Tooth and cranial disparity in the fossil relatives of Sphenodon (Rhynchocephalia) dispute the persistent 'living fossil' label

Journal of Evolutionary Biology

Volumn 25, Issue 11, 2012, Pages 2194-2209

  Meloro, C ^a   Jones, M E H ^b  

^a UNIVERSITY OF HULL   (United Kingdom)

^b UNIVERSITY COLLEGE LONDON   (United Kingdom)

Author keywords

Comparative methods; Disparity through time; Geometric morphometrics; Partial least square; Tuatara; Vertebrate

Indexed keywords

CRANIUM; FOSSIL RECORD; MESOZOIC; MORPHOLOGY; MORPHOMETRY; PHYLOGENETICS; PHYLOGENY; REPTILE; TOOTH;

ANIMAL; ARTICLE; CLASSIFICATION; DENTITION; EVOLUTION; FACTUAL DATABASE; FOSSIL; HISTOLOGY; MORPHOGENESIS; PHYLOGENY; PHYSIOLOGY; PRINCIPAL COMPONENT ANALYSIS; REPTILE; SKULL; SPECIES DIFFERENCE; TIME; TOOTH;

ANIMALS; BIOLOGICAL EVOLUTION; DATABASES, FACTUAL; DENTITION; FOSSILS; MORPHOGENESIS; PHYLOGENY; PRINCIPAL COMPONENT ANALYSIS; REPTILES; SKULL; SPECIES SPECIFICITY; TIME FACTORS; TOOTH;

SPHENODON; SPHENODON PUNCTATUS; SPHENODONTIA; VERTEBRATA;

EID: 84868208884     PISSN: 1010061X     EISSN: 14209101     Source Type: Journal    
DOI: 10.1111/j.1420-9101.2012.02595.x     Document Type: Article

References (131)

