Functional and phylogenetic constraints in Rhinocerotinae craniodental morphology

Evolutionary Ecology Research

Volumn 12, Issue 8, 2010, Pages 897-928

  Piras, Paolo ^a   Maiorino, Leonardo ^a   Raia, Pasquale ^a,b   Marcolini, Federica ^a   Salvi, Daniele ^a,c   Vignoli, Leonardo ^a   Kotsakis, Tassos ^a  

^a ROMA TRE UNIVERSITY   (Italy)

^b UNIVERSITY OF NAPLES FEDERICO II   (Italy)

^c CENTRO DE INVESTIGAÇÃO EM BIODIVERSIDADE E RECURSOS GENÉTICOS   (Portugal)

Author keywords

Comparative methods; Craniodental morphology; Europe; Feeding habits; Geometric morphometrics; Plio Pleistocene; Rhinocerotinae

Indexed keywords

RHINOCEROTIDAE;

EID: 79955925381     PISSN: 15220613     EISSN: None     Source Type: Journal    
DOI: None     Document Type: Article

References (151)

1. Ackermann, R.R. and Cheverud, J.M. 2000. Phenotypic covariance structure in tamarins (genus Saguinus): A comparison of variation patterns using matrix correlation and common principal component analysis. Am. J. Phys. Anthropol., 111: 489-501. (Pubitemid 30185893)
2. Ackermann, R.R. and Cheverud, J.M. 2004a. Morphological integration in primate evolution. In Phenotypic Integration: Studying the Ecology and Evolution of Complex Phenotypes (M. Pigluicci and K. Preston, eds.), pp. 302-319. New York: Oxford University Press.
3. Ackermann, R.R. and Cheverud, J.M. 2004b. Detecting genetic drift versus selection in human evolution. Proc. Natl. Acad. Sci. USA, 101: 17946-17951. (Pubitemid 40054055)
4. Adams, D.C. 2008. Phylogenetic meta-analysis. Evolution, 62: 567-572. (Pubitemid 351348258)
5. Adams, D.C., Rohlf, F.J. and Slice, D.E. 2004. Geometric morphometrics: Ten years of progress following the 'revolution'. Ital. J. Zool., 71: 5-16. (Pubitemid 38531303)
6. Agusti, J. and Anton, M. 2002. Mammoths, Sabertooths and Hominids: 65 Million Years of Mammalian Evolution in Europe. New York: Columbia University Press.
7. Anderson, M.J. 2003. PCO: A FORTRAN Computer Program for Principal Coordinate Analysis. Auckland: Department of Statistics, University of Auckland.
8. Antoine, P.O. 2002. Phylogénie et évolution des Elasmotheriina (Mammalia, Rhinocerotidae). Mém. Mus. Nat. Hist. Nat. Paris, 188: 5-350.
9. Arnold, S.J. 1994. Investigating the origins of the performance advantage: Adaptation, exaptation and lineage effects. In Phylogenetics and Ecology (P. Eggleton and R. Vane-Wright, eds.), pp. 123-168. London: Academic Press.
10. Arthur, W. 2001. Developmental drive: An important determinant of the direction of phenotypic evolution. Evol. Develop., 3: 271-278. (Pubitemid 32665254)
11. Atchley, W.R. 1993. Genetic and developmental aspects of variability in the mammalian mandible. In The Skull, Vol. 1: Development (J. Hanken and B.K. Hall, eds.), pp. 207-247. Chicago, IL: University of Chicago Press.
12. Atchley, W.R. and Hall, B.K. 1991. A model for development and evolution of complex morphological structures. Biol. Rev. Cambr. Phil. Soc., 66: 101-157. (Pubitemid 121000606)
13. Atchley, W.R., Nordheim, E.V., Gunsett, F.C. and Crump, P.L. 1982. Geometric and probabilistic aspects of statistical distance functions. Syst. Zool., 31: 445-460.
