Professor Richard Butler PhD

Professor Richard Butler

School of Geography, Earth and Environmental Sciences
Professor of Palaeobiology

Contact details

School of Geography, Earth and Environmental Sciences
University of Birmingham
B15 2TT

Richard is a vertebrate palaeontologist with expertise in the systematics, evolution and biogeography of late Palaeozoic to Mesozoic reptiles. His current research programme aims to establish major patterns and drivers of terrestrial biodiversity change, as exemplified by four-limbed vertebrates (tetrapods), over the last 375 million years. He is also interested in the origin and dramatic evolutionary radiation of dinosaurs and closely related fossil groups in the aftermath of the Permo-Triassic mass extinction, the largest extinction event in the history of life on Earth.


2007 – PhD, University of Cambridge
2002 – BSc Geology, University of Bristol


  • 2017–. Professor of Palaeobiology, School of Geography, Earth and Environmental Sciences (GEES), University of Birmingham.
  • 2015–2017: Senior Birmingham Fellow, GEES, University of Birmingham.
  • 2013–2015: Birmingham Fellow, GEES, University of Birmingham.
  • 2011–2013. Junior Research Group Leader (DFG Emmy Noether Programme), GeoBio-Center, Ludwig-Maximilians-Universität München, Munich, Germany.
  • 2009–2011. Alexander von Humboldt Foundation Research Fellowship, Bayerische Staatssammlung für Paläontologie und Geologie, Munich, Germany.
  • 2008–2009. NERC Researcher Co-Investigator, Department of Palaeontology, Natural History Museum.
  • 2006–2008. NERC Research Assistant, Department of Palaeontology, Natural History Museum, London.


  • 2016. Award for Excellence in Doctoral Research Supervision, University of Birmingham Graduate School (overall winner for the College of Life & Environmental Sciences)
  • 2011. Palaeontological Association Hodson Award (annual award to a palaeontologist under the age of 35 who has made a notable early contribution to the science)
  • 2005. Society of Vertebrate Paleontology Predoctoral Fellowship (annual award intended to promote a professional career in vertebrate palaeontology)


Richard supervises major/advanced projects, and contributes to teaching in the following areas:

  • Advanced projects
  • Evolution of vertebrates
  • Field skills

Postgraduate supervision

Richard leads a highly active research group including postdoctoral, PhD, and MSci researchers. 

Postdoctoral Researchers

  • Dr Roger Close (European Research Council)
  • Dr Terri Cleary (European Research Council)
  • Dr Gemma Benevento (European Research Council)
  • Dr Christopher Dean (European Research Council)

Current PhD students

  • Emma Dunne (2015–2019, European Research Council) 
  • Daniel Cashmore (2015–2019, European Research Council)
  • Amy Jones (co-supervisor; 2016–2020, NERC CENTA) 
  • Romy Rayner (2018–2022, self)
  • Luke Meade (co-supervisor; 2018–2022, NERC CENTA)
  • Fion Ma (co-supervisor; 2018–2022, self)
  • Lisa Schnetz (co-supervisor; 2018–2022, NERC CENTA)
  • Struan Henderson (co-supervisor; 2018–2022, Royal Society)
  • Emily Brown (2019–2023; NERC CENTA)

External PhD students

  • Jordan Bestwick (University of Leicester)

Research group alumni

  • Dr Pedro Godoy (PhD student, 2014–2018, now a postdoctoral fellow at Stony Brook University, New York)
  • Dr Andrew Jones (PhD student, 2014–2018, now Digital Technologies Officer at the Lapworth Museum of Geology)
  • Dr Julio Marsola (visiting PhD student, 2016–2017)
  • Juan Benito (Research Assistant, 2015–2017; now a PhD student at University of Bath)
  • Dr David Button (Research Fellow, 2015–2016; now a postdoc at the Natural History Museum, London)
  • Dr Felipe Montefeltro (UoB Brazilian Visiting Fellow, 2014; now Associate Professor at São Paulo State University, Ilha Solteira)
  • Dr Martin Ezcurra (PhD student, 2012–2015; now a permanent CONICET-funded researcher, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires)
  • Dr Roland Sookias (PhD student, 2012–2015; now a Humboldt Postdoctoral Fellow at the Museum für Naturkunde, Berlin)


Research interests

  • Systematics, taxonomy, and anatomy of archosaurs, particularly dinosaurs, and closely related groups
  • Terrestrial recovery from the Permo-Triassic mass extinction event
  • Phanerozoic and Mesozoic diversification patterns among vertebrates, and fossil record quality
  • Late Palaeozoic–Mesozoic vertebrate biogeography
  • Body size evolution and its drivers in deep time
  • Early evolution of the avian respiratory system, and lung ventilation among fossil archosaurs