1. Adams, D.C., Rohlf, F.J. & Slice, D.E. 2004. Geometric morphometrics: ten years of progress following the 'revolution'. Ital. J. Zool. 71: 5-16.
2. Alfaro, M.E., Santinia, F., Brock, C., Alamillo, H., Dornburg, A., Rabosky, D.L. et al. 2009. Nine exceptional radiations plus high turnover explain species diversity in jawed vertebrates. Proc. Nat. Acad. Sci. 106: 13410-13414.
3. Alibardi, L. & Toni, M. 2006. Distribution and characterization of keratins in the epidermis of the Tuatara (Sphenodon punctatus; Lepidosauria, Reptilia). Zool. Sci. 23: 801-807.
4. Anderson, P.S.L. 2009. The effects of trapping and blade angle of notched dentitions on fracture of biological tissues. J. Exp. Biol. 212: 3627-3632.
5. Apesteguía, S. & Jones, M.E.H. 2012. A Late Cretaceous 'tuatara' (Lepidosauria: Sphenodontinae) from South America. Cretac. Res. 24: 154-160.
6. Apesteguía, S. & Novas, F.E. 2003. Large Cretaceous sphenodontian from Patagonia provides insights into lepidosaur evolution in Gondwana. Nature 459: 609-612.
7. Apesteguía, S. & Rougier, G.W. 2007. A Late Campanian sphenodontid maxilla from northern Patagonia. Am. Mus. Nov. 3581: 1-11.
8. Asimov, I. 1990. Preserving 'living fossils' in New Zealand. The Day June 24: H6.
9. Benton, M.J. 1987. Progress and competition in macroevolution. Biol. Rev. 62: 305-338.
10. Bhullar, B.-A., Marugán-Lobón, J., Fernando, R., Bever, G.S., Rowe, T.B., Norell, M.A. et al. 2012. Birds have paedomorphic dinosaur skulls. Nature 487: 223-226.
11. Blomberg, S.P., Garland, T. Jr & Ives, A.R. 2003. Testing for phylogenetic signal in comparative data: behavioral traits are more labile. Evolution 57: 717-745.
12. Bogert, C.M. 1953. Why is it a lone survivor? Sci. Month. 76: 163-170.
13. Bonaparte, J.F. & Sues, H.-D. 2006. A New Species of Clevosaurus (Lepidosauria: Rhynchocephalia) From The Upper Triassic of Rio Grande Do Sul, Brazil. Palaeontology 49: 917-923.
14. Bookstein, F.L. 1989. 'Size and shape': a comment on semantics. Syst. Zool. 38: 173-180.
15. Brusatte, S.L., Benton, M.J., Ruta, M. & Graeme, T. 2008. Superiority, competition, and opportunism in the evolutionary radiation of dinosaurs. Science 321: 1485-1488.
16. Brusatte, S.L., Sakamoto, M., Montanari, S. & Harcourt-Smith, W.E.H. 2012. The evolution of cranial form and function in theropod dinosaurs: insights from geometric morphometrics. J. Evol. Biol. 25: 365-377.
17. Burbrink, F.T. & Pyron, R.A. 2009. How does ecological opportunity influence rates of speciation, extinction and morphological diversification in New World rattlesnakes. Evolution 64: 934-943.
18. Burton, M. 1956. Living Fossils. Readers Union Thames & Hudson, London.
19. Cardini, A. & Elton, S. 2008. Does the skull carry a phylogenetic signal? Evolution and modularity in the guenons. Biol. J. Linn. Soc. 93: 813-834.
20. Carroll, R.C. & Wild, R. 1994. Marine members of the Sphenodontia. In: In the Shadow of the Dinosaurs: Early Mesozoic Tetrapods (N.C. Fraser & H.-D. Sues, eds), pp. 70-83. Cambridge University Press, Cambridge.
21. Caumul, R. & Polly, P.D. 2005. Phylogenetic and environmental components of morphological variation: skull, mandible and molar shape in marmots (Marmota, Rodentia). Evolution 59: 2460-2472.
22. Ciampaglio, C.N., Kemp, M. & McShea, D.W. 2001. Detecting changes in morphospace occupation patterns in the fossil record: characterization and analysis of measures of disparity. Paleobiology 27: 695-715.
23. Cocude-Michel, M. 1963. Les rhynchocéphales et les sauriens des calcaires lithographiques (Jurassique Supérieur) d'Europe occidentale. Nouv. Arch. Mus. Hist. Nat. 7: 1-187.
