The long necks of sauropods did not evolve primarily through sexual selection

Journal of Zoology

Volumn 285, Issue 2, 2011, Pages 150-161

  Taylor, Michael P ^a   Hone, D W E ^b   Wedel, M J ^c   Naish, D ^d  

^a UNIVERSITY OF BRISTOL   (United Kingdom)

^b UNIVERSITY COLLEGE DUBLIN   (Ireland)

^c WESTERN UNIVERSITY OF HEALTH SCIENCES   (United States)

^d UNIVERSITY OF PORTSMOUTH   (United Kingdom)

Author keywords

Behaviour; Dinosaurs; Ecology; Feeding; Giraffes; Necks; Sauropods; Sexual selection

Indexed keywords

ADAPTATION; ALLOMETRY; DATA SET; DINOSAUR; FEEDING BEHAVIOR; FEEDING ECOLOGY; FITNESS; GRAZING; MORPHOLOGY; ONTOGENY; PHYLOGENY; SEXUAL DIMORPHISM; SEXUAL SELECTION; SURVIVAL;

ANIMALIA; DINOSAURIA; EUHELOPODIDAE; GIRAFFIDAE; SAUROPODA; TETRAPODA;

EID: 80053053743     PISSN: 09528369     EISSN: 14697998     Source Type: Journal    
DOI: 10.1111/j.1469-7998.2011.00824.x     Document Type: Article

References (96)

