, W. (1984) Erstfund eines fossilen Kugelfingergeckos (Sauria: Gekkonidae: Sphaerodactylinae) aus Dominikanischem Bernstein (Oligozän von Hispaniola, Antillen). Salamandra, 20 (4), 212–220.
  4. Brown, M.B. & Forsythe, A.B. (1974) The small sample behavior of some statistics which test the equality of several means. Technometrics, 16 (1), 129–132. https://doi.org/10.1080/00401706.1974.10489158
  5. Cameron, E.K. & Taylor, G.A. (1989) Oaia Island, South Muriwai. Auckland Botanical Society Journal, 44 (1), 11–12.
  6. Cangelosi, R. & Goriely, A. (2007) Component retention in principal component analysis with application to cDNA microarray data. Biology Direct, 2 (1), 1–21. https://doi.org/10.1186/1745-6150-2-2
  7. Chan, K.O. & Grismer, L.L. (2021) A standardized and statistically defensible framework for quantitative morphological analyses in taxonomic studies. Zootaxa, 5023 (2), 293–300. https://doi.org/10.11646/zootaxa.5023.2.9
  8. Chappell, P.R. (2013) The climate and weather of Auckland. 2nd Edition. NIWA Science and Technology Series Number 60. Niwa, Auckland, 40 pp.
  9. Cope, E.D. (1869) Observations on reptiles of the Old World. Art. II. Proceedings of the Academy of Natural Sciences of Philadelphia, 20 (5), 316–323. [1868] https://doi.org/10.1038/scientificamerican05151869-316
  10. Cree, A. & Hare K.M. (2016) Reproduction and life history of New Zealand lizards. In: Chapple, D. (Ed.), New Zealand Lizards. Springer International Publishing, Cham, pp. 169–206. https://doi.org/10.1007/978-3-319-41674-8_7
  11. Daza, J.D. & Bauer, A.M. (2012) A new amber-embedded sphaerodactyl gecko from Hispaniola, with comments on morphological synapomorphies of the Sphaerodactylidae. Breviora, 529 (1), 1–28. https://doi.org/10.3099/529.1
  12. Daza, J.D., Bauer, A.M. & Snively, E.D. (2014) On the fossil record of the Gekkota. The Anatomical Record, 297 (3), 433–462. https://doi.org/10.1002/ar.22772
  13. de Queiroz, K. (1998) The general lineage concept of species: species criteria and the process of speciation. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless Forms: Species and Speciation. Oxford University Press, Oxford, pp. 57–75.
  14. de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56 (6), 879–886. https://doi.org/10.1080/10635150701701083
  15. De Vis, C.W. (1888) A contribution to the herpetology of Queensland. Proceedings of the Linnean Society of. New South Wales, Series 2, 2, 811–826. https://doi.org/10.5962/bhl.part.29213
  16. Drummond, A.J. & Rambaut, A. (2007) BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology, 7 (1), 1–8. https://doi.org/10.1186/1471-2148-7-214
  17. Drummond, A.J., Suchard, M.A., Xie, D. & Rambaut, A. (2012) Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecular Biology and Evolution, 29 (8), 1969–1973. https://doi.org/10.1093/molbev/mss075
  18. Duméril, A.M.C. & Bibron, G. (1836) Erpetologie Générale ou: Histoire Naturelle Complete des Reptiles. Vol. 3. Libr. Encyclopédique Roret, Paris, 528 pp.
  19. Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32 (5), 1792–1797. https://doi.org/10.1093/nar/gkh340
  20. Fitzinger, L. (1843) Systema Reptilium. Fasciculus primus: Amblyglossae. Braumüller und Seidel, Vindobonae [Vienna], 106 pp. https://doi.org/10.5962/bhl.title.4694
  21. Fleming, C.A. (1975) The geological history of New Zealand and its biota. In: Kuschel, G. (Ed.), Biogeography and Ecology in New Zealand. Dr W. Junk., The Hague, pp. 1–86. https://doi.org/10.1007/978-94-010-1941-5_1
  22. Fleming, C.A. (1979) The Geological History of New Zealand and its Life. Auckland University Press, Auckland, 141 pp.
  23. Frost, D.R. & Hillis, D.M. (1990) Species in concept and practice: herpetological applications. Herpetologica, 46 (1), 87–104.
  24. Gamble, T., Bauer, A.M., Greenbaum, E. & Jackman, T.R. (2008) Evidence for Gondwanan vicariance in an ancient clade of gecko lizards. Journal of Biogeography, 35 (1), 88–104. https://doi.org/10.1111/j.1365-2699.2007.01770.x
  25. Garman, S. (1901) Some reptiles and batrachians from Australasia. Bulletin of the Museum of Comparative Zoology at Harvard College, 39, 1–14.