Personal research group webpage

Key research grants

2018–2020. Royal Society International Exchange Grant (Russia): "The rise of reptilian apex predators following the Permo-Triassic mass extinction" (Principal Investigator)

2015–2020. ERC Starting Grant: "TERRA. 375 Million Years of the Diversification of Life on Land: Shifting the Paradigm?" (Principal Investigator) 

2014-2018. Marie Curie Career Integration Grant: "The early Mesozoic rise of archosaurs: New insights into an exemplar evolutionary radiation" (Principal Investigator)

2011-2014. German Research Foundation (DFG) Emmy Noether Programme: “Dawn of the dinosaurs: Archosauromorph evolution in the terrestrial Triassic” (Principal Investigator)

2010-2011. German Research Foundation (DFG) Research Grant: “Terrestrial vertebrates from near the Triassic-Jurassic boundary in Portugal: Excavation, geological context, and faunal change” (Principal Investigator)

2008-2009. NERC Small Grant: “Origin of the avian respiratory system: A CT-study of postcranial pneumaticity in basal archosaurs” (Researcher Co-Investigator)

Richard has also received additional small grant funding (each < £5000) from the Rutherford Fund, Royal Society, Synthesys, Lamont-Doherty Earth Observatory, Jurassic Foundation, Palaeontological Association, Systematics Association, and the Percy Sladen Memorial Fund, among others.

Other activities

  • 2018–. REF lead, Earth Systems and Environmental Sciences, GEES, University of Birmingham
  • 2014–. Honorary Senior Research Fellow, Evolutionary Studies Institute, University of the Witwatersrand, Johannesburg, South Africa
  • 2014–2018. Academic Keeper, Lapworth Museum of Geology, University of Birmingham
  • 2015–2018. Chair of Steering Group, Symposium on Vertebrate Palaeontology and Comparative Anatomy (SVPCA)
  • 2012–2017. Program Committee, Society of Vertebrate Paleontology
  • 2014–2016. Member of Council, the Palaeontological Association
  • 2013–2017. Executive Committee, the Paleobiology Database
  • 2013–2018. Editorial Board, Journal of Vertebrate Paleontology
  • 2013–. Editorial Board, Acta Palaeontologica Polonica
  • 2012–2014. Editorial Board, PLOS ONE
  • 2010–2013. Associate Editor, Paläontologische Zeitschrift
  • 2008–2011. Section Editor, Fossil Reptiles, Zootaxa



118. Cashmore DD, Butler RJ. In press. Skeletal completeness of the non-avian theropod dinosaur fossil record. Palaeontology.

​117. Kammerer CF, Viglietti PA, Hancox J, Butler RJ, Choiniere JN. In press. A new kannemeyeriiform dicynodont (Ufudocyclops mukanelai gen. et sp. nov.) from Subzone C of the Cynognathus Assemblage Zone (Triassic of South Africa) with implications for biostratigraphic correlation with other African Triassic faunas. Journal of Vertebrate Paleontology.

​116. Lehmann OER, Ezcurra MD, Butler RJ, Lloyd GT. In press. Biases in disparity analyses with the generalized Euclidean distance. Palaeontology.

115. Butler RJ, Jones AS, Buffetaut E, Mandl G, Scheyer T, Schultz O. 2019. Description and phylogenetic placement of a new marine species of phytosaur (Archosauriformes: Phytosauria) from the Late Triassic of Austria. Zoological Journal of the Linnean Society.

114. Brown EE, Cashmore DD, Simmons NB, Butler RJ. 2019 Quantifying the completeness of the bat fossil record. Palaeontology.

113. Butler RJ, Ezcurra MD, Liu J, Sookias RB, Sullivan C. 2019. The anatomy and phylogenetic position of the erythrosuchid archosauriform Guchengosuchus shiguaiensis from the earliest Middle Triassic of China. PeerJ 7:e6435.

112. Close RA, Benson RBJ, Alroy J, Behrensmeyer AK, Benito J, Carrano MT, Cleary TJ, Dunne EM, Mannion PD, Uhen MD, Butler RJ. 2019. Diversity dynamics of Phanerozoic terrestrial tetrapods at the local-community scale. Nature Ecology & Evolution.

111. Marsola JCA, Bittencourt JS, Butler RJ, Da Rosa AAS, Sayão JM, Langer MC. 2019. A new dinosaur with theropod affinities from the Late Triassic Santa Maria Formation, South Brazil. Journal of Vertebrate Paleontology e1531878.

