Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 5311 No. 3: 30 Jun. 2023
Type: Article
Published: 2023-06-30
DOI: 10.11646/zootaxa.5311.3.1
Page range: 301-339
Abstract views: 2665
PDF downloaded: 39

Whip it into shape: Revision of the Demansia psammophis (Schlegel, 1837) complex (Squamata: Elapidae), with a description of a new species from central Australia

School of Biological Sciences; University of Adelaide; Adelaide; South Australia 5005; AUSTRALIA; Faculty of Science & Engineering; Flinders University; Bedford Park; South Australia 5042; AUSTRALIA
Research Associate; Collections & Research; Western Australian Museum; 49 Kew Street; Welshpool; Western Australia 6106; AUSTRALIA
Snakes Harmful & Harmless; 9 Birch Place; Stoneville; Western Australia 6081; AUSTRALIA
South Australian Museum; North Terrace; Adelaide; South Australia 5000; AUSTRALIA; Faculty of Science & Engineering; Flinders University; Bedford Park; South Australia 5042; AUSTRALIA
Reptilia Demansia cyanochasma morphology mtDNA nDNA phylogeny Reptiliasynonymy taxonomy whip snakes

Abstract

The genus Demansia Günther is the most diverse genus of Australian terrestrial elapids. A phylogenetic framework for the familiar but problematic taxa D. psammophis and D. reticulata (Gray) has been long overdue to ascertain interspecific relationships and resolve unclear taxonomic issues. Here we present an integrated molecular phylogenetic and morphological analyses to review species delineation, resulting in confirmation that both D. psammophis and D. reticulata are full species and that some populations referred to D. r. cupreiceps Storr are not distinguishable from more typical D. reticulata. We also find the widespread central Australian population (treated by most authors as part of cupreiceps) to be specifically distinct. We redescribe D. psammophis and D. reticulata to clarify morphological and geographical boundaries and describe D. cyanochasma sp. nov. based on a combination of molecular genetic markers, details of colour and pattern, adult total length and a few morphometric attributes. We also designate a lectotype for D. psammophis from the original syntype series and comment on the necessity for further taxonomic refinement of this distinctive group.

 

References

  1. Adler, K. (2012) Herpetologists of the past, part 3. Biographies. In: Adler, K. (Ed.), Contributions to the History of Herpetology, Vol 3. Society for the Study of Amphibians and Reptiles, Vancouver, pp. 9‒360.
  2. Allen, L. & Vogel, G. (2019) Venomous Snakes of Australia and Oceania. Terralog Volume 18. Edition Chimaira, Frankfurt am Main, 168 pp.
  3. Arévalo, E., Davis, S.K. & Sites, J.W. (1994) Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology, 43, 387‒418. https://doi.org/10.1093/sysbio/43.3.387 DOI: https://doi.org/10.1093/sysbio/43.3.387
  4. Ashman, L.G., Bragg, J.G., Doughty, P., Hutchinson, M.N., Bank, S., Matzke, N.J., Oliver, P. & Moritz, C. (2018) Diversification across biomes in a continental lizard radiation. Evolution, 72, 1553–1569. https://doi.org/10.1111/evo.13541 DOI: https://doi.org/10.1111/evo.13541
  5. Bonnemains, J., Forsyth, E. & Smith, B. (Eds.) (1988) The Artwork of the French Voyage of Discovery to the Southern Lands, 1801–1804. Oxford University Press, Melbourne, 360 pp.
  6. Boulenger, G.A. (1896) Catalogue of the Snakes in the British Museum (Natural History).
  7. Vol. III. Trustees of the British Museum (Natural History), London, 727 pp.
  8. Brandle, R. (2010) A biological survey of the Eyre Peninsula, South Australia. Department for Environment and Heritage, South Australia, 114 pp.
  9. Burbidge, A.A., Fuller, P.J. & McKenzie, N.L. (1995) Vertebrate fauna. In: The Biological Survey of the Eastern Goldfields of Western Australia. Part 12. Barlee‒Menzies Study Area. Records of the Western Australian Museum, 49 (Supplement), 208‒245.
