Evolutionary Ecology

, Volume 17, Issue 3, pp 293–312 | Cite as

The fast life of a dwarfed giant

  • Pasquale RaiaEmail author
  • Carmela Barbera
  • Maurizio Conte


In the first half of the 1960s, a rich paleontological site was discovered at Spinagallo caves (Eastern Sicily, Southern Italy). A very abundant fossil population (at least 104 specimens) of the dwarf elephant Elephas falconeri, the smallest elephant that ever lived, was recovered. We computed the survivorship curve for this fossil population in order to investigate both the great juvenile abundance and high calf mortality which it shows. Through the analysis of E. falconeri survivorship, of some reconstructed life-history traits, and of its ecology, and taking into account the Island rule (Foster, 1964), we concluded that E. falconeri moved somewhat toward the ‘fast’ extreme of the slow-fast continuum in life-history traits in regards to its mainland ancestor E. antiquus, that is, it was somehow r-selected. In keeping with our findings, we propose a new explanation for the common occurrence of dwarfism in large mammals living on islands. We suggest the interplay of competition, resource allocation shift and feeding niche width could successfully explain this pattern.

dwarfism elephants island rule life history resource allocation 


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  1. Abe, E. (1996) Tusklessness among the Queen Elizabeth National Park elephants, Uganda. Pachyderm 22, 46-47.Google Scholar
  2. Adler, G.H. and Levins, R. (1994) The island syndrome in rodent populations. Quart. Rev. Biol. 69(4), 473-490.Google Scholar
  3. Alcover, J.A. and McMinn, M. (1994) Bioscience 44, 12-19.Google Scholar
  4. Ambrosetti, P. (1968) The Pleistocene dwarf elephants of Spinagallo (Siracusa, South-Eastern Sicily). Geol. Rom. 7, 277-398.Google Scholar
  5. Bada, J.L., Belluomini, G., Bonfiglio, L., Branca, M., Bugio, E. and Delitala, L. (1991) Isoleucine epimerization ages of Quaternari mammals of Sicily. Il Quaternario 4(1a), 5-11.Google Scholar
  6. Belluomini, G. and Bada, J.L. (1985) Isoleucene epimerization ages of the dwarf elephants of Sicily. Geology 13, 451-452.Google Scholar
  7. Berger, J. and Gompper, M.E. (1999) Sex ratios in extant ungulates: products of contemporary predation or past life histories? J. Mammal. 80(4), 1084-1113.Google Scholar
  8. Bertoldi, R., Rio, D. and Thunell, R. (1989) Pliocene-Pleistocene vegetational and climatic evolution of the South-Central Mediterranean. Palaeog. Palaeocl. Palaeoe. 72, 263-275.Google Scholar
  9. Bonfiglio, L., Di Geronimo, S., Insacco, G. and Marra, A.C. (1996) Riv. Ital. Pal. Strat. 102(3), 375-384.Google Scholar
  10. Bonfiglio, L., Marra, A.C. and Masini, F. (2000) The contribution of Quaternary vertebrates to paleoenvironmental and paleoclimatological reconstructions in Sicily. In M.B. Hart, (ed.) Climates: Past and Present. Geological Society, London. Special publications, Vol 181, pp. 171-184.Google Scholar
  11. Brommer, J.E. (2000) The evolution of fitness in life-history theory. Biol. Rev. 75, 377-404.Google Scholar
  12. Brown, J.H., Marquet, P.A. and Taper, M.L. (1993) Evolution of body size: consequences of an energetic definition of fitness. Am. Nat. 142, 573-584.Google Scholar
  13. Calder, W. (1996) Size, Function and Life History. Dover Publications, Olineole, New York, USA.Google Scholar
  14. Calef, G.W. (1988) Maximum rate of increase in the African elephant. Afr. J. Ecol. 26, 323-327.Google Scholar