110. Marsola JCA, Ferreira GS, Langer MC, Button DJ, Butler RJ. 2019. Increases in sampling support the southern Gondwanan hypothesis for the origin of dinosaurs. Palaeontology.

109. Ezcurra MD, Gower DJ, Sennikov A, Butler RJ. 2019. The osteology of the holotype of the early erythrosuchid Garjainia prima Ochev, 1958 (Diapsida: Archosauromorpha) from the upper Lower Triassic of European Russia. Zoological Journal of the Linnean Society 185:717–783.

108. Hedge J, Shillito AP, Davies NS, Butler RJ, Sansom IJ. 2019. Invertebrate trace fossils from the Alveley Member, Salop Formation (Pennsylvanian, Carboniferous), Shropshire, UK. Proceedings of the Geologists’ Association 130:103–111.


107. Jones AS, Butler RJ. 2018. A new phylogenetic analysis of Phytosauria (Archosauria: Pseudosuchia) with the application of continuous and geometric morphometric character coding. PeerJ 6:e5901.

106. Bestwick J, Unwin DM, Butler RJ, Henderson DM, Purnell MA. 2018. Pterosaur dietary hypotheses: a review of ideas and approaches. Biological Reviews 93:2021–2048.

105. Ezcurra MD, Butler RJ. 2018. The rise of the ruling reptiles and ecosystem recovery from the Permian-Triassic mass extinction. Proceedings of the Royal Society B 285:20180361.

104. Nesbitt SJ, Butler RJ, Ezcurra MD, Charig AJ, Barrett PM. 2018. The anatomy of Teleocrater rhadinus, an early avemetatarsalian from the lower portion of the Lifua Member of the Manda Beds (Middle Triassic); pp. 142–177 in C. A. Sidor, S. J. Nesbitt (eds.), Vertebrate and Climatic Evolution in the Triassic Rift Basins of Tanzania and Zambia. Society of Vertebrate Paleontology Memoir 17. Journal of Vertebrate Paleontology 37 (6, Supplement).

103. Butler RJ, Nesbitt SJ, Charig AJ, Gower DJ, Barrett PM. 2018. Mandasuchus tanyauchen, gen. et sp. nov., a pseudosuchian archosaur from the Manda Beds (?Middle Triassic) of Tanzania; pp. 96–121 in C. A. Sidor, S. J. Nesbitt (eds.), Vertebrate and Climatic Evolution in the Triassic Rift Basins of Tanzania and Zambia. Society of Vertebrate Paleontology Memoir 17. Journal of Vertebrate Paleontology 37 (6, Supplement).

102. Panciroli E, Benson RBJ, Butler RJ. 2018. New partial dentaries of amphitheriid mammalian Palaeoxonodon ooliticus from Scotland, and posterior dentary morphology in early cladotherians. Acta Palaeontologica Polonica 63:197–206.

101. Godoy PL, Ferreira GS, Montefeltro FC, Vila Nova BC, Butler RJ, Langer MC. 2018. Evidence for heterochrony in the cranial evolution of fossil crocodyliforms. Palaeontology 61:543–558.

100. Close RA, Alroy J, Evers SW, Butler RJ. 2018. How should we estimate diversity in the fossil record? Testing richness estimators using sampling-standardised discovery curves. Methods in Ecology & Evolution 9:1386–1400.

99. Dunne EM, Close RA, Button DJ, Brocklehurst N, Cashmore DD, Lloyd GT, Butler RJ. 2018. Diversity change during the rise of tetrapods and the impact of the ‘Carboniferous Rainforest Collapse’. Proceedings of the Royal Society B 285:20172730.


98. Tutin S, Butler RJ. 2017. The completeness of the fossil record of plesiosaurs, marine reptiles from the Mesozoic. Acta Palaeontologica Polonica 62:563–573.

97. Button DJ, Lloyd GT, Ezcurra MD, Butler RJ. 2017. Mass extinctions drove global faunal cosmopolitanism on the supercontinent Pangaea. Nature Communications 8:733.

96. Close RA, Benson RBJ, Upchurch P, Butler RJ. 2017. Controlling for the species-area effect supports constrained long-term Mesozoic terrestrial vertebrate diversification. Nature Communications 8:15381.

95. Nesbitt SJ, Butler RJ, Ezcurra MD, Barrett PM, Stocker MR, Angielczyk KD, Smith RMH, Sidor CA, Niedźwiedzki G, Sennikov A, Charig AJ.2017.The earliest bird-line archosaurs and the assembly of the dinosaur body plan. Nature 544:484–487.


94. Meade LE, Jones AS, Butler RJ. 2016. A revision of tetrapod footprints from the late Carboniferous of the West Midlands, UK. PeerJ 4:e2718. 