  10. Bush, B. (2017) Additions to the description of Paroplocephalus atriceps (Serpentes: Elapidae) with a discussion on pupil shape in it and other Australian snakes. Zootaxa, 4344 (2), 333‒344. https://doi.org/10.11646/zootaxa.4344.2.8 DOI: https://doi.org/10.11646/zootaxa.4344.2.8
  11. Bush, B., Maryan, B., Browne-Cooper, R. & Robinson, D. (2007) Reptiles and Frogs in the Bush: Southwestern Australia. University of Western Australia Press, Perth, 302 pp.
  12. Bush, B., Maryan, B., Browne-Cooper, R. & Robinson, D. (2010) Field Guide to Reptiles & Frogs of the Perth Region. Western Australian Museum, Welshpool, 183 pp.
  13. Bush, B. & Maryan, B. (2011) Field Guide to Snakes of the Pilbara Western Australia. Western Australian Museum, Welshpool, 112 pp.
  14. Chapman, A. & Dell, J. (1985) Biology and zoogeography of the amphibians and reptiles of the Western Australian Wheatbelt. Records of the Western Australian Museum, 12, 1‒46.
  15. Chapple, D.G., Tingley, R., Mitchell, N.J., Macdonald, S.L., Keogh, J.S., Shea, G.M., Bowles, P., Cox, N.A. & Woinarski, J.C.Z. (2019) The Action Plan for Australian Lizards and Snakes 2017. CSIRO Publishing, Melbourne, 663 pp. https://doi.org/10.1071/9781486309474 DOI: https://doi.org/10.1071/9781486309474
  16. Cogger, H.G. (1967) Australian Reptiles in Colour. Reed Books, Sydney, 112 pp.
  17. Cogger, H.G. (1979) Reptiles & Amphibians of Australia. 2nd Edition. Reed Books, Sydney, 608 pp.
  18. Cogger, H.G. (1983) Reptiles & Amphibians of Australia. 3rd Edition. Reed Books, Sydney, 660 pp.
  19. Cogger, H.G. (2018) Reptiles & Amphibians of Australia. 7th Edition (Updated). CSIRO Publishing, Melbourne, 1080 pp.
  20. https://doi.org/10.1071/9781486309702 DOI: https://doi.org/10.1071/9781486309702
  21. Cogger, H.G., Cameron, E.E. & Cogger, H.M. (1983) Zoological Catalogue of Australia. Vol. 1. Amphibia and Reptilia. Bureau of Flora and Fauna, Canberra, vi + 313 pp.
  22. Dell, J. & How, R.A. (1984) Vertebrate Fauna. In: The Biological Survey of the Eastern Goldfields of Western Australia. Part 2. Widgiemooltha‒Zanthus Area. Records of the Western Australian Museum, 18 (Supplement), pp. 57‒81.
  23. Dell, J. & How, R.A. (1988) Vertebrate Fauna. In: The Biological Survey of the Eastern Goldfields of Western Australia. Part 5. Edjudina-Menzies Study Area. Records of the Western Australian Museum, 31 (Supplement), pp. 38‒68.
  24. Dowling, H.G. (1951) A proposed standard of counting ventrals in snakes. British Journal of Herpetology, 1, 97‒99. https://doi.org/10.2307/1437542 DOI: https://doi.org/10.2307/1437542
  25. Duméril, A.M.C., Bibron, G. & Duméril, A.H.A. (1854) Erpétologie généraleou histoire naturelle complete des reptiles. VII. Librairie Encyclopédique de Roret, Paris, xii + 1536 pp. DOI: https://doi.org/10.5962/bhl.title.118797
  26. Duyker, E. (2014) Dumont d’Urville: Explorer & Polymath. Otago University Press,
  27. Dunedin, 671 pp.
  28. Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32, 1792‒1797. https://doi.org/10.1093/nar/gkh340 DOI: https://doi.org/10.1093/nar/gkh340
  29. Ehmann, H. (2005) South Australian Rangelands and Aboriginal Lands Wildlife Management Manual: a resource handbook. Department of Water, Land and Biodiversity Conservation, South Australia, 516 pp.
  30. Eipper, S. (2012) A Guide to Australian Snakes in Captivity Elapids and Colubrids. Reptile Publications, Burleigh BC, 279 pp.