14. Badyaev, A.V. and Foresman, K.R. 2000. Extreme environmental change and evolution: Stressinduced morphological variation is strongly concordant with patterns of evolutionary divergence in shrew mandibles. Proc. R. Soc. Lond. B, 267: 371-377. (Pubitemid 30133061)
15. Badyaev, A.V. and Foresman, K.R. 2004. Evolution of morphological integration. I. Functional units channel stress-induced variation in shrew mandibles. Am. Nat., 163: 868-879. (Pubitemid 38852140)
16. Badyaev, A.V., Foresman, K.R. and Young, R.L. 2005. Evolution of morphological integration. II. Developmental accommodation of stress-induced variation in shrew mandibles. Am. Nat., 166: 382-395. (Pubitemid 41266025)
17. Bales, G.S. 1996. Skull evolution in the Rhinocerotidae (Mammalia, Perissodactyla): Cartesian transformations and functional interpretations. J. Mammal. Evol., 3: 261-279. (Pubitemid 127017827)
18. Bastir, M. and Rosas, A. 2005. The hierarchical nature of morphological integration and modularity in the human posterior face. Am. J. Phys. Anthropol., 128: 26-34. (Pubitemid 41216057)
19. Bastir, M. and Rosas, A. 2006. Correlated variation between the lateral basicranium and the face: A geometric morphometric study in different human groups. Arch. Oral Biol., 51: 814-824. (Pubitemid 44233480)
20. Blomberg, S.P., Garland, T., Jr. and Ives, A.R. 2003. Testing for phylogenetic signal in comparative data: Behavioural traits are more labile. Evolution, 57: 717-745. (Pubitemid 36579712)
21. Bookstein, F.L. 1986. Size and shape spaces for landmark data in two dimensions. Stat. Sci., 1: 181-242.
22. Bookstein, F.L. 1991. Morphometric Tools for Landmark Data: Geometry and Biology. Cambridge: Cambridge University Press.
23. Bookstein, F.L., Streissguth, A.P., Sampson, P.D., Connor, P.D. and Barr, H.H. 2002. Corpus callosum shape and neuropsychological deficits in adult males with heavy fetal alcohol exposure. NeuroImage, 15: 233-251. (Pubitemid 34852223)
24. Borcard, D., Legendre, P., and Drapeau, P. 1992. Partialling out the spatial component of ecological variation. Ecology, 73: 1045-1055.
25. Cerdeño, E. 1998. Diversity and evolutionary trends of the family Rhinocerotidae (Perissodactyla). Palaeogeogr. Palaeoclimatol. Palaeoecol., 141: 13-34. (Pubitemid 28450809)
26. Cheverud, J.M. 1982. Phenotypic, genetic, and environmental morphological integration in the cranium. Evolution, 36: 499-516.
27. Cheverud, J.M. 1989. Evolution of morphological integration. In Trends in Vertebrate Morphology: Proceedings of the 2nd International Symposium on Vertebrate Morphology (H. Splechtna and H. Helge, eds.). Fortschr. Zool., 35: 196-197.
28. Cheverud, J.M. 1995. Morphological integration in the saddle-back tamarin (Saguinus fuscicollis) cranium. Am. Nat., 145: 63-89.
29. Cheverud, J.M. 1996. Developmental integration and the evolution of pleiotropy. Am. Zool., 36: 44-50. (Pubitemid 126043153)
30. Cheverud, J.M. 2004. Modular pleiotropic effects of quantitative trait loci on morphological traits. In Modularity in Development and Evolution (G. Schlosser & G.P. Wagner, eds.), pp. 132-153.
31. Chicago, IL: University of Chicago Press.
32. Cheverud, J.M., Hartman, S.E., Richtsmeier, J.T. and Atchley, W.R. 1991. A quantitative genetic analysis of localized morphology in mandibles of inbred mice using finite element scaling analysis. J. Craniofac. Genet. Dev. Biol., 11: 122-137.
33. Cheverud, J.M., Leamy, L.J. and Routman, E.J. 1997. Pleiotropic effects of individual gene loci on mandibular morphology. Evolution, 51: 2004-2014.