93. McPhee BW, Upchurch P, Mannion PD, Sullivan C, Butler RJ, Barrett PM. 2016. A revision of Sanpasaurus yaoi Young, 1944 from the Early Jurassic of China, and its relevance to the early evolution of Sauropoda (Dinosauria). PeerJ 4:e2578

92. Foth C, Ezcurra M, Sookias RB, Brusatte SL, Butler RJ. 2016. Unappreciated diversification of stem archosaurs during the Middle Triassic predated the dominance of dinosaurs. BMC Evolutionary Biology 16:188. 

91. Lautenschlager S, Butler RJ. 2016. Neural and endocranial anatomy of Triassic phytosaurian reptiles and convergence with fossil and modern crocodylians. PeerJ 4:e2251.

90. Sobral G, Sookias RB, Bhullar B-AS, Smith RMH, Butler RJ, Müller J. 2016. New information on the braincase and inner ear of Euparkeria capensis Broom: implications for diapsid and archosaur evolution. Royal Society Open Science 3:160072.

89. Barrett PM, Butler RJ, Yates AM, Baron MG, Choniere JN. New specimens of the basal ornithischian dinosaur Lesothosaurus diagnosticus Galton, 1978 from the Early Jurassic of South Africa. Palaeontologia Africana 50:48–63. 

88. Pinheiro FL, De França MAG, Lacerda MB, Butler RJ, Schultz CL. 2016. An exceptional fossil skull from South America illuminates the origins of the archosauriform radiation. Scientific Reports 6:22817.  

87. Dean CD, Mannion PD, Butler RJ. 2016. Preservational bias controls the fossil record of pterosaurs. Palaeontology 59:225–247. 

86. Benson RBJ, Butler RJ, Alroy J, Mannion PD, Carrano MT, Lloyd GT. 2016. Near-stasis in the long-term diversification of Mesozoic tetrapods. PLoS Biology 14:e1002359. 

85. Kammerer, CF, Butler RJ, Bandyopadhyay S, Stocker MR. 2016. Relationships of the Indian phytosaur Parasuchus hislopi Lydekker, 1885. Papers in Palaeontology 2:1–23.  

84. Ezcurra MD, Montefeltro F, Butler RJ. 2016. The early evolution of rhynchosaurs. Frontiers in Ecology and Evolution 3:142. 


83. Mannion PD, Benson RBJ, Carrano MT, Tennant JP, Judd J, Butler RJ. 2015. Climate constrains the evolutionary history and biodiversity of crocodylians. Nature Communications 6:8438. 

82. Liu J, Butler RJ, Sullivan C, Ezcurra MD. 2015. “Chasmatosaurus ultimus”, a putative proterosuchid archosauriform from the Middle Triassic, is an indeterminate crown archosaur. Journal of Vertebrate Paleontology 35:e965779. 

81. Upchurch P, Andres B, Butler RJ, Barrett PM. An analysis of pterosaurian biogeography: implications for the evolutionary history and fossil record quality of the first flying vertebrates. Historical Biology 27:697–717. 

80. Butler RJ, Ezcurra MD, Montefeltro FC, Samathi A, Sobral G. 2015. A new species of basal rhynchosaur (Diapsida: Archosauromorpha) from the early Middle Triassic of South Africa, and the early evolution of Rhynchosauria. Zoological Journal of the Linnean Society 174:571–588. 

79. Brusatte SL, Butler RJ, Mateus O, Steyer JS. 2015. A new species of Metoposaurus from the Late Triassic of Portugal and comments on the systematics and biogeography of metoposaurid temnospondyls. Journal of Vertebrate Paleontology 35:e912988. 

78. Ezcurra MD, Butler RJ. 2015. Post-hatchling cranial ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusiJournal of Anatomy 226:387–402. 

77. Brusatte SL, Butler RJ, Barrett PM, Carrano MT, Evans DC, Lloyd GT, Mannion PD, Norell MA, Peppe DJ, Upchurch P, Williamson TE. 2015. The extinction of the dinosaurs. Biological Reviews 90:628–642. 

76. Ezcurra MD, Butler RJ. Taxonomy of the proterosuchid archosauriforms (Diapsida: Archosauromorpha) from the earliest Triassic of South Africa, and implications for the early archosauriform radiation. Palaeontology 58:141–170. 


75. Sookias RB, Sullivan C, Liu J, Butler RJ. 2014. Systematics of putative euparkeriids (Diapsida: Archosauriformes) from the Triassic of China. PeerJ 2:e658. 