  26. Goldberg, J., Trewick, S.A. & Paterson, A.M. (2008) Evolution of New Zealand’s terrestrial fauna: a review of molecular evidence. Philosophical Transactions of the Royal Society of London B Biological Sciences, 363 (1508), 3319–3334. https://doi.org/10.1098/rstb.2008.0114
  27. Gray, J.E. (1842) Description of two hitherto unrecorded species of reptiles from New Zealand; presented to the British Museum by Dr. Dieffenbach. In: Gray, J.E. (Ed.), The Zoological Miscellany. Part II. Treuttel, Würtz & Co, London, pp. 57–72.
  28. Gray, J.E. (1845) Catalogue of the Specimens of Lizards in the Collection of the British Museum. Trustees of die British Museum and Edward Newman, London, xxvii + 289 pp.
  29. Grimaldi, D. (1995) The age of Dominican amber. In: Anderson, K.B. & Crelling, J.C. (Eds.), Amber, Resinite, and Fossil Resins. American Chemical Society, Washington, D.C., pp. 203–217. https://doi.org/10.1021/bk-1995-0617.ch011
  30. Grismer, L.L., Perry, L.W., Grismer, M.S., Quah, E.S., Thura, M.K., Oaks, J.R., Lin, A. & Lim, D.Y. (2020) Integrative taxonomic and geographic variation analyses in Cyrtodactylus aequalis (Squamata: Gekkonidae) from southern Myanmar (Burma): one species, two different stories. Israel Journal of Ecology and Evolution, 66 (3–4), 151–179. https://doi.org/10.1163/22244662-20191082
  31. Hammer, Ø., Harper, D.A.T. & Ryan, P.D. (2001) PAST: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontologia Electronica, 4 (1), 9.
  32. Han, D., Zhou, K. & Bauer, A.M. (2004) Phylogenetic relationships among gekkotan lizards inferred from C-mos nuclear DNA sequences and a new classification of the Gekkota. Biological Journal of the Linnean Society of London, 83 (3), 353–368. https://doi.org/10.1111/j.1095-8312.2004.00393.x
  33. Hare, K.M., Daugherty, C.H. & Chapple, D.G. (2008) Comparative phylogeography of three skink species (Oligosoma moco, O. smithi, O. suteri; Reptilia: Scincidae) in northeastern New Zealand. Molecular Phylogenetics and Evolution, 46 (1), 303–315. https://doi.org/10.1016/j.ympev.2007.08.012
  34. Hare, K.M., Chapple, D.G., Towns, D.R. & van Winkel, D. (2016) The ecology of New Zealand’s lizards. In: Chapple, D. (Ed.), New Zealand Lizards. Springer International Publishing, Cham, Switzerland, pp. 133–168. https://doi.org/10.1007/978-3-319-41674-8_6
  35. Heinicke, M.P., Nielsen, S.V., Bauer, A.M., Kelly, R., Geneva, A.J., Daza, J.D., Keating, S.E. & Gamble, T. (2023) Reappraising the evolutionary history of the largest known gecko, the presumably extinct Hoplodactylus delcourti, via high-throughput sequencing of archival DNA. Scientific Reports, 13 (1), 9141. https://doi.org/10.1038/s41598-023-35210-8
  36. Hillis, D.M. (2019) Species delimitation in herpetology. Journal of Herpetology, 53 (1), 3–12. https://doi.org/10.1670/18-123
  37. Hitchmough, R.A. (1997) A systematic revision of the New Zealand Gekkonidae. Unpublished Ph.D. Dissertation, Victoria University, Wellington, 370 pp.
  38. Hitchmough, R.A., Patterson, G.B. & Chapple, D.G. (2016a) Putting a name to diversity: taxonomy of the New Zealand lizard fauna. In: Chapple, D. (Ed.), New Zealand Lizards. Springer International Publishing, Cham, pp. 87–108. https://doi.org/10.1007/978-3-319-41674-8_4
  39. Hitchmough, R., Barr, B., Lettink, M., Monks, J., Reardon, J., Tocher, M., van Winkel, D. & Rolfe, J. (2016b) Conservation status of New Zealand reptiles, 2015. In: New Zealand Threat Classification Series. Vol. 17. Department of Conservation, Wellington, 14 pp.