  31. Georges, A., Gruber, B., Pauly, G.B., White, D., Adams, M., Young, M.J., Kilian, A., Zhang, X., Shaffer, H.B. & Unmack, P.J. (2018) Genomewide SNP markers breathe new life into phylogeography and species delimitation for the problematic short-necked turtles (Chelidae: Emydura) of eastern Australia. Molecular Ecology, 27, 5195‒5213. https://doi.org/10.1111/mec.14925 DOI: https://doi.org/10.1111/mec.14925
  32. Girard, C. (1858) United States exploring expedition during the years 1838, 1839, 1840, 1841, 1842, under the command of Charles Wilkes, U.S.N. Vol. 20. Herpetology.C. Sherman & Son, Philadelphia, Pennsylvania, xv + 492 pp.
  33. Glauert, L. (1957) A Handbook of the Snakes of Western Australia. 2nd Edition. Western
  34. Australian Naturalists’ Club, Perth, 60 pp.
  35. Gray, J.E. (1842) Description of some hitherto unrecorded species of Australian reptiles and batrachians. In: Zoological Miscellany. Treuttel, Wȕrtz & Co., London, pp. 51‒57.
  36. Greer, A.E. (1997) The Biology and Evolution of Australian Snakes. Surrey, Beatty & Sons, Sydney, 358 pp.
  37. Griffiths, K. (2006) Frogs & Reptiles of the Sydney Region. Reed New Holland, Sydney, 128 pp.
  38. Gruber, B., Unmack, P.J., Berry, O.F. & Georges, A. (2018) dartr: An r package to facilitate analysis of SNP data generated from reduced representation genome sequencing. Molecular Ecology Resources, 18, 691‒699. https://doi.org/10.1111/1755-0998.12745 DOI: https://doi.org/10.1111/1755-0998.12745
  39. Guibé, J. & Roux-Estevé, R. (1972) Les types de Schlegel (Ophidiens), présents dans les collections du Muséum National d’Histoire Naturelle de Paris. Zoologische Mededelingen, 47, 129‒134.
  40. Günther, A. (1858) Catalogue of colubrine snakes in the collection of the British Museum. British Museum (Natural History), London, 281 pp. https://doi.org/10.1080/00222936308681373
  41. Günther, A. (1863) On new species of snakes in the collection of the British Museum. Annals Magazine of Natural History, 11, 20‒25. DOI: https://doi.org/10.1080/00222936308681373
  42. How, R.A. & Dell, J. (2000) Ground vertebrate fauna of Perth’s vegetation remnants: impact of 170 years of urbanization. Pacific Conservation Biology, 6, 198‒217. https://doi.org/10.1071/PC000198 DOI: https://doi.org/10.1071/PC000198
  43. Hutchinson, M. & Tyler, M. (1996) Reptiles and amphibians. In: Davies, M., Twidale, C.R. & Tyler, M.J. (Eds.), Natural History of the Flinders Ranges. Royal Society of South Australia, Adelaide, pp. 149‒158.
  44. Jan, G. (1859) Prodrome d’une iconographie descriptive des ophidiens et description sommaire de nouvelles espèces de serpents venimeux. Veuve Bouchard-Huzard, Paris, 2, 1‒32. https://doi.org/10.5962/bhl.title.12473 DOI: https://doi.org/10.5962/bhl.title.12473
  45. Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K., Von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods, 14, 587‒589. https://doi.org/10.1038/nmeth.4285 DOI: https://doi.org/10.1038/nmeth.4285
  46. Kendrick, P.G. (1993) Biogeography of the vertebrates of the Cape Range Peninsula, Western Australia. In: Humphreys, W.F. (Ed.), The Biogeography of Cape Range, Western Australia. Records of the Western Australian Museum, 45 (Supplement), pp. 193‒206.
  47. Keogh, J.S. (1999) Evolutionary implications of hemipenial morphology in the terrestrial Australian elapid snakes. Zoological Journal of the Linnean Society, 125, 239−278. https://doi.org/10.1111/j.1096-3642.1999.tb00592.x DOI: https://doi.org/10.1111/j.1096-3642.1999.tb00592.x
  48. Kilian, A., Wenzl, P., Huttner, E., Carling, J., Xia, L., Blois, H., Caig, V., Heller-Uszynska, K., Jaccoud, D., Hopper, C. & Aschenbrenner-Kilian, M. (2012) Diversity arrays technology: a generic genome profiling technology on open platforms. In: Data production and analysis in population genomics. Humana Press, Totowa, New Jersey, pp. 67‒89. https://doi.org/10.1007/978-1-61779-870-2_5 DOI: https://doi.org/10.1007/978-1-61779-870-2_5