34. Cheverud, J.M., Ehrich, T.H., Vaughn, T.T., Koreishi, S.F., Linsey, R.B. and Pletscher, L.S. 2004. Pleiotropic effects on mandibular morphology II: Differential epistasis and genetic variation in morphological integration. J. Exp. Zool. B: Molec. Develop. Evol., 302B: 424-435. (Pubitemid 39428814)
35. Claude, J. 2008. Morphometrics with R. New York: Springer.
36. Clauss, M. and Hummel, J. 2005. The digestive performance of mammalian herbivores: Why big may not be that much better. Mammal. Rev., 35: 174-187. (Pubitemid 40900014)
37. Clauss, M., Frey, R., Kiefer, B., Lechner-Doll, M., Loehlein, W., Polster, Cö. et al. 2003. The maximum attainable body size of herbivorous mammals: Morphophysiological constraints on foregut, and adaptations of hindgut fermenters. Oecologia, 136: 14-27. (Pubitemid 36837257)
38. Codron, D., Codron, J., Lee-Thorp, J.A., Sponheimer, D., de Ruiter, D., Sealy, J. et al. 2007. Diets of savanna ungulates from stable carbon isotope composition of faeces. J. Zool., 273: 21-29. (Pubitemid 47300726)
39. Cubo, J., Ponton, F., Laurin, M., De Margerie, E. and Castanet, J. 2005. Phylogenetic signal in bone microstructure of Sauropsids. Syst. Biol., 54: 562-574. (Pubitemid 41356714)
40. Cubo, J., Legendre, P., de Ricqlès, A., Montes, L., de Margerie, E., Castanet, J. et al. 2008. Phylogenetic, functional and structural components of variation in bone growth rate of amniotes. Evol. Develop., 10: 217-227. (Pubitemid 351341482)
41. Cumming, D.H.M., Du Toit, R.F. and Stuart, S.N. 1990. African Elephants and Rhinos: Status Survey and Conservation Action Plan. IUCN/SSC African Elephant and Rhino Specialist Group. Gland, Switzerland: IUCN.
42. Dayan, T., Wool, D. and Simberloff, D. 2002. Variation and covariation of skulls and teeth: Modern carnivores and the interpretation of fossil mammals. Paleobiology, 28: 508-526. (Pubitemid 36029654)
43. Demment, M.W. and Van Soest, P.J. 1985. A nutritional explanation for body-size patterns of ruminant and non-ruminant herbivores. Am. Nat., 125: 1-641.
44. Desdevises, Y., Legendre, P., Azouzi, L. and Morand, S. 2003. Quantifying phylogenetically structured environmental variation. Evolution, 57: 2647-2652. (Pubitemid 37502567)
45. Dierenfeld, E. 1995. Rhinoceros nutrition: An overview with special reference to browsers. Verh. Ber. Erkrg. Zootiere, 37: 7-14.
46. Diniz-Filho, J.A.F., Sant'ana, C.E.R. and Bini, L.M. 1998. An eigenvector method for estimating phylogenetic inertia. Evolution, 52: 1247-1262. (Pubitemid 28537236)
47. Drake, A.G. and Klingenberg, C.P. 2010. Large-scale diversification of skull shape in domestic dogs: Disparity and modularity. Am. Nat., 175: 289-301.
48. Ehrich, T.H., Vaughn, T.T., Koreishi, S.F., Linsey, R.B., Pletscher, L.S. and Cheverud, J.M., 2003. Pleiotropic effects on mandibular morphology II. Developmental morphological integration and differential dominance. J. Exp. Zool. B: Molec. Develop. Evol., 296B: 58-79. (Pubitemid 37529151)
49. Emslie, R. and Brooks, M. 1999. African Rhino: Status Survey and Conservation Action Plan. Gland, Switzerland: IUCN.
50. Escoufier, Y. 1973. Le traitement des variables vectorielles. Biometrics, 29: 751-760.
51. Felsenstein, J. 1985. Phylogenies and the comparative method. Am. Nat., 125: 1-15.
52. Feranec, R.S. 2007. Ecological generalization during adaptive radiation: Evidence from Neogene mammals. Evol. Ecol., 9: 555-577. (Pubitemid 46998128)
53. Fortelius, M. 1981. Functional aspects of occlusal cheek-tooth morphology in hypsodont, non-ruminant ungulates. In International Symposium on Concept and Method in Paleontology: Contributed Papers (J. Martinell, ed.), pp. 153-162. Barcelona: University of Barcelona.