24. Cree, A. 1994. Low annual reproductive output in female reptiles from New Zealand. N. Z. J. Zool. 21: 351-372.
25. Cree, A., Daugherty, C.H., Schafer, S.F. & Brown, D. 1991. Nesting and Clutch Size of Tuatara (Sphenodon guntheri) on North Brother Island, Cook Strait. Tuatara 31: 9-16.
26. Daza, J.D., Herrera, A., Thomas, R. & Claudio, H.J. 2009. Are you what you eat? A geometric morphometric analysis of gekkotan skull shape. Biol. J. Linn. Soc. 97: 677-707.
27. Dryden, I. & Mardia, K.V. 1998. Statistical Shape Analysis. John Wiley and Sons Ltd, Chichester.
28. Evans, S.E. 1980. The skull of a new eosuchian reptile from the Lower Jurassic of South Wales. Zool. J. Linn. Soc. 70: 203-264.
29. Evans, S.E. 1985. Tooth replacement in the Lower Jurassic lepidosaur Gephyrosaurus bridensis. Neues. Jahr. Geol. Paläontol. Monat. 7: 411-420.
30. Evans, S.E. 1992. A sphenodontian (Reptilia: Lepidosauria) from the Middle Jurassic of England. Neues. Jahr. Geol. Paläontol. Abh. 1992: 449-457.
31. Evans, S.E. 1995. Lizards: evolution, early radiation and biogeography. In: Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota, Short Papers (A. Sun & Y. Wang, eds), pp. 271-283. China Ocean Press, Beijing.
32. Evans, S.E. 1998. Crown group lizards from the Middle Jurassic of Britain. Palaeont. Abt. A 250: 123-154.
33. Evans, S.E. 2003. At the feet of the dinosaurs: the early history and radiation of lizards. Biol. Rev. 78: 513-551.
34. Evans, S.E. 2008. The skull of lizards and tuatara. In: Morphology H: The Skull of Lepidosauria (C. Gans, A.S. Gaunt & K. Adler, eds). Society for the Study of Amphibians and Reptiles: Ithaca, New York. Biol. Reptilia 20: 1-344.
35. Evans, S.E. & Borsuk-Bialynicka, M. 2009. A small lepidosauromorph reptile from the Early Triassic of Poland. Palaeontol. Polonica 65: 179-202.
36. Evans, S.E. & Jones, M.E.H. 2010. The origin, early history and diversification of lepidosauromorph reptiles. In: New Aspects of Mesozoic Biodiversity (S. Bandyopadhyay, eds). Lect. Notes Earth Sci. 132: 27-44.
37. Evans, S.E. & Kermack, K.A. 1994. Assemblages of small tetrapods from the Early Jurassic of Britain. In: In the Shadow of the Dinosaurs: Early Mesozoic Tetrapods (N.C. Fraser & H.-D. Sues, eds), pp. 271-283. Cambridge University Press, Cambridge.
38. Evans, A.R. & Sanson, G.D. 1998. The effect of tooth shape on the breakdown of insects. J. Zool. Lond. 246: 391-400.
39. Evans, A.R. & Sanson, G.D. 2003. The tooth of perfection: functional and spatial constraints on mammalian tooth shape. Biol. J. Linn. Soc. 78: 173-191.
40. Evans, S.E., Prasad, G.V.R. & Manhas, B.K. 2001. Rhynchocephalia (Diapsida: Lepidosauria) from the Jurassic Kota Formation of India. Zool. J. Linn. Soc. 133: 309-334.
41. Evans, S.E., Prasad, G.V.R. & Manhas, B.K. 2002. Fossil lizards from the Jurassic Kota formation of India. J. Vertebr. Paleontol. 22: 299-312.
42. Figueirido, B., Serrano-Alarcón, F.J., Slater, G.J. & Palmqvist, P. 2010. Shape at the cross-roads: homoplasy and history in the evolution of the carnivoran skull towards herbivory. J. Evol. Biol. 23: 2579-2594.
43. Foote, M. 1992. Paleozoic record of morphological diversity in blastozoan echinoderms. Proc. Natl Acad. Sci. USA 89: 7325-7329.
44. Foote, M. 1993. Discordance and concordance between morphological and taxonomic diversity. Paleobiology 19: 185-204.
45. Foote, M. 1997. The evolution of morphological diversity. Annu. Rev. Ecol. Syst. 28: 129-152.
46. Foth, C. & Rauhut, O.W.M. In press. Macroevolutionary and morphofunctional patterns in theropod skulls: a morphometric approach. Acta Palaeontol. Pol. http:dx.doi.org/10.4202/app.2011.0145.