  49. Kinghorn, J.R. (1956) The Snakes of Australia. 2nd Edition. Halstead Press, Sydney, 197 pp.
  50. Klemmer, K. (1963) Liste der rezenten Giftschlangen: Elapidae, Hydropheidae, Viperidae und Crotalidae. In: Die Giftschlangen der Erde. N.G. Elwert Universitäts-U. Verlagsbuchhand, Marburg, pp. 256‒464.
  51. Krefft, G. (1866) On snakes observed in the neighbourhood of Sydney. Transactions of the Philosophical Society of New South Wales, 1862‒1865, 34‒60. https://doi.org/10.5962/p.345621 DOI: https://doi.org/10.5962/p.345621
  52. Laver, R.J., Doughty, P. & Oliver, P.M. (2018) Origins and patterns of endemic diversity in two specialized lizard lineages from the Australian Monsoonal Tropics (Oedura spp.). Journal of Biogeography, 45, 142–153. https://doi.org/10.1111/jbi.13127 DOI: https://doi.org/10.1111/jbi.13127
  53. Lee, M.S., Sanders, K.L., King, B. & Palci, A. (2016) Diversification rates and phenotypic evolution in venomous snakes (Elapidae). Royal Society Open Science, 3, 150277. https://doi.org/10.1098/rsos.150277 DOI: https://doi.org/10.1098/rsos.150277
  54. Lucas, A.H.S. & Frost, C. (1896) Reptilia. In: Spencer, B. (Ed.) Report on the work of the Horn Scientific Expedition to Central Australia. Part II. Zoology. Dulau and Co., London, pp. 112‒151, pls. 8‒12.
  55. Maryan, B. (1996) Herpetofauna of Dirk Hartog Island Shark Bay Area, Western Australia. Herpetofauna, 26, 8‒11.
  56. Maryan, B., Brennan, I.G., Hutchinson, M.N. & Geidans, L.S. (2020) What’s under the hood? Phylogeny and taxonomy of the snake genera Parasuta Worrell and Suta Worrell (Squamata: Elapidae), with a description of a new species from the Pilbara, Western Australia. Zootaxa, 4778 (1), 1‒47. https://doi.org/10.11646/zootaxa.4778.1.1 DOI: https://doi.org/10.11646/zootaxa.4778.1.1
  57. McDowell, S.B. & Cogger, H.G. (1967) Aspidomorphus, a genus of New Guinea snakes of the family Elapidae, with notes on related genera. Journal of Zoology, London, 151, 497‒543. https://doi.org/10.1111/j.1469-7998.1967.tb02130.x DOI: https://doi.org/10.1111/j.1469-7998.1967.tb02130.x
  58. McKenzie, N.L., Rolfe, J.K., Aplin, K.P., Cowan, M.A. & Smith, L.A. (2000) Herpetofauna of the southern Carnarvon Basin, Western Australia. In: Burbidge, A.H., Harvey, M.S. & McKenzie, N.L. (Eds.), Records of the Western Australian Museum, 61 (Supplement), pp. 335‒360. https://doi.org/10.18195/issn.0313-122x.61.2000.335-360 DOI: https://doi.org/10.18195/issn.0313-122x.61.2000.335-360
  59. Mengden, G.A. (1985) Australian elapid phylogeny: a summary of the chromosomal and electrophoretic data. In: Grigg, G., Shine, R. & Ehmann, H. (Eds.), Biology of Australasian Frogs and Reptiles. Surrey Beatty & Sons, Sydney, pp. 185‒192.
  60. Mitchell, F.J. (1961) Harmless or Harmful? 2nd Edition. South Australian Museum, Adelaide, 30 pp.
  61. Nguyen, L.T., Schmidt, H.A., Von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution,32, 268‒274. https://doi.org/10.1093/molbev/msu300 DOI: https://doi.org/10.1093/molbev/msu300
  62. Orange, P. (1991) Notes on the diet of the Yellow-faced Whip Snake Demansia psammophis psammophis in the Kambalda region of Western Australia. Herpetofauna, 21, 29‒30.