54. Fortelius, M. 1982. Ecological aspects of dental functional morphology in the Plio-Pleistocene rhinoceroses of Europe. In Teeth: Form, Function and Evolution (B. Kurten, ed.), pp. 163-181. New York: Columbia University Press.
55. Fortelius, M. and Solounias, N. 2000. Functional characterization of ungulate molars using the abrasion-attrition wear gradient: A new method for reconstructing paleodiets. Am. Mus. Nov., 3301: 1-36.
56. Fortelius, M., Mazza, P. and Sala, B. 1993. Stephanorhinus (Mammalia, Rhinocerotidae) of the western European Pleistocene, with a special revision of Stephanorhinus etruscus (Falconer, 1868). Pal. Ital., 80: 63-155.
57. Frontier, S. 1976. Étude de la décroissance des valeurs propres dans une analyse en composantes principales: Comparaison avec le modèle du bâton brisé. J. Exp. Mar. Biol. Ecol., 25: 67-75.
58. Garland, T., Jr. 1992. Rate tests for phenotypic evolution using phylogenetically independent contrasts. Am. Nat., 140: 509-519.
59. Garland, T., Jr., Harvey, P.H. and Ives, A.R. 1992. Procedures for the analysis of comparative data using phylogenetically independent contrasts. Syst. Biol., 41: 18-32.
60. Garland, T., Jr., Dickerman, A.W., Janis, C.M. and Jones, J.A. 1993. Phylogenetic analysis of covariance by computer simulation. Syst. Biol., 42: 265-292.
61. Garland, T., Jr., Bennett, A.F. and Rezende, E.L. 2005. Phylogenetic approaches in comparative physiology. J. Exp. Biol., 208: 3015-3035. (Pubitemid 41345680)
62. Geraads, D. 2005. Pliocene Rhinocerotidae (Mammalia) from Hadar e Dikika (Lower Awash, Ethiopia) and a revision of the origin of modern African rhinos. J. Vert. Paleontol., 25: 451-461. (Pubitemid 41057643)
63. Giaourtsakis, I., Theodorou, G., Roussiakis, S., Athanassiou, A. and Iliopoulos, G. 2006. Late Miocene horned rhinoceroses (Rhinocerotinae, Mammalia) from Kerassia (Euboea, Greece). N. Jb. Geol. Pal. Abh., 239: 367-398. (Pubitemid 43676208)
64. Gordon, I.J. and Illius, A.W. 1988. Incisor arcade structure and diet selection in ruminants. Funct. Ecol., 2: 15-22.
65. Goswami, A. 2006. Cranial modularity shifts during mammalian evolution. Am. Nat., 168: 270-280.
66. Goswami, A. and Polly, P.D. 2010. The influence of modularity on cranial morphological disparity in Carnivora and Primates (Mammalia). PLoS One, 5(3): E9517, 1-8.
67. Goswami, A. and Prochel, J. 2007. Ontogenetic morphology and cranial allometry of the common European mole (Talpa europaea). J. Mammal., 88: 667-677. (Pubitemid 47001221)
68. Gould, S.J. 2002. The Structure of Evolutionary Theory. Cambridge, MA: Belknap Press of Harvard University Press.
69. Gould, S.J. and Lewontin, R.C. 1979. The spandrels of San Marco and the Panglossian paradigm: A critique of the adaptationist programme. Proc. R. Soc. Lond. B, 205: 581-598. (Pubitemid 10248042)
70. Gould, S.J. and Vrba, E.S. 1982. Exaptation - a missing term in the science of form. Paleobiology, 8: 4-15. (Pubitemid 13281606)
71. Groves, C.P. and Kurt, F. 1972. Dicerorhinus sumatrensis. Mammal. Species, 21: 1-6.
72. Guérin, C. 1980. Les rhinocéros (Mammalia, Perissodactyla) du Miocène terminal au Pléistocène supérieur en Europe occidentale. Comparaison avec les espèces actuelles. Doc. Lab. Géol. Lyon, 79: Fasc. 1, 2, 3, 1185 pp.