74. Sookias RB, Sennikov AG, Gower DJ, Butler RJ.  2014. The monophyly of Euparkeriidae (Reptilia: Archosauriformes) and the origins of crown Archosauria: a revision of Dorosuchus neoetus from the Middle Triassic of Russia. Palaeontology 57:1177–1202.  

73. Niedźwiedzki G, Brusatte SL, Sulej T, Butler RJ. 2014. Basal dinosauriform and theropod dinosaurs from the middle-late Norian (Late Triassic) of Poland: implications for Triassic dinosaur evolution and distribution. Palaeontology 57:1121–1142. 

72. Gower DJ, Hancox PJ, Botha-Brink J, Sennikov AG, Butler RJ. A new species of Garjainia Ochev, 1958 (Diapsida: Archosauriformes: Erythrosuchidae) from the Early Triassic of South Africa. PLoS ONE 9(11): e111154. 

71. Barrett PM, Butler RJ, Mundil R, Scheyer TM, Irmis RB, Sánchez-Villagra MR. 2014. A palaeoequatorial ornithischian and new constraints on early dinosaur diversification. Proceedings of the Royal Society B 281: 20141147.  

70. Mateus O, Butler RJ, Brusatte SL, Steyer JS, Whiteside JH. 2014. The first phytosaur (Diapsida, Archosauriformes) from the Late Triassic of the Iberian Peninsula. Journal of Vertebrate Paleontology 34:970-975. 

69. Butler RJ, Sullivan C, Ezcurra MD, Liu J, Lecuona A, Sookias RB. 2014. New clade of enigmatic early archosaurs yields insights into early pseudosuchian phylogeny and the biogeography of the archosaur radiation. BMC Evolutionary Biology 14:128.

68. Benson RBJ, Frigot RA, Goswami A, Andres B, Butler RJ. 2014. Competition and constraint drove Cope's rule in the evolution of giant flying reptiles. Nature Communications 5:3567.

67. Ezcurra MD, Scheyer T, Butler RJ. 2014. The origin and early evolution of Sauria: reassessing the Permian saurian fossil record and the timing of the crocodile-lizard divergence. PLOS ONE 9:e89165.

66. Butler RJ, Rauhut OWM, Stocker MR, Bronowicz R. 2014. Redescription of the phytosaurs Paleorhinus ("Francosuchus") angustifrons and Ebrachosuchus neukami from Germany, with implications for Late Triassic biochronology. Zoological Journal of the Linnean Society 170:155–208. 


65. Nesbitt SJ, Butler RJ, Gower DJ. 2013. A new archosauriform (Reptilia: Diapsida) from the Manda beds (Middle Triassic) of southwestern Tanzania. PLOS ONE 8: e72753.

64. Niedźwiedzki G, Brusatte SL, Butler RJ. 2013. Prorotodactylus and Rotodactylus tracks: an ichnological record of dinosauromorphs from the Early–Middle Triassic of Poland. In: Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and their Kin, Nesbitt SJ, Desojo JB, Irmis RB (eds). Geological Society, London, Special Publications 379: 319-351.

63. Stocker MR, Butler RJ. Phytosauria. 2013. In: Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and their Kin, Nesbitt SJ, Desojo JB, Irmis RB (eds). Geological Society, London, Special Publications 379: 91-117.

62. Sookias RB, Butler RJ. Euparkeriidae. 2013. In: Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and their Kin, Nesbitt SJ, Desojo JB, Irmis RB (eds). Geological Society, London, Special Publications 379: 35-48.

61. Ezcurra MD, Butler RJ, Gower DJ. 2013. Proterosuchia: the origin and early history of Archosauriformes. In: Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and their Kin, Nesbitt SJ, Desojo JB, Irmis RB (eds). Geological Society, London, Special Publications 379: 9-33.

60. Butler RJ. 2013. “Francosuchus” trauthi is not Paleorhinus: implications for Late Triassic vertebrate biostratigraphy. Journal of Vertebrate Paleontology 33: 858–864.

59. Toljagić O, Butler RJ. 2013. The Triassic/Jurassic mass extinction as trigger for the Mesozoic radiation of crocodylomorphs. Biology Letters 9: 20130095.

58. Benson RBJ, Mannion PD, Butler RJ, Upchurch P, Goswami A, Evans SE. 2013. Cretaceous tetrapod fossil record sampling and faunal turnover: implications for biogeography and the rise of modern clades. Palaeogeography, Palaeoclimatology, Palaeoecology 372: 88–107.

57. Butler RJ, Benson RBJ, Barrett PM. 2013. Pterosaur diversity: untangling the influences of sampling biases, Lagerstätten, and genuine biodiversity signals. Palaeogeography, Palaeoclimatology, Palaeoecology 372: 78–87.