47. Foth, C., Brusatte, S.L. & Butler, R.J. 2012. Do different disparity proxies converge on a common signal? Insights from the cranial morphometrics and evolutionary history of Pterosauria (Diapsida: Archosauria). J. Evol. Biol. 25: 904-915.
48. Fraser, N.C. 1982. A new rhynchocephalian from the British Upper Trias. Palaeontology 25: 709-725.
49. Fraser, N.C. 1985. Vertebrate faunas from Mesozoic fissure deposits of South West Britain. Mod. Geol. 9: 273-300.
50. Fraser, N.C. 1986. New Triassic sphenodontids from South West England and a review of their classification. Palaeontology 29: 165-186.
51. Fraser, N.C. 1988. The osteology and relationships of Clevosaurus (Reptilia: Sphenodontida). Philos. Trans. R. Soc. Lond. B 321: 125-178.
52. Fraser, N.C. 1994. Assemblages of small tetrapods from British Late Triassic fissure deposits. In: In the Shadow of the Dinosaurs: Early Mesozoic Tetrapods (N.C. Fraser & H.-D. Sues, eds), pp. 214-226. Cambridge University Press, Cambridge.
53. Fraser, N.C. & Benton, M.J. 1989. The Triassic reptiles Brachyrhinodon and Polysphenodon and the relationships of the sphenodontids. Zool. J. Linn. Soc. 96: 413-445.
54. Fraser, N.C. & Walkden, G.M. 1983. The ecology of a Late Triassic reptile assemblage from Gloucestershire, England. Palaeogeog. Palaeoclimat. Palaeolecol. 42: 341-365.
55. Fraser, N.C. & Walkden, G.M. 1984. The postcranial skeleton of the Upper Triassic sphenodontid Planocephalosaurus robinsonae. Palaeontology 24: 575-595.
56. Freeman, P.W. & Lemen, C. 2006. Puncturing ability of idealized canine teeth: edged and non-edged shanks. J. Zool. 269: 51-56.
57. Garland, T. Jr, Bennett, A.F. & Rezende, E.L. 2005. Phylogenetic approaches in comparative physiology. J. Exp. Biol. 208: 3015-3035.
58. Günther, A. 1867. Contribution to the anatomy of Hatteria (Rhynchocephalus, Owen). Philos. Trans. R. Soc. Lond. B 157: 1-34.
59. Harmon, L.J., Schulte, J.A., Losos, J.B. & Larson, A. 2003. Tempo and mode of evolutionary radiation in iguanian lizards. Science 301: 961-964.
60. Harmon, L.J., Weir, J., Brock, C., Glor, R.E. & Challenger, W. 2008. GEIGER: investigating evolutionary radiations. Bioinformatics 24: 129-131.
61. Hay, J.M., Sarre, S.D., Lambert, D.M., Allendorf, F.W. & Daughery, C.H. 2010. Genetic diversity and taxonomy: a reassessment of species designation in tuatara (Sphenodon: Reptilia). Conserv. Genet. 11: 1063-1081.
62. Heckert, A.B. 2004. Late Triassic microvertebrates from the lower Chinle Group (Otischalkian-Adamanian: Carnian) southwestern USA. N. Mex. Mus. Nat. Hist. Sci. Bull. 27: 1-170.
63. Herrel, A., Schaerlaeken, V., Moravec, J. & Ross, C.F. 2009. Sexual shape dimorphism in tuatara. Copeia 2009: 727-731.
64. Ivanović, A., Sotiropoulos, K., Üzüm, M., Olgun, K., Cogǎlniceanu, D. & Kalezić, M.L. 2011. A phylogenetic view on skull size and shape variation in the smooth newt (Lissotriton vulgaris, Caudata, Salamandridae). J. Zool. Syst. Evol. Res. 50: 116-124.
65. Jones, M.E.H. 2004. Skull evolution in Rhynchocephalia. J. Vertebr. Palaeontol. 24S: 76-77.
66. Jones, M.E.H. 2006a. Tooth diversity and function in the Rhynchocephalia (Diapsida: Lepidosauria). In: Ninth International Symposium on Mesozoic Terrestrial Ecosystems and Biota (P.M. Barrett & S.E. Evans, eds), pp. 55-58. Natural History Museum, London, UK.
67. Jones, M.E.H. 2006b. The Early Jurassic clevosaurs from China (Diapsida: Lepidosauria). In: The Triassic/Jurassic Terrestrial Transition (J.D. Harris, S. Lucas, J. Kirkland & A.R.C. Milner eds). N. Mex. Mus. Nat. Hist. Sci. Bull. 37: 548-562.
68. Jones, M.E.H. 2008. Skull shape and feeding strategy in Sphenodon and other Rhynchocephalia (Diapsida: Lepidosauria). J. Morphol. 269: 945-966.