73. Hallgrímsson, B., Willmore, K. and Hall, B. 2002. Canalization, developmental stability, and morphological integration in primate limbs. Yearbook Phys. Anthropol., 45: 131-158. (Pubitemid 36109385)
74. Harmon, L., Weir, J., Brock, C., Glor, R., Challenger, W. and Hunt, G. 2009. GEIGER: Analysis of Evolutionary Diversification. R package version 1.3-1 (http://CRAN.R-project.org/package=Geiger).
75. Harvey, P.H. and Pagel, M.D. 1991. The Comparative Method in Evolutionary Biology. Oxford: Oxford University Press.
76. Heissig, K. 1989. The Rhinocerotidae. In The Evolution of Perissodactyls (D.R. Prothero and R.M. Schoch, eds.), pp. 399-417. New York: Oxford University Press.
77. Janis, C.M. 1976. The evolutionary strategy of the Equidae and the origin of rumen and cecal digestion. Evolution, 30: 757-774.
78. Janis, C.M. 1995. Correlations between craniodental morphology and feeding behaviour in ungulates: Reciprocal illumination between living and fossil taxa. In Functional Morphology in Vertebrate Paleontology (J.J. Thomason, ed.), pp. 76-98. Cambridge: Cambridge University Press.
79. Janis, C.M. 2008. An evolutionary history of browsing and grazing ungulates. In The Ecology of Browsing and Grazing (I.J. Gordon and H.H.T. Prins, eds.), pp. 21-45. Berlin: Springer.
80. Josse, S., Moreau, T. and Laurin, M. 2006. Stratigraphic Tools for Mesquite (http://mesquiteproject .org/packages/stratigraphicTools).
81. Kahlke, R.D. and Kaiser, T.M. in press. Generalism as a subsistence strategy: Advantages and limitations of the highly flexible feeding traits of Pleistocene Stephanorhinus hundsheimensis (Rhinoceritidae, Mammalia). Quat. Sci. Rev.
82. Kahlke, R.D. and Lacombat, F. 2008. The earliest immigration of woolly rhinoceros (Coelodonta tologoijensis, Rhinocerotidae, Mammalia) into Europe and its adaptive evolution in Palaearctic cold stage mammal faunas. Quat. Sci. Rev., 27: 1951-1961.
83. Klingenberg, C.P. 2005. Developmental constraints, modules and evolvability. In Variation: A Central Concept in Biology (B. Hallgrímsson and B.K. Hall, eds.), pp. 219-247. Burlington, MA: Elsevier.
84. Klingenberg, C.P. 2009. Morphometric integration and modularity in configurations of landmarks: Tools for evaluating a-priori hypotheses. Evol. Develop., 11: 405-421.
85. Klingenberg, C.P. 2011. MorphoJ: An integrated software package for geometric morphometrics. Molec. Ecol. Res. (DOI: 10.1111/j.1755-0998.2010.02924. x).
86. Klingenberg, C.P. and Leamy, L.J. 2001. Quantitative genetics of geometric shape in the mouse mandible. Evolution, 55: 2342-2352. (Pubitemid 34045563)
87. Klingenberg, C.P., Leamy, L.J., Routman, E.J. and Cheverud, J.M. 2001. Genetic architecture of mandible shape in mice: Effects of quantitative trait loci analyzed by geometric morphometrics. Genetics, 157: 785-802. (Pubitemid 32165034)
88. Klingenberg, C.P., Barluenga, M. and Meyer, A. 2003. Body shape variation in cichlid fishes of the Amphilophus citrinellus species complex. Biol. J. Linn. Soc., 80: 397-408. (Pubitemid 37435945)
89. Klingenberg, C.P., Leamy, L.J. and Cheverud, J.M. 2004. Integration and modularity of quantitative trait locus effects on geometric shape in the mouse mandible. Genetics, 166: 1909-1921. (Pubitemid 38621070)
90. Lacombat, F. 2003. Étude des rhinocéroses du Pléistocène de l'Europe méditerranéenne et du Massif Central. Paléontologie, phylogénie et biostratigraphie. Unpublished DPhil thesis. Paris : Museum National d'Histoire Naturelle.
91. Laurie, W.A. 1982. Behavioural ecology of the greater one-horned rhinoceros (Rhinoceros unicornis). J. Zool., 196: 307-341.