1. Bakker, R.T. (1971). Ecology of the brontosaurs. Nature 229, 172-174.
2. Bakker, R.T. (1986). The dinosaur heresies. New York: Morrow.
3. Barnes, R.F.W. (1995). Elephants. In The encyclopedia of mammals, 2nd edn: 452-461. McDonald, D.W. (Ed.) Oxford: Oxford University Press.
4. Bonduriansky, R. (2007). Sexual selection and allometry: a critical reappraisal of the evidence and ideas. Evolution 61, 838-849.
5. Britt, B.B. & Naylor, B.G. (1994). An embryonic Camarasaurus (Dinosauria, Sauropoda) from the Upper Jurassic Morrison Formation (Dry Mesa Quarry, Colorado). In Dinosaur eggs and babies: 256-264. Carpenter, K., Hirsch, K.F. & Horner, J.R. (Eds) Cambridge: Cambridge University Press.
6. Cameron, E.Z. & du Toit, J.T. (2007). Winning by a neck: tall giraffes avoid competing with shorter browsers. Am. Nat. 169, 130-135.
7. Caro, T.M., Graham, C.M., Stoner, C.J. & Flores, M.M. (2003). Correlates of horn and antler shape in bovids and cervids. Behav. Ecol. Sociobiol. 55, 32-41.
8. Christian, A. & Dzemski, G. (2007). Reconstruction of the cervical skeleton posture of Brachiosaurus brancai Janensch, 1914 by an analysis of the intervertebral stress along the neck and a comparison with the results of different approaches. Foss. Rec. 10, 38-49.
9. Coombs, W.P. (1975). Sauropod habits and habitats. Palaeogeogr. Palaeoclimatol. Palaeoecol. 17, 1-33.
10. Dagg, A.I. (1971). Giraffa camelopardalis. Mamm. Species 5, 1-8.
11. Darwin, C. (1859). On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. London: John Murray.
12. Darwin, C. (1871). The descent of man in relation to sex. London: John Murray.
13. Ezenwa, V.O. & Jolles, A.E. (2008). Horns honestly advertise parasite infection in male and female African buffalo. Anim. Behav. 75, 2013-2021.
14. Farlow, J.O. & Pianka, E.R. (2000). Body form and trackway pattern in Australian desert monitors (Squamata:Varanidae): comparing zoological and ichnological diversity. Palaios 15, 235-247.
15. Folch, A. (1992). Family Casuariidae (Cassowaries). In Handbook of the birds of the World. Volume 1: ostriches to ducks: 90-97. Del Hoyo, J., Elliott, A. & Sargatal, J. (Eds) Barcelona: Lynx Edicions.
16. Foster, J.B. (1966). The giraffe of Nairobi national Park: home range, sex ratios, the herd and food. East Afri Wildl. J. 4, 139-148.
17. Fritts, T.H. (1984). Evolutionary divergence of giant tortoises in Galapagos. Biol. J. Linn. Soc. 21, 165-176.
18. Gauthier, J. (1986). Saurischian monophyly and the origin of birds. In The origin of birds and the evolution of flight (Memoirs of the California Academy of Sciences, 8): 1-55. Padian, K. (Ed.). San Francisco: California Academy of Sciences.
19. Ginnett, T.F. & Demment, M.W. (1997). Sex differences in giraffe foraging behavior at two spatial scales. Oecologica 110, 291-300.
20. Ginnett, T.F. & Demment, M.W. (1999). Sexual segregation by Masai giraffes at two spatial scales. Afr. J. Ecol. 37, 93-106.
21. Grimaldi, D. & Engel, M.S. (2005). Evolution of the insects. New York: Cambridge University Press.
22. Hatcher, J.B. (1903). Osteology of Haplocanthosaurus with description of a new species, and remarks on the probable habits of the Sauropoda and the age and origin of the Atlantosaurus beds; additional remarks on Diplodocus. Mem. Carnegie Mus. 2, 1-75, and plates I-VI.
23. Hoefs, M. (2000). The thermoregulatory potential of Ovis horn cores. Can. J. Z. 78, 1419-1426.
24. Hone, D.W.E. & Rauhut, O.W.M. (2009). Feeding behaviour and bone utilization by theropod dinosaurs. Lethaia 43, 232-244.
25. Hosken, D.J., Minder, A.M. & Ward, P.I. (2005). Male genital allometry in Scathophagidae (Diptera). Evol. Ecol. 19, 501-515.
26. Hunt, S., Cuthill, I.C., Bennett, A.T.D. & Griffiths, R.M. (1999). Preference for ultraviolet partners in the blue tit. Anim. Behav. 58, 809-815.
27. Huxley, J.S. (1914). The courtship habits of the great crested grebe Podiceps cristatus; with an addition to the theory of sexual selection. Proc. Zool. Soc. Lond. 35, 491-562.