69. Jones, M.E.H. 2009. Dentary tooth shape in Sphenodon and its fossil relatives (Diapsida: Lepidosauria: Rhynchocephalia). Front. Oral Biol. 13: 9-15.
70. Jones, M.E.H. & Lappin, A.K. 2009. Bite-force performance of the last rhynchocephalian (Lepidosauria: Sphenodon). J. R. Soc. N. Z. 39: 71-83.
71. Jones, M.E.H., Curtis, N., O'Higgins, P., Fagan, M. & Evans, S.E. 2009a. Head and neck muscles associated with feeding in Sphenodon (Reptilia: Lepidsauria: Rhynchocephalia). Palaeontologia Electr. 12: 1-56.
72. Jones, M.E.H., Tennyson, A.J.D., Worthy, J.P., Evans, S.E. & Worthy, T.H. 2009b. A sphenodontine (Rhynchocephalia) from the Miocene of New Zealand and palaeobiogeography of the tuatara (Sphenodon). Proc. R. Soc. Lond. B Biol. Sci. 276: 1385-1390.
73. Jones, M.E.H., Curtis, N., Fagan, M.J., O'Higgins, P. & Evans, S.E. 2011. Hard tissue anatomy of the cranial joints in Sphenodon (Rhynchocephalia): sutures, kinesis, and skull mechanics. Palaeontologia Electr. 14: 1-92.
74. Jones, M.E.H., O'Higgins, P., Fagan, M., Evans, S.E. & Curtis, N. 2012. Shearing mechanics and the influence of a flexible symphysis during oral food processing in Sphenodon (Lepidosauria: Rhynchocephalia). Anat. Rec. 295: 1075-1091.
75. Kirkland, J.I. 2006. Fruita Paleontological Area (Upper Jurassic, Morrison Formation), western Colorado: an example of terrestrial taphofacies analysis. In: Paleontology and Geology of the Upper Jurassic Morrison Formation (J.R. Foster & S.G. Lucas, eds), pp. 67-95. Museum of Natural History & Science, New Mexico, Bulletin 36.
76. Klingenberg, C.P. 2011. morphoj: an integrated software package for geometric morphometrics. Mol. Ecol. Resour. 11: 353-357.
77. Klingenberg, C.P. & Gidazwiski, N.A. 2010. Testing and quantifying phylogenetic signals and homoplasy in morphometric data. Syst. Biol. 59: 245-261.
78. Losos, J.B. 2011a. Seeing the forest for the trees: the limitations of phylogenies in comparative biology. Am. Nat. 177: 709-727.
79. Losos, J.B. 2011b. Convergence, adaptation, and constraint. Evolution 65: 1827-1840.
80. Lucas, P.W. & Luke, D.A. 1984. Chewing it over: basic principles of food breakdown. In: Food Acquisition and Processing in Primates (D.K. Chivers, B.A. Wood & A. Bilsborough, eds), pp. 283-301. Plenum Press, New York.
81. Maddison, W.P. 1991. Squared-change parsimony reconstructions of ancestral states for continuous-valued characters on a phylogenetic tree. Syst. Zool. 40: 304-314.
82. Martin, T. & Krebs, B. 2000. Guimarota: A Jurassic Ecosystem. Dr Friedrich Pfeil, Munich.
83. Meloro, C. 2011. Morphological disparity in Plio-Pleistocene large carnivore guilds from Italian peninsula. Acta Palaeontol. Pol. 56: 33-44.
84. Meloro, C. 2012. Mandibular shape correlates of tooth fracture in extant Carnivora. Implications to infer feeding behaviour of Rancho La Brea predators. Biol. J. Linn. Soc. 106: 70-80.
85. Meloro, C. & Raia, P. 2010. Cats and dogs down the tree: the tempo and mode of evolution in the lower carnassial of fossil and living Carnivora. Evol. Biol. 37: 177-186.
86. Meloro, C., Raia, P., Piras, P., Barbera, C. & O'Higgins, P. 2008. The shape of the mandibular corpus in large fissiped carnivores: allometry, function and phylogeny. Zool. J. Linn. Soc. 154: 832-845.
87. Meloro, C., Raia, P., Carotenuto, F. & Cobb, S.N. 2011. Phylogenetic signal, function and integration in the subunits of the carnivoran mandible. Evol. Biol. 38: 465-475.
88. Metzger, K.A. & Herrel, A. 2005. Correlations between lizard cranial shape and diet: a quantitative, phylogenetically informed analysis. Biol. J. Linn. Soc. 86: 433-466.