92. Laurie, W.A., Lang, E.M. and Groves, C.P. 1983. Rhinoceros unicornis. Mammal. Species, 211: 1-6.
93. Lavin, S.R., Karasov, W.H., Ives, A.R., Middleton, K.M. and Garland, T., Jr. 2008. Morphometrics of the avian small intestine compared with that of nonflying mammals: A phylogenetic approach. Physiol. Biochem. Zool., 81: 526-550.
94. Leamy, L.J., Routman, E.J. and Cheverud, J.M. 1999. Quantitative trait loci for early and late developing skull characters in mice: A test of the genetic independence model of morphological integration. Am. Nat., 153: 201-214. (Pubitemid 29139436)
95. Leamy, L.J., Routman, E.J. and Cheverud, J.M. 2002. An epistatic genetic basis for fluctuating asymmetry of mandible size in mice. Evolution, 56: 642-653. (Pubitemid 34299838)
96. Lieberman, D.E., Ross, C.F. and Ravosa, M.J. 2000. The primate cranial base: Ontogeny, function, and integration. Yearbook Phys. Anthropol., 43: 117-169. (Pubitemid 32057404)
97. Loose, H. 1975. Pleistocene Rhinocerotidae of W. Europe with reference to the recent two-horned species of Africa and S.E. Asia. Scr. Geol., 33: 1-59.
98. Louys, J., Curnoe, D. and Tong, H. 2007. Characteristics of Pleistocene megafauna extinctions in Southeast Asia. Palaeogeogr. Palaeoclimatol. Palaeoecol., 243: 152-173. (Pubitemid 44909221)
99. Maddison, W.P. and Maddison, D.R. 2007. Mesquite: A Modular System for Evolutionary Analysis, Version 2.01 (http://mesquiteproject.org).
100. Marcus, L.F., Hingst-Zaher, E. and Zaher, H. 2000. Application of landmarks morphometrics to skull representing the orders of living mammals. Hystrix, 11: 27-47.
101. Márquez, E.J. 2008. A statistical framework for testing modularity in multidimensional data. Evolution, 62: 2688-2708.
102. Marroig, G. and Cheverud, J.M. 2001. A comparison of phenotypic variation and covariation patterns and the role of phylogeny, ecology, and ontogeny during cranial evolution of New World monkeys. Evolution, 55: 2576-2600. (Pubitemid 34115687)
103. Marroig, G., De Vivo, M. and Cheverud, J.M. 2004. Cranial evolution in sakis (Pithecia, Platyrrhini) II: Evolutionary processes and morphological integration. J. Evol. Biol., 17: 144-155. (Pubitemid 38162189)
104. Martins, E.P., Diniz-Filho, J.A.F. and Housworth, E.A. 2002. Adaptive constraints and the phylogenetic comparative method: A computer simulation test. Evolution, 56: 1-13. (Pubitemid 34243797)
105. Maynard Smith, J., Burian, R., Kaufman, S., Alberch, P. and Campbell, J. 1985. Developmental constraints and evolution. Quart. Rev. Biol., 60: 265-287.
106. Mazza, P. and Azzaroli, A. 1993. Ethological inferences on Pleistocene rhinoceroses of Europe. Rend. Fis. Accad. Lincei, 4: 127-137.
107. McCullagh, P. 1980. Regression models for ordinal data. J. R. Stat. Soc. Lond. B (Methodological), 42: 109-142.
108. Meloro, C., Raia, P., Piras, P., Barbera, C. and 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.
109. Mendoza, M. and Palmqvist, P. 2008. Hypsodonty in ungulates: An adaptation for grass consumption or for foraging in open habitat? J. Zool., 274: 134-142. (Pubitemid 351160851)
110. Mendoza, M., Janis, C.M. and Palmqvist, P. 2002. Characterizing complex craniodental patterns related to feeding behaviour in ungulates: A multivariate approach. J. Zool., 258: 223-246. (Pubitemid 35255069)
111. Mullin, S.K. and Taylor, P.J. 2002. The effects of parallax on geometric morphometric data. Comp. Biol. Med., 32: 455-464. (Pubitemid 35287135)
112. Nanda, A.C. 2002. Upper Siwalik mammalian faunas of India and associated events. J. Asian Earth Sci., 21: 47-58. (Pubitemid 35376301)
113. Nowak, R.M. 1991. Walker's Mammals of the World. Baltimore, MD: Johns Hopkins University Press.
114. 2 in regression analysis. Econ. Model., 17: 473-483.