  15. Caloi, L. and Palombo, M.R. (1994) Functional aspects and ecological implications in Pleistocene endemic herbivores of Mediterranean islands. Hist. Biol. 8, 151-172.Google Scholar
  16. Caloi, L., Kotsakis, T., Palombo, M.R. and Petronio, C. (1993) The Pleistocene dwarf elephants of mediterranean islands. In J. Shoshani and P. Tassy, (eds) The Proboscidea: Evolution and Palaeoecology of Elephants and Their Relative. Oxford Science Publications, Oxford, UK.Google Scholar
  17. Case, T.J. (1978) A general explanation for insular body size trends in terrestrial vertebrates. Ecology 59, 1-18.Google Scholar
  18. Charnov, E.L. (1991) Evolution of life history variations among female mammals. Proc. Nat. Acad. Sci. USA 88, 1134-1137.Google Scholar
  19. Charnov, E.L. (2001) Evolution of mammal life histories. Evol. Ecol. Res. 3, 521-535.Google Scholar
  20. Chicon, M. (2001) Diversity of age-specific reproductive rates may result from ageing and optimal resource allocation. J. Evol. Biol. 14, 180-185.Google Scholar
  21. Clutton-Brock, T.H., Albon, S.D. and Guinness, F.E. (1985) Parental investment and sex differences in juvenile mortality in birds and mammals. Nature 313, 331-133.Google Scholar
  22. Damuth, J. (1982) Analysis of the preservation of community structure in assemblage of fossil mammals. Paleobiology 8, 434-446.Google Scholar
  23. Damuth, J. (1993) Cope's rule, the island rule and the scaling of mammalian population density. Nature 365, 748-750.Google Scholar
  24. Damuth, J.D. and MacFadden, B.J. (eds) (1990) Body Size in Mammalian Paleobiology. Cambridge University Press, 397pp.Google Scholar
  25. Dayan, T. and Simberlo., D. (1998) Size patterns among competitors: ecological character displacement and character release in mammals, with special reference to island populations. Mammal Rev. 28(3), 99-124.Google Scholar
  26. Demetrius, L. (2000) Directionality theory and the evolution of body size. Proc. Roy. Soc. Lond. B 267, 2385-2391.Google Scholar
  27. Demment, M.W. and Van Soest, P.J. (1985) A nutritional explanation for body-size patterns of ruminant and nonruminant herbivores. Am. Nat. 125, 641-672.Google Scholar
  28. Dodd, J.R. and Stanton, R.J. (1990) Paleoecology, Concepts and Applications. John Wiley and Sons, New York, USA, 502pp.Google Scholar
  29. Dudley, J.P., Criag, G.C., Gibson, D.St.C., Haynes, G. and Klimowicz, J. (2001) Drought mortality of bush elephants in Hwange National Park, Zimbabwe. Afr. J. Ecol. 39, 187-194.Google Scholar
  30. Ehrlich, P.R. (1986) Which animal will invade? In H.A. Mooney and J.A. Drake, (eds) Ecology of Biological Invasions of North American and Hawaii. Springer Verlag, New York, pp. 79-95.Google Scholar
  31. Foster, J.B. (1964) The evolution of mammals on island. Nature 202, 234-235.Google Scholar
  32. Gasser, M., Kaiser, M., Berrigan, D. and Stearns, S.C. (2000) Life-history correlates of evolution under high and low adult mortality. Evolution 54(4), 1260-1272.Google Scholar
  33. Geffen, E., Gomper, M., Gittleman, J.L., Hang-Kwang Luh, MacDonald, D.W. and Wayne, R.K. (1996) Size, life-history traits, and social organization in the canidae: a reevaluation. Am. Nat. 147(1), 140-160.Google Scholar
  34. Gilinsky, N.L. and Bennington, J.B. (1994) Estimating number of whole individuals from collections of body parts: a taphonomic limitation of the paleontological record. Paleobiology 20(2), 245-258.Google Scholar
  35. Greenwood, P.J. (1980) Mating systems, philopatry, and dispersal in birds and mammals. Anim. Behav. 28, 1140-1162.Google Scholar