  40. Hitchmough, R.A., Nielsen, S.V. & Bauer, A.M. (2020) Earning your stripes: a second species of striped gecko in the New Zealand gecko genus Toropuku (Gekkota: Diplodactylidae). Zootaxa, 4890 (4), 578–588. https://doi.org/10.11646/zootaxa.4890.4.9
  41. Hitchmough, R.A., Nielsen, S.V., Lysaght, J.A. & Bauer, A.M. (2021a) A new species of Naultinus from the Te Paki area, northern New Zealand. Zootaxa, 4915 (3), 389–400. https://doi.org/10.11646/zootaxa.4915.3.7
  42. Hitchmough, R.A., Barr, B., Knox, C., Lettink, M., Monks, J.M., Patterson, G.B., Reardon, J.T., van Winkel, D., Rolfe, J. & Michel, P. (2021b) Conservation status of New Zealand reptiles, 2021. In: New Zealand Threat Classification Series. Vol. 35. Department of Conservation, Wellington, pp. 1–23.
  43. Holloway, B.A. (1961) A systematic revision of the New Zealand Lucanidae (Insecta: Coleoptera). Bulletin of the Dominion Museum, Wellington, 20, 1–139.
  44. Iturralde-Vinent, M.A. & MacPhee, R.D.E. (1996) Age and paleogeographical origin of Dominican amber. Science, 273 (5283), 1850–1852. https://doi.org/10.1126/science.273.5283.1850
  45. Jewell, T.J. (2017) Oligosoma awakopaka n. sp. (Reptilia: Scincidae) from Fiordland National Park, New Zealand. BioGecko, 4, 4–19.
  46. Jewell, T.J. (2019a) Skinks of Southern New Zealand: A Field Guide. Edition 4. Jewell Publications, Invercargill, 102 pp.
  47. Jewell, T.R. (2019b) New Zealand forest-dwelling skinks of the Oligosoma oliveri (McCann) species-complex (Reptilia: Scincidae): reinstatement of O. pachysomaticum (Robb) and an assessment of historical distribution ranges. Zootaxa, 4688 (3), 382–398. https://doi.org/10.11646/zootaxa.4688.3.5
  48. Jombart, T., Devillard, S. & Balloux, F. (2010) Discriminant analysis of principal components: a new method for the analysis of genetically structured populations. BMC Genetics, 11 (1), 1–15. https://doi.org/10.1186/1471-2156-11-94
  49. Jombart, T. & Collins, C. (2015) A tutorial for discriminant analysis of principal components (DAPC) using adegenet 2.0.0. Available from: http://adegenet.r-forge.r-project.org/files/tutorial-dapc.pdf (accessed 30 December 2021)
  50. King, M. (1987) Origin of the Gekkonidae: chromosomal and albumin evolution suggests Gondwanaland. Search, 18 (5), 252–254.
  51. Kluge, A.G. (1995) Cladistic relationships of sphaerodactyl lizards. American Museum Novitates, 3139, 1–23.
  52. Knox, C., Hitchmough, R.A., Nielsen, S.V., Jewell, T. & Bell, T. (2021) A new, enigmatic species of black-eyed gecko (Reptilia: Diplodactylidae: Mokopirirakau) from North Otago, New Zealand. Zootaxa, 4964 (1), 140–156. https://doi.org/10.11646/zootaxa.4964.1.7
  53. Knox, C., Patterson, G.B. & Chapple, D.G. (2023). Oligosoma aureocola sp. nov. (Reptilia: Scincidae) from the northern Southland high country of Aotearoa/ New Zealand. Zootaxa, 5285 (2), 271–292. https://doi.org/10.11646/zootaxa.5285.2.3
  54. Lanfear, R., Calcott, B., Ho, S.Y. & Guindon, S. (2012) PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29 (6), 1695–1701. https://doi.org/10.1093/molbev/mss020
  55. Lee, M.S., Hutchinson, M.N., Worthy, T.H., Archer, M., Tennyson, A.J., Worthy, J.P. & Scofield, R.P. (2009) Miocene skinks and geckos reveal long-term conservatism of New Zealand’s lizard fauna. Biology Letters, 5 (6), 833–837. https://doi.org/10.1098/rsbl.2009.0440
  56. Lettink, M. & Whitaker, T. (2006) Hoplodactylus maculatus (common gecko): longevity. Herpetological Review, 37 (2), 223–224.
  57. Lleonart, J., Salat, J. & Torres, G.J. (2000) Removing allometric effects of body size in morphological analysis. Journal of Theoretical Biology, 205 (1), 85–93. https://doi.org/10.1006/jtbi.2000.2043
  58. Melzer, S., Hitchmough, R., van Winkel, D., Wedding, C., Chapman, S. & Rixon, M. (2022) Conservation status of reptile species in Tāmaki Makaurau/Auckland. Auckland Council Technical Report, TR2022/3, 1–20.