115. Oksanen, J., Kindt, R., Legendre, P., O'Hara, B., Simpson, G.L., Solymos, P. et al. 2008. Vegan: Community Ecology Package. R Package version 1.15-1 (http:// cran.r-project.org/, http:// vegan.r-forge.r-project.org/).
116. Orlando, L., Leonard, J.A., Thenot, A., Laudet, V., Guérin, C. and Hänni, C. 2003. Ancient DNA analysis reveals woolly rhino evolutionary relationships. Molec. Phylogenet. Evol., 28: 485-499. (Pubitemid 37436117)
117. Owen-Smith, R.N. 1988. Megaherbivores: The Influence of Very Large Body Size on Ecology. Cambridge: Cambridge University Press.
118. Peres-Neto, P.R., Legendre, P., Dray, S. and Borcard, D. 2006. Variation partitioning of species data matrices: Estimation and comparison of fractions. Ecology, 87: 2614-2625. (Pubitemid 44885972)
119. Perez, S.I., Bernal, V. and Gonzalez, P.N. 2006. Differences between sliding semi-landmark methods in geometric morphometrics, with an application to human craniofacial and dental variation. J. Anat., 208: 769-784. (Pubitemid 43805744)
120. Perez-Barberia, F.J. and Gordon, I.J. 1999. The functional relationship between feeding type and jaw and cranial morphology in ungulates. Oecologia, 118: 157-165. (Pubitemid 29112044)
121. Perez-Barberia, F.J. and Gordon, I.J. 2001. Relationships between oral morphology and feeding style in the Ungulata: A phylogenetically controlled evaluation. Proc. R. Soc. Lond. B, 268: 1023-1032. (Pubitemid 32500475)
122. Piras, P., Colangelo, P., Adams, D.C., Buscalioni, A., Cubo, J., Kotsakis, T. et al. 2010. The Gavialis- Tomistoma debate: The contribution of skull ontogenetic allometry and growth trajectories to the study of crocodylian relationships. Evol. Develop., 12: 568-579.
123. Polly, P.D. 2005. Development and phenotypic correlations: The evolution of tooth shape in Sorex araneus. Evol. Develop., 7: 29-41. (Pubitemid 40210830)
124. Pratiknyo, H. 1991. The diet of the Javan rhino. Voice Nat., 93: 12-13.
125. Prithiviraj, F., Polet, G., Foead, N., Ng, L., Pastorini, J. and Melnick, D.J. 2006. Genetic diversity, phylogeny and conservation of the Javan rhinoceros (Rhinoceros sondaicus). Conserv. Genet., 7: 439-448. (Pubitemid 43873296)
126. Raia, P., Carotenuto, F., Meloro, C., Piras, P. and Pushkina, D. 2010. The shape of contention: Adaptation, history and contingency in ungulate mandibles. Evolution, 64: 1489-1503.
127. Ramette, A. and Tiedje, J. 2007. Multiscale responses of microbial life to spatial distance and environmental heterogeneity in a patchy ecosystem. Proc. Natl. Acad. Sci. USA, 104: 2761-2766. (Pubitemid 46327953)
128. R Development Core Team (2010). R: A Language and Environment for Statistical Computing. Vienna, Austria: R Foundation for Statistical Computing (URL: Http://www.R-project.org).
129. Rohlf, F.J. 2001. Comparative methods for the analysis of continuous variables: Geometric interpretations. Evolution, 55: 2143-2160. (Pubitemid 34049746)
130. Rohlf, F.J. 2005. tpsDig, Digitize Landmarks and Outlines. Version 2.05. Department of Ecology and Evolution, State University of New York at Stony Brook.