28. Ikejiri, T., Tidwell, V. & Trexler, D.K. (2005). New adult specimens of Camarasaurus lentus highlight ontogenetic variation within the species. In Thunder-lizards: the Sauropodomorph dinosaurs: 154-179. Tidwell, V. & Carpenter, K. (Eds) Bloomington: Indiana University Press.
29. Jain, S.L. & Bandyopadhyay, S. (1997). New titanosaurid (Dinosauria:Sauropoda) from the Late Cretaceous of central India. J. Vertebr. Paleontol. 17, 114-136.
30. Janensch, W. (1936). Ein aufgestelltes Skelett von Dicraeosaurus hansemanni. Palaeontographica 7 (Suppl.), 301-308, plates XIV-XVI.
31. Janensch, W. (1950). Die Skelettrekonstruktion von Brachiosaurus brancai. Palaeontographica 7 (Suppl.), 97-103, plates VI-VIII.
32. van der Jeugd, H.T. & Prins, H.H.P. (2000). Movements and group structure of giraffe (Giraffa camelopardalis) in Lake Manyara National Park, Tanzania. J. Zool. (Lond.) 251, 15-21.
33. Jones, I.L. & Hunter, F.M. (1993). Mutual sexual selection in a monogamous seabird. Nature 362, 238-239.
34. Jones, I.L. & Hunter, F.M. (1999). Experimental evidence for mutual inter- and intrasexual selection favouring a crested auklet ornament. Anim. Behav. 57, 521-528.
35. Knell, R.J. & Sampson, S. (2010). Bizarre structures in dinosaurs: species recognition or sexual selection? A response to Padian and Horner. J. Zool. (Lond.) 283, 18-22.
36. Kotrba, M. (2004). Baltic amber fossils reveal early evolution of sexual dimorphism in stalk-eyed flies (Diptera: Diopsidae). Org. Divers. Evol. 4, 265-275.
37. Kraaijeveld, K., Gregurke, J., Hall, C., Komdeur, J. & Mulder, R.A. (2004). Mutual ornamentation, sexual selection, and social dominance in the black swan. Behav. Ecol. 15, 380-389.
38. Lammers, A.R. & German, R.Z. (2002). Ontogenetic allometry in the locomotor skeleton of specialized half-bounding mammals. J. Zool. (Lond.) 258, 485-495.
39. Larson, P. (2008). Variation and sexual dimorphism in Tyrannosaurus rex. In Tyrannosaurus rex, the tyrant king: 103-128. Larson, P. & Carpenter, K. (Eds). Bloomington: Indiana University Press.
40. Leuthold, W. (1978). Ecological separation among browsing ungulates in Tsavo East National Park, Kenya. Oecologica 35, 241-252.
41. Leuthold, W. (1979). Social organisation and behaviour of giraffe in Tsavo East National Park. Afr. J. Ecol. 17, 19-34.
42. Lovelace, D.M., Hartman, S.A. & Wahl, W.R. (2008). Morphology of a specimen of Supersaurus (Dinosauria, Sauropoda) from the Morrison Formation of Wyoming, and a re-evaluation of diplodocid phylogeny. Arq. Mus. Nac. Rio J. 65, 527-544.
43. Lydekker, R. (1894). Royal natural history, volume II. London: Frederick Warne and Co.
44. Mack, A.L. & Jones, J. (2003). Low-frequency vocalizations by cassowaries Casuarius spp. Auk 120, 1062-1068.
45. Mannion, P.D. & Upchurch, P. (2010). A quantitative analysis of environmental associations in sauropod dinosaurs. Paleobiology 36, 253-282.
46. Martin, J. (1987). Mobility and feeding of Cetiosaurus (Saurischia, Sauropoda) - why the long neck? In Fourth Symposium on Mesozoic Terrestrial Ecosystems, Short Papers: 154-159. Currie, P.J. & Koster, E.H. (Eds) Drumheller: Boxtree Books.
47. Mateus, O., Maidment, S.C.R. & Christiansen, N.A. (2009). A new long-necked 'sauropod-mimic' stegosaur and the evolution of the plated dinosaurs. Proc. Roy. Soc. Lond. Ser. B 276, 1815-1821.
48. McIntosh, J.S. (1990). Species determination in sauropod dinosaurs with tentative suggestions for the their classification. In Dinosaur systematics: approaches and perspectives: 53-69. Carpenter, K. & Currie, P.J. (Eds) Cambridge: Cambridge University Press.
49. McIntosh, J.S. (2005). The genus Barosaurus Marsh (Sauropoda, Diplodocidae). In Thunder-lizards: the Sauropodomorph dinosaurs: 38-77. Tidwell, V. & Carpenter, K. (Eds) Bloomington: Indiana University Press.
50. McNaughton, S.J. & Georgiadis, N.J. (1986). Ecology of African grazing and browsing mammals. Ann. Rev. Ecol. Syst. 17, 39-65.