  59. Morgan-Richards, M. (1997) Intraspecific karyotype variation is not concordant with allozyme variation in the Auckland tree weta of New Zealand, Hemideina thoracica (Orthoptera: Stenopelmatidae). Biological Journal of the Linnean Society, 60 (4), 423–442. https://doi.org/10.1111/j.1095-8312.1997.tb01505.x
  60. Morgan-Richards. M., Trewick, S.A. & Wallis, G.P. (2001) Chromosome races with Pliocene origins: evidence from mtDNA. Heredity, 86 (3), 303–312. https://doi.org/10.1046/j.1365-2540.2001.00828.x
  61. Myers, S.S., Holwell, G.I. & Buckley, T. R. (2017) Genetic and morphometric data demonstrate alternative consequences of secondary contact in Clitarchus stick insects. Journal of Biogeography, 44 (9), 2069–2081. https://doi.org/10.1111/jbi.13004
  62. Naish, T.R. (2005) New Zealand’s shallow-marine record of Pliocene-Pleistocene global sea-level and climate change. Journal of the Royal Society of New Zealand, 35 (1–2), 1–8.
  63. Naish, T.R. & Wilson, G.S. (2009) Constraints on the amplitude of Mid-Pliocene (3.6–2.4 Ma) eustatic sea-level fluctuations from the New Zealand shallow-marine sediment record. Philosophical Transactions of the Royal Society A: Mathematical, Physical and Engineering Sciences, 367 (1886), 169–187. https://doi.org/10.1098/rsta.2008.0223
  64. Nielsen, S.V., Bauer, A.M., Jackman, T.R., Hitchmough, R.A. & Daugherty, C.H. (2011) New Zealand geckos (Diplodactylidae): cryptic diversity in a post-Gondwanan lineage with trans-Tasman affinities. Molecular Phylogenetics and Evolution, 59 (1), 1–22. https://doi.org/10.1016/j.ympev.2010.12.007
  65. Nelson, N.J., Romijn, R.L., Dumont, T., Reardon, J.T., Monks, J.M., Hitchmough, R.A., Empson, R. & Briskie, J.V. (2016) Lizard conservation in mainland sanctuaries. In: Chapple, D.G. (Ed.), New Zealand Lizards. Springer International Publishing, Cham, pp. 321−339. https://doi.org/10.1007/978-3-319-41674-8_12
  66. Padial, J.M., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7 (1), 1–14. https://doi.org/10.1186/1742-9994-7-16
  67. Patterson, G.B. & Hitchmough, R.A. (2021) A new alpine skink species (Scincidae: Eugongylinae: Oligosoma) from Kahurangi National Park, New Zealand. Zootaxa, 4920 (4), 495–508. https://doi.org/10.11646/zootaxa.4920.4.2
  68. Powell, A.W.B. (1949) The species problem in New Zealand land snails. Transactions of the Royal Society of New Zealand, 77, 202–208.
  69. Purdie, S. (2022) A Naturalist’s Guide to the Reptiles & Amphibians of New Zealand. John Beaufoy Publishing Ltd, Oxford, 176 pp.
  70. Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67 (5), 901–904. https://doi.org/10.1093/sysbio/syy032
  71. Robb, J. (1980) Three new species of gekkonid lizards, genera Hoplodactylus Fitzinger and Heteropholis Fischer, from New Zealand. National Museum of New Zealand Records, 1 (19), 305–310.