131. Rohlf, F.J. 2006. A comment on phylogenetic corrections. Evolution, 60: 1509-1515.
132. Schwenk, K. 1995. A utilitarian approach to evolutionary constraint. Zoology, 98: 251-262.
133. Shrader, A.M., Owen-Smith, N. and Ogutu, J.O. 2006. How a mega-grazer copes with the dry season: Food and nutrient intake rates by white rhinoceros in the wild. Funct. Ecol., 20: 376-384. (Pubitemid 43725643)
134. Solounias, N., Teaford, M. and Walker, A. 1988. Interpreting the diet of extinct ruminants: The case of a non-browsing giraffid. Paleobiology, 14: 287-300. (Pubitemid 19309976)
135. Solounias, N., Moelleken, S.M.C. and Plavcan, J.M. 1995. Predicting the diet of extinct bovids using masseteric morphology. J. Vert. Paleontol., 15: 795-805. (Pubitemid 126024582)
136. Spencer, LM. 1995. Morphological correlates of dietary resource partitioning in the African Bovidae. J. Mammal., 76: 448-471.
137. Sponheimer, M., Lee-Thorp, J.A., DeRuiter, D.J., Smith, J.M., van der Merwe, N.J., Reed, K. et al. 2003. Diets of southern African bovidae: Stable isotope evidence. J. Mammal., 84: 471-479. (Pubitemid 36689356)
138. Strait, D.S. 2001. Integration, phylogeny, and the hominid cranial base. Am. J. Phys. Anthropol., 114: 273-297. (Pubitemid 32275968)
139. Strömberg, C.A.E. 2006. The evolution of the hypsodonty in equids: Testing a hypothesis of adaptation. Paleobiology, 32: 236-258.
140. Tong, H. 2001. Rhinocerotids in China, systematics and material analysis. Geobios, 34: 585-591. (Pubitemid 33109556)
141. Tougard, C., Delefosse, T., Hänni, C. and Montgelard, C. 2001. Phylogenetic relationships of the five extant rhinoceros species (Rhinocerotidae, Perissodactyla) based on mitochondrial cytochrome b and 12S rRNA genes. Molec. Phylogenet. Evol., 19: 34-44. (Pubitemid 32996698)
142. Wagner, G.P. and Altenberg, L. 1996. Complex adaptations and the evolution of evolvability. Evolution, 50: 967-976.
143. Westoby, M., Leishmann, M.R. and Lord, J.M. 1995. On misinterpreting the 'phylogenetic correction'. J. Ecol., 83: 531-534.
144. Williams, S.H. and Kay, R.F. 2001. A comparative test for adaptive explanations for hypsodonty in ungulates and rodents. J. Mammal. Evol., 8: 207-229. (Pubitemid 33815230)
145. Young, N.M. and Hallgrímsson, B. 2005. Serial homology and the evolution of mammalian limb covariation structure. Evolution, 59: 2691-2704. (Pubitemid 43069949)
146. Zelditch, M.L. 1988. Developmental and functional constraints on phenotypic covariation during growth and evolution. Dissert. Abstr. Intern. B Sci. Engineer., 48: 1912-1913.
147. Zelditch, M.L. and Carmichael, A.C. 1989a. Ontogenetic variation in patterns of developmental and functional integration in skulls of Sigmodon fulviventer. Evolution, 43: 814-824.
148. Zelditch, M.L. and Carmichael, A.C. 1989b. Growth and intensity of integration through postnatal growth in the skull of Sigmodon fulviventer. J. Mammal., 70: 477-484.
149. Zelditch, M.L. and Moscarella, R.A. 2004. Form, function, and life history: Spatial and temporal dynamics of integration. In Phenotypic Integration: Studying the Ecology and Evolution of Complex Phenotypes (M. Pigluicci and K. Preston, eds.), pp. 274-301. New York: Oxford University Press.
150. Zelditch, M.L., Wood, A.R. and Swiderski, D.L. 2009. Building developmental integration into functional systems: Function-induced integration of mandibular shape. Evol. Biol. (New York), 36: 71-87.
151. Zeuner, F.E. 1934. Die Beziehungen zwischen Schädelform und Lebensweise bei den rezenten und fossilen Nashörnern. Ber. Naturf. Ges. Freiburg i. B, 34: 31-80.