56. Nesbitt SJ, Butler RJ. 2013. Redescription of the archosaur Parringtonia gracilis from the Middle Triassic Manda Beds of Tanzania, and the antiquity of Erpetosuchidae. Geological Magazine 150: 225–238.

55. Butler RJ, Yates AM, Rauhut OWM, Foth C. A pathological tail in a basal sauropodomorph dinosaur from South Africa: evidence of traumatic amputation? Journal of Vertebrate Paleontology 33: 224–228.

54. Brusatte SL, Butler RJ, Niedźwiedzki G, Sulej T, Bronowicz R, Satkunas J. 2013. First record of Mesozoic terrestrial vertebrates from Lithuania: phytosaurs (Diapsida: Archosauriformes) of probable Late Triassic age, with a review of phytosaur biogeography. Geological Magazine 150: 110–121.


53. Han F-L, Barrett PM, Butler RJ, Xu X. 2012. Postcranial anatomy of Jeholosaurus shangyuanensis (Dinosauria, Ornithischia) from the Lower Cretaceous Yixian Formation of China. Journal of Vertebrate Paleontology 32: 1370–1395.

52. Ősi A, Prondvai E, Butler RJ, Weishampel DB. 2012. Phylogeny, histology and inferred body size evolution in a new rhabdodontid dinosaur from the Late Cretaceous of Hungary. PLOS ONE 7(9): e44318.

51. Mannion PD, Benson RBJ, Upchurch P, Butler RJ, Carrano MT, Barrett PM. 2012. A temperate palaeodiversity peak in Mesozoic dinosaurs and evidence for Late Cretaceous geographical partitioning. Global Ecology and Biogeography 21: 898–908.

50. Sookias RB, Benson RBJ, Butler RJ. 2012. Biology, not environment, drives major temporal patterns in terrestrial tetrapod body size. Biology Letters 8: 674–677.

49. Brusatte SL, Butler RJ, Prieto-Marquez A, Norell MA. 2012. Dinosaur morphological diversity and the end-Cretaceous extinction. Nature Communications 3: article 804.

48. Foth C, Brusatte SL, Butler RJ. 2012. Do different disparity proxies converge on a common signal? Insights from the cranial morphometrics and evolutionary history of Pterosauria (Diapsida: Archosauria). Journal of Evolutionary Biology 25: 904–915.

47. Sookias RB, Butler RJ, Benson RBJ. 2012. Rise of dinosaurs reveals major body size transitions are driven by passive processes of trait evolution. Proceedings of the Royal Society B 279: 2180–2187.

46. Butler RJ, Porro LB, Galton PM, Chiappe LM. 2012. Anatomy and cranial functional morphology of the small-bodied dinosaur Fruitadens haagarorum (Ornithischia: Heterodontosauridae) from the Upper Jurassic of the USA. PLOS ONE 7: e31556.

45. Butler RJ, Barrett PM, Gower DJ. 2012. Reassessment of the evidence for postcranial skeletal pneumaticity in Triassic archosaurs, and the early evolution of the avian respiratory system. PLOS ONE 7: e34094.

44. Butler RJ, Brusatte SL, Andres B, Benson RBJ. 2012. How do rock record biases affect studies of disparity in deep time? A case study of the Pterosauria (Reptilia: Archosauria). Evolution 66: 147–162.

43. Benson RBJ, Butler RJ, Carrano MT, O’Connor PM. 2012. Air-filled postcranial bones in theropod dinosaurs: physiological implications and the ‘reptile’-bird transition. Biological Reviews 87: 168­–193.


42. Butler RJ, Brusatte SL, Reich M, Nesbitt SJ, Schoch RR, Hornung JJ. 2011. The sail-backed reptile Ctenosauriscus from the latest Early Triassic of Germany and the timing and biogeography of the early archosaur radiation. PLOS ONE 6: e25693 (28 pages).

41. Barrett PM, Butler RJ, Twitchett RJ, Hutt S. 2011. New material of Valdosaurus canaliculatus (Ornithischia: Ornithopoda) from the Lower Cretaceous of southern England. Special Papers in Palaeontology 86: 131–163.

40. Upchurch P, Mannion PD, Benson RBJ, Butler RJ, Carrano MT. 2011. Geological and anthropogenic controls on the sampling of the terrestrial fossil record: a case study from the Dinosauria. In: Comparing the Geological and Fossil Records: Implications for Biodiversity Studies, McGowan AJ, Smith AB (eds). Geological Society, London, Special Publication 358: 209–240.