89. Milner, A.C., Milner, A.R. & Evans, S.E. 2000. Global changes and biota: amphibians, reptiles and birds. In: Biotic Response to Global Change: The Last 145 Million Years (S. Culver & P. Rawson, eds), pp. 316-332. Cambridge University Press, Cambridge.
90. Nelson, N.J., Keall, S.N., Brown, D. & Daugherty, C.H. 2002. Establishing a new wild population of tuatara (Sphenodon guntheri). Cons. Biol. 16: 887-894.
91. O'Higgins, P. & Jones, N. 1998. Facial growth in Cercocebus torquatus: an application of three dimensional geometric morphometric techniques to the study of morphological variation. J. Anat. 193: 251-272.
92. Pierce, S.E., Angielczyk, K.D. & Rayfield, E.J. 2009. Morphospace occupation in thalattosuchian crocodylomorphs: skull shape variation, species delineation, and temporal patterns. Palaeontology 52: 1057-1097.
93. Piras, P., Marcolini, F., Claude, J., Ventura, J., Kotsakis, T. & Cubo, J. 2012. Ecological and functional correlates of molar shape variation in European populations of Arvicola (Arvicolinae, Rodentia). Zool. Anz. http:dx.doi.org/10.1016/j.jcz.2011.12.002.
94. Rasmussen, T.E. & Callison, G. 1981. A new herbivorous sphenodontid (Rhynchocephalia: Reptilia) from the Jurassic of Colorado. J. Paleontol. 55: 1109-1116.
95. Reilly, S.M. & McBrayer, L.D. 2007. Prey capture and prey processing behaviour and the evolution of lingual and sensory characteristics: divergences and convergences in lizard feeding biology. In: Lizard Ecology. The Evolutionary Consequences of Foraging Mode (S.M. Reilly, L.D. McBrayer & D.B. Miles, eds), pp. 302-333. Cambridge University Press, UK.
96. Reynoso, V.H. 1996. A Middle Jurassic Sphenodon-like sphenodontian (Diapsida: Lepidosauria) from Huizachal Canyon, Tamaulipas, Mexico. J. Vertebr. Paleontol. 16: 210-221.
97. Reynoso, V.H. 2000. An unusual aquatic sphenodontian (Reptilia: Diapsida) from the Tlayua Formation (Albian), central Mexico. J. Paleontol. 74: 133-148.
98. Reynoso, V.H. 2005. Possible evidence of a venom apparatus in a Middle Jurassic sphenodontian from the Huizachal red beds of Tamaulipas, México. J. Vertebr. Paleontol. 25: 646-654.
99. Reynoso, V.H. & Clark, J.M. 1998. A dwarf sphenodontian from the Jurassic la Boca Formation of Tamaulipas, Mexico. J. Vertebr. Paleontol. 18: 333-339.
100. Robb, J. 1977. The Tuatara. Meadowfield Press Limited, Durham.
101. Robinson, P.L. 1962. Gliding reptiles from the Upper Keuper of Great Britain. Proc. Geol. Soc. 1601: 137-146.
102. Robinson, P.L. 1976. How Sphenodon and Uromastix grow their teeth and use them. In: Morphology and Biology of the Reptiles (A.d'.A. Bellairs & C.B. Cox, eds), pp. 43-64. Academic Press, London.
103. Rohlf, F.J. 2001. Comparative methods for the analysis of continuous variables: geometric interpretations. Evolution 55: 2143-2160.
104. Rohlf, F.J. 2002. Geometric morphometrics in phylogeny. In: Morphology, Shape and Phylogenetics (P. Forey & N. Macleod, eds), pp. 175-193. Francis & Taylor, London.
105. Rohlf, F.J. 2006a. tpsdig 2.10. Department of Ecology and Evolution. State University of New York, Stony Brook, New York.
106. Rohlf, F.J. 2006b. tpspls 1.18. Department of Ecology and Evolution. State University of New York, Stony Brook, New York.
107. Rohlf, F.J. 2006c. A comment on phylogenetic correction. Evolution 60: 1509-1515.
108. Rohlf, F.J. 2007. tpstree 1.21. Department of Ecology and Evolution. State University of New York, Stony Brook, New York.
109. Rohlf, F.J. 2008. tpsrelw 1.46. Department of Ecology and Evolution. State University of New York, Stony Brook, New York.
110. Rohlf, F.J. & Corti, M. 2000. Use of two-block partial least squares to study covariation in shape. Syst. Biol. 49: 740-753.