  63. Orange, P. (1994) A note regarding colour change in Demansia psammophis psammophis (Serpentes: Elapidae). Herpetofauna, 24, 41.
  64. OZCAM (2022) Online Zoological Collections of Australian Museums. Available from: https://OZCAM.org.au/search Demansia+psammophis (accessed 5 July 2022)
  65. Powney, G.D., Grenyer, R., Orme, C.D.L., Owens, P.F. & Meiri, S. (2010) Hot, dry and different: Australian lizard richness is unlike that of mammals, amphibians and birds. Global Ecology and Biogeography, 19, 386–396. https://doi.org/10.1111/j.1466-8238.2009.00521.x DOI: https://doi.org/10.1111/j.1466-8238.2009.00521.x
  66. Rambaut, A. (2012) FigTree v1.4.4, a graphical viewer of phylogenetic trees. Available from: http://tree.bio.ed.ac.uk/software/figtree/ (accessed 8 May 2023)
  67. Reid, J.R.W., Kerle, J.A., Baker, L. & Jones, K.R. (1993) Reptiles and frogs. In: Uluru Fauna The Distribution and Abundance of Vertebrate Fauna of Uluru (Ayers Rock-Mount Olga) National Park, N.T. Australian National Parks and Wildlife Service series, Kowari, 4, pp. 58‒68.
  68. Robertson, P. & Coventry, A.J. (2019) Reptiles of Victoria A Guide to Identification and Ecology. CSIRO Publishing, Melbourne, 323 pp. https://doi.org/10.1071/9781486309993 DOI: https://doi.org/10.1071/9781486309993
  69. Scanlon, J.D. (2003) The Australian elapid genus Cacophis: morphology and phylogeny of rainforest crowned snakes. Herpetological Journal, 13, 1–20.
  70. Scanlon, J.D. & Lee, M.S.Y. (2004) Phylogeny of Australasian venomous snakes (Colubroidea, Elapidae, Hydrophiinae) based on phenotypic and molecular evidence. Zoologica Scripta, 33, 335–366. https://doi.org/10.1111/j.0300-3256.2004.00151.x DOI: https://doi.org/10.1111/j.0300-3256.2004.00151.x
  71. Schätti, B., Ineich, I. & Kucharzewski, C. (2010) Nominal taxa of Spalerosophis diadema (Schlegel, 1837) from Iraq to Pakistan—two centuries of confusion (Reptilia: Squamata: Colubrinae). Revue Suisse de Zoologie, 117, 637‒664. https://doi.org/10.5962/bhl.part.145579 DOI: https://doi.org/10.5962/bhl.part.145579
  72. Schlegel, H. (1837a) Essai sur la physionomie des serpens. I. Partie Générale. M.H. Schonekat, Amsterdam, xxviii + 251 pp. https://doi.org/10.5962/bhl.title.4273
  73. Schlegel, H. (1837b) Essai sur la physionomie des serpens. II. Partie Descriptive. M.H. Schonekat, Amsterdam, 606 + xv pp. https://doi.org/10.5962/bhl.title.4693 DOI: https://doi.org/10.5962/bhl.title.4273
  74. Shea, G.M. (1998) Geographic variation in scalation and size of the black whip snakes (Squamata: Elapidae: Demansia vestigiata complex): evidence for two broadly sympatric species. The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 14, 41‒61. https://doi.org/10.5962/p.287491 DOI: https://doi.org/10.5962/p.287491
  75. Shea, G.M. & Scanlon, J.D. (2007) Revision of the small tropical whipsnakes previously referred to Demansia olivacea (Gray, 1842) and Demansia torquata (Gȕnther, 1862) (Squamata: Elapidae). Records of the Australian Museum, 59, 117‒142. https://doi.org/10.3853/j.0067-1975.59.2007.1488 DOI: https://doi.org/10.3853/j.0067-1975.59.2007.1488
  76. Shea, G., Gaikhorst, G. & Cowan, M. (2017) Demansia psammophis. The IUCN Red List of Threatened Species, 2017, e.T42492989A42492999.
  77. Shine, R. (1980) Ecology of eastern Australian whipsnakes of the genus Demansia. Journal of Herpetology, 14, 381‒389. https://doi.org/10.2307/1563694 DOI: https://doi.org/10.2307/1563694
  78. Storr, G.M. (1978) Whip snakes (Demansia: Elapidae) of Western Australia. Records of the Western Australian Museum, 6, 287‒301.