39. Benson RBJ, Butler RJ. 2011. Uncovering the diversification history of marine tetrapods: ecology influences the effect of geological sampling biases. In: Comparing the Geological and Fossil Records: Implications for Biodiversity Studies, McGowan AJ, Smith AB (eds). Geological Society, London, Special Publication 358: 191–207.

38. Norman DB, Crompton AW, Butler RJ, Porro LB, Charig AC. 2011. The Lower Jurassic ornithischian dinosaur Heterodontosaurus tucki Crompton and Charig, 1962: cranial anatomy, functional morphology, taxonomy and relationships. Zoological Journal of the Linnean Society 163: 182–276.

37. Butler RJ, Jin L, Chen J, Godefroit P. 2011. The postcranial osteology and phylogenetic position of the small ornithischian dinosaur Changchunsaurus parvus from the Quantou Formation (Cretaceous: Aptian-Cenomanian) of Jilin Province, northeastern China. Palaeontology 54: 667–683.

36. Butler RJ, Benson RBJ, Carrano MT, Mannion PD, Upchurch P. 2011. Sea-level, dinosaur diversity, and sampling biases: investigating the ‘common cause’ hypothesis in the terrestrial realm. Proceedings of the Royal Society B 278: 1165–1170.

35. Brusatte SL, Niedźwiedzki G, Butler RJ.2011. Footprints pull origin and diversification of dinosaur stem-lineage deep into Early Triassic. Proceedings of the Royal Society B 278: 1107–1113.

34. Porro LB, Butler RJ, Barrett PM, Moore-Fay S, Abel RL. 2011. New heterodontosaurid specimens from the Lower Jurassic of southern Africa and the early ornithischian dinosaur radiation. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 101: 351–366.

33. Barrett PM, Butler RJ, Nesbitt SJ. 2011. The roles of herbivory and omnivory in early dinosaur evolution. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 101: 383–396.


32. Wang X, Pan R, Butler RJ, Barrett PM. 2010. The postcranial skeleton of the iguanodontian ornithopod Jinzhousaurus yangi from the Lower Cretaceous Yixian Formation of western Liaoning, China. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 101: 135–159.

31. Butler RJ. 2010. The anatomy of the basal ornithischian dinosaur Eocursor parvus from the lower Elliot Formation (Late Triassic) of South Africa. Zoological Journal of the Linnean Society 160: 648–684.

30. Brusatte SL, Nesbitt SJ, Irmis RB, Butler RJ, Benton MJ, Norell MA. 2010. The origin and early radiation of dinosaurs. Earth-Science Reviews 101: 68–100.

29. Ősi A, Butler RJ, Weishampel DB. 2010. A Late Cretaceous ceratopsian dinosaur from Europe with Asian affinities. Nature 465: 466–468.

28. Butler RJ, Barrett PM, Kenrick P, Penn MG. 2010. Testing coevolutionary hypotheses over geological timescales: interactions between Cretaceous dinosaurs and plants. Biological Journal of the Linnean Society 100: 1–15.

27. Benson RBJ, Butler RJ, Lindgren J, Smith AS. 2010. Mesozoic marine tetrapod diversity: mass extinctions and temporal heterogeneity in geological megabiases affecting vertebrates. Proceedings of the Royal Society B 277: 829–834.

26. Jin L, Chen J, Zan S, Butler RJ, Godefroit P. 2010. Cranial anatomy of the small ornithischian dinosaur Changchunsaurus parvus from the Quantou Formation (Cretaceous: Aptian–Cenomanian) of Jilin Province, northeastern China. Journal of Vertebrate Paleontology 30: 196–214.

25. Butler RJ, Galton PM, Porro LB, Chiappe LM, Henderson DM, Erickson GM. 2010. Lower limits of ornithischian dinosaur body size inferred from a diminutive new Upper Jurassic heterodontosaurid from North America. Proceedings of the Royal Society B 277: 375–381.


24. Butler RJ, Barrett PM, Abel RL, Gower DJ. 2009. A possible ctenosauriscid archosaur from the Middle Triassic Manda Beds of Tanzania. Journal of Vertebrate Paleontology 29: 1022–1031.

23. Butler RJ, Barrett PM, Gower DJ. 2009. Postcranial skeletal pneumaticity and air-sacs in the earliest pterosaurs. Biology Letters 5: 557–560.

22. Brusatte SL, Butler RJ, Sulej T, Niedźwiedzki G. 2009. The taxonomy and anatomy of rauisuchian archosaurs from the Late Triassic of Germany and Poland. Acta Palaeontologica Polonica 54: 221–230.

21. Butler RJ, Barrett PM, Nowbath S, Upchurch P. 2009. Estimating the effects of the rock record on pterosaur diversity patterns: implications for hypotheses of bird/pterosaur competitive replacement. Paleobiology 35: 432–446.