111. Rohlf, F.J. & Slice, D.E. 1990. Extensions of the procrustes method for the optimal superimposition of landmarks. Syst. Zool. 39: 40-59.
112. Ruta, M., Wagner, P.J. & Coates, I.M. 2006. Evolutionary patterns in early tetrapods. I. Rapid initial diversification followed by decrease in rates of character change. Proc. R. Soc. Lond. B Biol. Sci. 273: 2107-2111.
113. Slater, G.J., Price, S.A., Santini, F. & Alfaro, M.A. 2010. Diversity vs disparity and the evolution of modern cetaceans. Proc. R. Soc. Lond. B Biol. Sci. 277: 3097-3104.
114. Stayton, C.T. 2005. Morphological evolution of the lizard skull: a geometric morphometrics survey. J. Morph. 263: 47-59.
115. Stayton, C.T. 2006. Testing hypotheses of convergence with multivariate data: morphological and functional convergence among herbivorous lizards. Evolution 60: 824-841.
116. Stayton, C.T. & Ruta, M. 2006. Geometric morphometrics of the skull roof of stereospondyls (Amphibia: Temnospondyli). Palaeontology 49: 307-337.
117. Stein, K., Palmer, C., Gill, P.G. & Benton, M.J. 2008. The aerodynamics of the British late Triassic Kuehneosauridae. Palaeontology 51: 967-981.
118. Subramanian, S., Hay, J.M., Mohandesan, E., Millar, C.D. & Lambert, D.M. 2009. Molecular and morphological evolution in tuatara are decoupled. Trends Genet. 25: 16-18.
119. Sues, H.-D., Shubin, N.H. & Olsen, P.E. 1994. A new sphenodontian (Lepidosauria: Rhynchocephalia) from the McCoy Brook Formation (Lower Jurassic) of Nova Scotia. Canad. J. Vertebr. Paleontol. 14: 327-340.
120. Taylor, M.A. 1987. How tetrapods feed in the water: a functional analysis by paradigm. Zool. J. Linn. Soc. 91: 171-195.
121. Throckmorton, G.S., Hopson, J.A. & Parks, P. 1981. A redescription of Toxolophosaurus cloudi Olson, a Lower Cretaceous herbivorous sphenodontian reptile. J. Paleontol. 55: 586-597.
122. Towns, D.R., Daugherty, C.H. & Cree, A. 2001. Raising the prospects for a forgotten fauna: a review of 10 years of conservation effort for New Zealand reptiles. Biol. Conserv. 99: 3-16.
123. Towns, D.R., Parrish, G.R., Tyrrell, C.L., Ussher, G.T., Cree, A., Newman, D.G. et al. 2007. Responses of tuatara (Sphenodon punctatus) to removal of introduced Pacific rats from Islands. Cons. Biol. 21: 1021-1031.
124. Waldman, M. & Evans, S.E. 1994. Lepidosauromorph reptiles from the Middle Jurassic of Skye. Zool. J. Linn. Soc. 112: 135-150.
125. Walls, G.Y. 1981. Feeding ecology of the tuatara (Sphenodon punctatus) on Stephens Island, Cook Strait. N. Z. J. Ecol. 4: 89-97.
126. Whiteside, D.I. 1986. The head skeleton of the Rhaetian sphenodontid Diphydontosaurus avonis gen. et sp. nov., and the modernising of a living fossil. Philos. Trans. R. Soc. Lond. B 312: 379-430.
127. Wills, M.A., Briggs, D.E.G. & Fortey, R.A. 1994. Disparity as an evolutionary index: a comparison of Cambrian and Recent arthropods. Paleobiology 20: 93-130.
128. Worthy, T.H. & Grant-Mackie, J.A. 2003. Late-Pleistocene avifaunas from Cape Wanbrow, Otago, South Island, New Zealand. J. R. Soc. N. Z. 33: 427-485.
129. Young, M.T., Brusatte, S.L., Ruta, M. & Andrade, M.B. 2010. The evolution of Metriorhynchoidea (Mesoeucrocodylia: Thalattosuchia): an integrated approach using geometric morphometrics, analysis of disparity and biomechanics. Zool. J. Linn. Soc. 158: 801-859.
130. Young, M.T., Bell, M.A. & Brusatte, S.L. 2011. Craniofacial form and function in Metriorhynchidae (Crocodylomorpha: Thalattosuchia): modelling phenotypic evolution with maximum likelihood methods. Biol. Lett. 7: 913-916.
131. Zelditch, M.L., Swiderski, D.L., Sheets, H.D. & Fink, W.L. 2004. Geometric Morphometrics for Biologists. A Primer. Elsevier, USA.