51. Mitchell, G., van Sittert, S.J. & Skinner, J.D. (2009). Sexual selection is not the origin of long necks in giraffes. J. Zool. (Lond.) 278, 281-286.
52. Moll, E.O. (1986). Turtles and tortoises. In Animals of the World: reptiles and amphibians: 72-81. Halliday, T. & Adler, A. (Eds) Wembley: The Leisure Circle.
53. Novas, F.E., Salgado, L., Calvo, J. & Agnolin, F. (2005). Giant titanosaur (Dinosauria, Sauropoda) from the Late Cretaceous of Patagonia. Rev. Mus. Argent. Cienc. Nat. 7, 37-41.
54. Osborn, H.F. & Mook, C.C. (1921). Camarasaurus, Amphicoelias and other sauropods of Cope. Memoirs of the American Museum of Natural History, new series 3, 247-387 and plates LX-LXXXV.
55. Owen, M. (1980). Wild geese of the World. London: B. T. Batsford Ltd.
56. Owen-Smith, N. (2007). Changing vulnerability to predation related to season and sex in an African ungulate assemblage. Oikos 117, 602-610.
57. Padian, K. & Horner, J.R. (2010). The evolution of 'bizarre structures' in dinosaurs: biomechanics, sexual selection, social selection or species recognition? J. Zool. (Lond.) 283, 3-17.
58. Paul, G.S. (1998). Terramegathermy and Cope's Rule in the land of titans. Mod. Geol. 23, 179-217.
59. Raath, M.A. (1990). Morphological variation in small theropods and its meaning in systematics: evidence from Syntarsus rhodesiensis. In Dinosaur systematics: approaches and perspectives: 91-105. Carpenter, K. & Currie, P.J. (Eds) Cambridge: Cambridge University Press.
60. Rauhut, O.W.M., Remes, K., Fechner, R., Cladera, G. & Puerta, P. (2005). Discovery of a short-necked sauropod dinosaur from the Late Jurassic period of Patagonia. Nature 435, 670-672.
61. Russell, D.A. & Zheng, Z. (1993). A large mamenchisaurid from the Junggar Basin, Xinjiang, People's Republic of China. Can. J. Earth Sci. 30, 2082-2095.
62. Ruxton, G.D. & Wilkinson, D.M. (2011). The energetics of low browsing in sauropods. Biol. Lett. Online DOI: DOI: 10.1098/rsbl.2011.0116.
63. Sander, P.M. & Clauss, M. (2008). Sauropod gigantism. Science 322, 200-201.
64. Senter, P. (2006). Necks for sex: sexual selection as an explanation for sauropod dinosaur neck elongation. J. Zool. (Lond.) 271, 45-53.
65. Sereno, P.C. (1991). Basal archosaurs: phylogenetic relationships and functional implications. Mem. Soc. Vertebr. Paleontol. 2, 1-53.
66. Sereno, P.C., Wilson, J.A., Witmer, L.M., Whitlock, J.A., Maga, A., Ide, O. & Rowe, T.A. (2007). Structural extremes in a cretaceous dinosaur. PLoS ONE 2, e1230.
67. Seymour, R.S. (2009). Raising the sauropod neck: it costs more to get less. Biol. Lett. 5, 317-319.
68. Siegwarth, J.D., Smith, C.N. & Redman, P.D. (2010). An alternative sauropod physiology and cardiovascular system that eliminates high blood pressures. Lethaia 44, 46-57.
69. Simmons, L.W. & Tomkins, J.L. (1996). Sexual selection and the allometry of earwig forceps. Evol. Ecol. 10, 97-104.
70. Simmons, R.E. & Altwegg, R. (2010). Necks-for-sex or competing browsers? A critique of ideas on the evolution of giraffe. J. Zool. (Lond.) 282, 6-12.
71. Simmons, R.E. & Scheepers, L. (1996). Winning by a neck: sexual selection in the evolution of giraffe. Am. Nat. 148, 771-786.
72. Sinclair, A.R.E. (1985). Does interspecific competition or predation shape the African ungulate community? J. Anim. Ecol. 54, 899-918.
73. Stein, K., Csiki, Z., Curry Rogers, K., Weishampel, D.B., Redelstorff, R., Carballido, J.L. & Sander, P.M. (2010). Small body size and extreme cortical bone remodeling indicate phyletic dwarfism in Magyarosaurus dacus (Sauropoda:Titanosauria). PNAS 107, 9258-9263.
74. Stevens, K.A. & Parrish, J.M. (1999). Neck posture and feeding habits of two Jurassic sauropod dinosaurs. Science 284, 798-800.
75. Stevens, K.A. & Parrish, M.J. (2005). Digital reconstructions of sauropod dinosaurs and implications for feeding. In The sauropods: evolution and paleobiology: 178-200. Wilson, J.A. & Curry Rogers, K. (Eds) Berkeley: University of California Press.