20. Barrett PM, Butler RJ, Wang X-L, Xu X. 2009. Cranial anatomy of the iguanodontoid ornithopod Jinzhousaurus yangi from the Lower Cretaceous Yixian Formation of China. Acta Palaeontologica Polonica 54: 35–48.

19. Butler RJ, Sullivan RM. 2009. The phylogenetic position of the ornithischian dinosaur Stenopelix valdensis from the Lower Cretaceous of Germany: implications for the early fossil record of Pachycephalosauria. Acta Palaeontologica Polonica 54: 21–34.

18. Butler RJ, Barrett PM, Kenrick P, Penn MG. 2009. Diversity patterns amongst dinosaurs and plants during the Cretaceous: implications for hypotheses of dinosaur/angiosperm co-evolution. Journal of Evolutionary Biology 22: 446–459.

17. Butler RJ,Barrett PM, Kenrick P, Penn MG. 2009. Testing co-evolutionary hypotheses over geological timescales: interactions between Mesozoic non-avian dinosaurs and cycads. Biological Reviews 84: 73–89.

16. Butler RJ, Zhao Q. 2009. The small-bodied ornithischian dinosaurs Micropachycephalosaurus hongtuyanensis and Wannanosaurus yansiensis from the Late Cretaceous of China. Cretaceous Research30: 63–77.


15. Barrett PM, Butler RJ, Edwards NP, Milner AR. 2008. Pterosaur distribution in time and space: an atlas. Zitteliana B 28: 61–107.

14. Butler RJ, Upchurch P, Norman DB. 2008. The phylogeny of the ornithischian dinosaurs. Journal of Systematic Palaeontology 6: 1–40.

13. Butler RJ, Porro LB, Norman DB. 2008. A juvenile skull of the primitive ornithischian Heterodontosaurus tucki from the ‘Stormberg’ of South Africa. Journal of Vertebrate Paleontology 28: 702–711.

12. Butler RJ, Goswami A. 2008. Body size evolution in Mesozoic birds: little evidence for Cope's rule. Journal of Evolutionary Biology 26: 1673–1682.

11. Butler RJ, Barrett PM. 2008. Palaeoenvironmental controls on the distribution of Cretaceous herbivorous dinosaurs. Naturwissenschaften 95: 1027–1032.

10. Butler RJ, Galton PM. 2008. The ‘dermal armour’ of the Wealden dinosaur Hypsilophodon: a reappraisal. Cretaceous Research29: 636–642.

9. Chen J, Butler RJ, Jin L. 2008. New material of large-bodied ornithischian dinosaurs, including an iguanodontian ornithopod, from the Quantou Formation (middle Cretaceous: Aptian–Cenomanian) of Jilin Province, northeastern China. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 248: 309–314.

8. Barrett PM, Butler RJ, Novas FE, Moore-Fay S, Moody JM, Clark JM, Sanchez-Villagra MR. 2008. Dinosaur remains from the La Quinta Formation (Lower or Middle Jurassic) of the Venezuelan Andes. Paläontologische Zeitschrift 82: 163–177.


7. Butler RJ, Smith RMH, Norman DB. 2007. A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia. Proceedings of the Royal Society B 274: 2041–2046.

6. Norman DB, Butler RJ, Maidment SCH. 2007. Reconsidering the status and affinities of the ornithischian dinosaur Tatisaurus oehleri Simmons, 1965. Zoological Journal of the Linnean Society 150: 865–874.

5. Butler RJ, Upchurch P. 2007. The problem of highly incomplete taxa and the phylogenetic relationships of the theropod dinosaur Juravenator starkiJournal of Vertebrate Paleontology27: 253–256.

4. Corfe IJ, Butler RJ. 2006. Comment on ‘A Well-Preserved Archaeopteryx Specimen with Theropod Features’. Science 313:1238. (doi:10.1126/science.1130800)

3. Butler RJ, Porro LB, Heckert AB. 2006. A supposed heterodontosaurid tooth from the Rhaetian of Switzerland and a reassessment of the European Late Triassic record of Ornithischia (Dinosauria). Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 10: 613–633.

2. Barrett PM, Butler RJ, Knoll F. 2005. Small-bodied ornithischian dinosaurs from the Middle Jurassic of Sichuan, China. Journal of Vertebrate Paleontology 25: 823–834.

1. Butler RJ. 2005. The ‘fabrosaurid’ ornithischian dinosaurs of the Upper Elliot Formation (Lower Jurassic) of South Africa and Lesotho. Zoological Journal of the Linnean Society 145: 175–218.

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Dinosaurs, vertebrate palaeontology, fossils