  72. Rodríguez, F.J.L.O.J., Oliver, J.L., Marín, A. & Medina, J.R. (1990) The general stochastic model of nucleotide substitution. Journal of Theoretical Biology, 142 (4), 485–501. https://doi.org/10.1016/s0022-5193(05)80104-3
  73. Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19 (12), 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
  74. Scarsbrook, L., Walton, K., Rawlence, N. J. & Hitchmough, R. A. (2023) Revision of the New Zealand gecko genus Hoplodactylus, with the description of a new species. Zootaxa, 5228 (3), 267–291. https://doi.org/10.11646/zootaxa.5228.3.3
  75. Skipwith, P.L., Bauer, A.M., Jackman, T.R. & Sadlier, R.A. (2016) Old but not ancient: coalescent species tree of New Caledonian geckos reveals recent post‐inundation diversification. Journal of Biogeography, 43 (6), 1266–1276. https://doi.org/10.1111/jbi.12719
  76. Skipwith, P.L., Bi, K. & Oliver, P.M. (2019) Relicts and radiations: Phylogenomics of an Australasian lizard clade with east Gondwanan origins (Gekkota: Diplodactyloidea). Molecular Phylogenetics and Evolution, 140, 106589. https://doi.org/10.1016/j.ympev.2019.106589
  77. Spencer, H.G., Brook, F.J. & Kennedy, M. (2006) Phylogeography of kauri snails and their allies from Northland, New Zealand (Mollusca: Gastropoda: Rhytididae: Paryphantinae). Molecular Phylogenetics and Evolution, 38 (3), 835–842. https://doi.org/10.1016/j.ympev.2005.10.015
  78. Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22 (21), 2688–2690. https://doi.org/10.1093/bioinformatics/btl446
  79. Steindachner, F. (1867) Reptilien. In: Reise der Österreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858,1859 unter den Befehlen des Commodore B. von Wüllerstorf-Urbair (Zoologie). Vol. 1. Part 3. Kaiserlich-Königliche Hof- und Staatsdruckerei; in Commission bei K. Gerold‘s Sohn, Wien, pp. 1–98.
  80. Sutherland, D.L., Hogg, I.D. & Waas, J.R. (2010) Phylogeography and species discrimination in the Paracalliope fluviatilis species complex (Crustacea: Amphipoda): can morphologically similar heterospecifics identify compatible mates? Biological Journal of the Linnean Society, 99 (1), 196–205. https://doi.org/10.1111/j.1095-8312.2009.01344.x
  81. Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution, 30 (12), 2725–2729. https://doi.org/10.1093/molbev/mst197
  82. Thorpe, R.S. (1975) Quantitative handling of characters useful in snake systematics with particular reference to interspecific variation in the Ringed Snake Natrix natrix (L.). Biological Journal of the Linnean Society, 7 (1), 27–43. https://doi.org/10.1111/j.1095-8312.1975.tb00732.x
  83. Thorpe, R.S. (1983) A review of the numerical methods for recognizing and analyzing racial differentiation. In: Felsenstein, J. (Ed.), Numerical Taxonomy. NATo ASI Series. Series G: Ecological Sciences. Vol. 1. Springer-Verlag, Berlin, pp. 404–423. https://doi.org/10.1007/978-3-642-69024-2_43
  84. Townsend, A.J., de Lange, P.J., Duffy, C.A., Miskelly, C.M., Molloy, J. & Norton, D.A. (2008) New Zealand Threat Classification System Manual. Department of Conservation, Wellington, 35 pp.
  85. Turan, C. (1999) A note on the examination of morphometric differentiation among fish populations: The Truss System. Turkish Journal of Zoology, 23 (3), 259–263.
  86. van Winkel, D., Baling, M. & Hitchmough, R. (2018) Reptiles and Amphibians of New Zealand. Auckland University Press, Auckland, 366 pp.
  87. Wallis, G.P. & Trewick, S.A. (2009) New Zealand phylogeography: evolution on a small continent. Molecular Ecology, 18 (17), 3548–3580. https://doi.org/10.1111/j.1365-294x.2009.04294.x
  88. Whitaker, A.H. (1973) Lizard populations on islands with and without Polynesian rats, Rattus exulans (Peale). Proceedings of the New Zealand Ecological Society, 20, 121–130.
  89. Whitaker, A.H. (1984) Hoplodactylus kahutarae n. sp. (Reptilia: Gekkonidae) from the Seaward Kaikoura Range, Marlborough, New Zealand. New Zealand Journal of Zoology, 11 (3), 259–270. https://doi.org/10.1080/03014223.1984.10428239
  90. Wightman, A.T. (1956) Birds on Oaia 1953-54. Notornis, 7 (2), 59–60.
  91. Worthy, T.H., Tennyson, A.J., Jones, C., McNamara, J.A. & Douglas, B.J. (2007) Miocene waterfowl and other birds from Central Otago, New Zealand. Journal of Systematic Palaeontology, 5 (1), 1–39. https://doi.org/10.1017/s1477201906001957
  92. Worthy, T.H. (2016) A Review of the Fossil Record of New Zealand Lizards. In: Chapple, D. (Ed.), New Zealand Lizards. Springer International Publishing, Cham, pp. 65–86. https://doi.org/10.1007/978-3-319-41674-8_3
  93. Wotton, D.M., Drake, D.R., Powlesland, R.G. & Ladley, J.J. (2016) The role of lizards as seed dispersers in New Zealand. Journal of the Royal Society of New Zealand, 46 (1), 40–65. https://doi.org/10.1080/03036758.2015.1108924