76. Swallow, J.G., Hayes, J.P., Koteja, P. & Garland, T. Jr (2009). Selection experiments and experimental evolution of performance and physiology. In Experimental evolution: concepts, methods, and applications of selection experiments: 301-352. Garland, T. Jr & Rose, M.R. (Eds) Berkeley: California University Press.
77. Taylor, M.P. (2006). Dinosaur diversity analysed by clade, age, place and year of description. In Ninth international symposium on Mesozoic terrestrial ecosystems and biota, Manchester, UK: 134-138. Barrett, P. (Ed.) Cambridge: Cambridge Publications.
78. Taylor, M.P. (2009). Range of motion in intervertebral joints: why we don't trust DinoMorph. Sauropod Vertebra Picture of the Week, 30 May 2009. Available at (accessed 4 May 2011).
79. Taylor, M.P. (2010). Sauropod dinosaur research: a historical review. In Dinosaurs and other extinct saurians: a historical perspective, geological society Special Publication 343: 361-386. Moody, R.T.J., Buffetaut, E., Naish, D. & Martill, D.M. (Eds) London: Geological Society.
80. Taylor, M.P., Wedel, M.J. & Naish, D. (2009). Head and neck posture in sauropod dinosaurs inferred from extant animals. Acta Palaeontol. Pol. 54, 213-220.
81. Upchurch, P. (2000). Neck posture of sauropod dinosaurs. Science 287, 547b.
82. Upchurch, P. & Barrett, P.M. (2000). The evolution of sauropod feeding mechanisms. In Evolution of herbivory in terrestrial vertebrates: 79-122. Sues, H.-D. (Ed.) Cambridge: Cambridge University Press.
83. Upchurch, P., Barrett, P.M. & Dodson, P. (2004). Sauropoda. In The dinosauria, 2nd edn: 259-322. Weishampel, D.B., Dodson, P. & Osmólska, H. (Eds) Berkeley: University of California Press.
84. Upchurch, P., Tomida, Y. & Barrett, P.M. (2005). A new specimen of Apatosaurus ajax (Sauropoda: Diplodocidae) from the Morrison Formation (Upper Jurassic) of Wyoming, USA. National Science Museum Monographs No. 26. Tokyo: National Science Museum.
85. Van Sittert, S.J., Skinner, J.D & Mitchell, G. (2010). From fetus to adult - an allometric analysis of the giraffe vertebral column. J. Exp. Zool. Mol. Dev. Evol. 314B, 469-479.
86. Vidal, P.P., Graf, W. & Berthoz, A. (1986). The orientation of the cervical vertebral column in unrestrained awake animals. Exp. Brain Res. 61, 549-559.
87. Wedel, M.J. (2005). Postcranial skeletal pneumaticity in sauropods and its implications for mass estimates. In The Sauropods: evolution and paleobiology: 201-228. Wilson, J.A. & Curry Rogers, K. (Eds) Berkeley: University of California Press.
88. Wedel, M.J. (2007). Postcranial pneumaticity in dinosaurs and the origin of the avian lung. PhD dissertation, Integrative Biology, University of California, Berkeley, CA, USA.
89. Wedel, M.J. & Cifelli, R.L. (2005). Sauroposeidon:: Oklahoma's native giant. Okla. Geol. Notes 65, 40-57.
90. Wedel, M.J., Cifelli, R.L. & Sanders, R.K. (2000). Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. Acta Palaeontol. Pol. 45, 343-388.
91. Wilson, J.A. & Sereno, P.C. (1998). Early evolution and higher-level phylogeny of sauropod dinosaurs. Mem. Soc. Vertebr. Paleontol. 5, 1-68.
92. Wiman, C. (1929). Die Kreide-Dinosaurier aus Shantung. Palaeontol. Sin. Ser. C 6, 1-67.
93. Woolnough, A.P. & Du Toit, J.T. (2001). Vertical zonation of browse quality in tree canopies exposed to a size-structured guild of African browsing ungulates. Oecologica 129, 585-590.
94. Yates, A.M. & Kitching, J.W. (2003). The earliest known sauropod dinosaur and the first steps towards sauropod locomotion. Proc. R. Soc. Lond. Ser. B 270, 1753-1758.
95. Young, C.-C. & Zhao, X. (1972). [Chinese title. Description of the type material of Mamenchisaurus hochuanensis]. Inst. Vertebr. Paleontol. Paleoanthropol. Monogr. Ser. I 8, 1-30.
96. Young, T.P. & Isbell, L.A. (1991). Sex differences in giraffe feeding ecology: energetic & social constraints. Ethology 87, 79-89.