  79. Storr, G.M. & Harold, G. (1990) Amphibians and reptiles of the Shark Bay area, Western Australia. In: Berry, P.F., Bradshaw, S.D. & Wilson, B.R. (Eds.), Research in Shark Bay: Report of the France-Australe Bicentenary Expedition Committee. Western Australian Museum, Perth, pp. 279‒285.
  80. Storr, G.M., Harold, G. & Barron, G. (1978) Amphibians and Reptiles of the Northern Swan Coastal Plain. In: How, R.A. (Ed.), Faunal Studies of the Northern Swan Coastal Plain—A Consideration of Past and Future Changes. Western Australian Museum Report to Department of Conservation and Environment, Perth, Western Australia, pp. 172‒203.
  81. Storr, G. M., Smith, L.A. & Johnstone, R.E. (1986). Snakes of Western Australia. Western Australian Museum, Perth, 187 pp.
  82. Storr, G.M., Smith, L.A. & Johnstone, R.E. (2002) Snakes of Western Australia.Revised Edition. Western Australian Museum, Perth, 309 pp.
  83. Strickland, J.L., Carter, S., Kraus, F. & Parkinson, C.L. (2016) Snake evolution in Melanesia: origin of the Hydrophiinae (Serpentes, Elapidae), and the evolutionary history of the enigmatic New Guinean elapid Toxicocalamus. Zoological Journal of the Linnean Society, 17, 663‒678. https://doi.org/10.1111/zoj.12423 DOI: https://doi.org/10.1111/zoj.12423
  84. Swan, G. & Wilson, S. (2015) Where do they all come from? Animal movement immediately following a hummock grassland fire. Australian Zoologist, 37, 485‒491. https://doi.org./10.7882/AZ.2015.012 DOI: https://doi.org/10.7882/AZ.2015.012
  85. Swan, G, Sadlier, R. & Shea, G. (2022) A Field Guide to Reptiles of New South Wales. Fourth Edition. Reed New Holland, Sydney, 288 pp.
  86. Thompson, S.A. & Thompson, G.G. (2006) Reptiles of the Western Australian Goldfields. Goldfields Environmental Management Group, Kalgoorlie, Boulder, 140 pp.
  87. Thompson, G.G. & Thompson, S.A. (2007) Usefulness of funnel traps in catching small reptiles and mammals, with comments on the effectiveness of the alternatives. Wildlife Research, 34, 491‒497. https://doi.org/10.1071/WR06081 DOI: https://doi.org/10.1071/WR06081
  88. Wallach, V., Williams K.L. & Boundy, J. (2014) Snakes of the World A Catalogue of Living and Extinct Species. CRC Press, Taylor & Francis Group, Boca Raton, Florida, 1209 pp.
  89. Weigel, J. (1990) Australian Reptile Park’s Guide to Snakes of South-East Australia. Australian Reptile Park, Gosford, 103 pp.
  90. Wells, R.W. & Wellington, C.R. (1985) A classification of the Amphibia and Reptilia of
  91. Australia. Australian Journal of Herpetology, Supplementary Series, 1, 1‒61.
  92. Williams, C. & Maier, C. (Eds.) 2019. A Tribute to the Reptiles and Amphibians of Australia and New Zealand. Australian Herpetological Society. Reed New Holland, Sydney, 272 pp.
  93. Wilson, S. (2022) A Field Guide to Reptiles of Queensland. 3rd Edition. Reed New Holland, Sydney, 288 pp.
  94. Wilson, S.K. & Knowles, D.G. (1988) Australia’s Reptiles. A Photographic Reference to the Terrestrial Reptiles of Australia. Collins, Sydney, 447 pp.
  95. Wilson, S. & Swan, G. (2013) A Complete Guide to Reptiles of Australia. Fourth Edition. Reed New Holland, Sydney, 592 pp.
  96. Wilson, S. & Swan, G. (2021) A Complete Guide to Reptiles of Australia. Sixth Edition. Reed New Holland, Sydney, 688 pp.
  97. Worrell, E. (1963) Reptiles of Australia. Angus & Robertson, Sydney, 207 pp.