  36. Guo, P., Mueller, L.D. and Ayala, F.J. (1991) Evolution of behaviour by density-dependent natural selection. Proc. Nat. Acad. Sci. 88, 10905-10906.Google Scholar
  37. Guthrie, R.D. (2001) Reconstruction of woolly mammoth life history. Proc. 1st International Congress The World of Elephants Rome, 16-20 October 2001.Google Scholar
  38. Harvey, P.H. and Zammuto, R.M. (1985) Patterns of mortality and age at first reproduction in natural populations of mammals. Nature 315, 319-320.Google Scholar
  39. Heaney, L.R. (1978) Island area and body size of insular mammals: evidence from tri-colored squirrel (Callosciurus prevosti) of SouthEast Asia. Evolution 32, 29.Google Scholar
  40. Holliday, R. (1989) Food, reproduction and longevity: is the extended lifespan of calorie.restricted animal an evolutionary adaptation? Bioessays 10, 125-127.Google Scholar
  41. Haynes, G. (1985) Age profiles in elephants and mammoth bone assemblage. Quat. Res. 24, 333-345.Google Scholar
  42. Jachmann, H. (1980) Population dynamics of the elephants in the Kasungu National Park, Malawi. Neth. J. Zool. 30, 622-634.Google Scholar
  43. Jachmann, H., Berry, P.S.M. and Imae, H. (1995) Tusklessness in African elephants: a future trend. Afr. J. Ecol. 33, 230-235.Google Scholar
  44. Janis, C. (1989) A climatic explanation for patterns of diversity in ungulate mammals. Paleontology 32, 463-481.Google Scholar
  45. Jarman, P. (1983) Mating system and sexual dimorphism in large, terrestrial mammalian herbivore. Biol. Rev. 58, 485-520.Google Scholar
  46. Keller, L. and Genoud, M. (1997) Extraordinary lifespans in ants a test of evolutionary theories of ageing. Nature 389, 958-960.Google Scholar
  47. Kelt, D. (1997) Assembly of local communities: consequences of an optimal body size for the organization of competitively structured communities. Biol. J. Linn. Soc. 69, 15-37.Google Scholar
  48. Kelt, D.A. and Brown, J.H. (1998) Diversification of body sizes: patterns and processes in the assembly of terrestrial mammal faunas. In McKinney and J.A. Drake (eds) Biodiversity Dynamics, Turnover of Populations, Taxa and Communities. Columbia UP, New York, USA, pp. 109-132.Google Scholar
  49. Koch, P.L., Hoppe, K.A. and Webb, D.S. (1998) The isotopic ecology of late Pleistocene mammals in North America Part 1. Florida. Chem. Geol. 152, 119-138.Google Scholar
  50. Krukonis, G. and Schaffer, W.M. (1991) Population cycles in mammals and birds: does periodicity scale with body size? J Theoretical Biology 148, 469-493.Google Scholar
  51. Kurten, B. (1954) Population dynamics-a new method in Paleontology. J. Paleont. 28, 286-292.Google Scholar
  52. Laursen, L. and Beko., M. (1978) Loxodonta africana. Mammal. Species 92, 1-8.Google Scholar
  53. Lawlor, T.E. (1982) The evolution of body size in Mammals: evidence from insular population in Mexico. Am. Nat. 119(1), 54-72.Google Scholar
  54. Lister, A.M. (1989) Rapid dwarfing of red deer on Jersey in the last interglacial. Nature 342, 539-542.Google Scholar
  55. Lister, A.M. (1993) Mammoths in miniature. Nature 362, 288.Google Scholar
  56. Lomolino, M.V. (1985) Body size of mammals on islands: the island rule reexamined. Am. Nat. 125, 310-316.Google Scholar
  57. MacArthur, R.H. and Wilson, E.O. (1967) The Theory of Island Biogeography. Princeton University Press, Princeton (NJ), USA.Google Scholar
  58. Mangel, M. and Stamps, J. (2001) Trade-offs between growth and mortality and the manteinance of individual variation in growth. Evol. Ecol. Res. 3, 583-593.Google Scholar
  59. McComb, K., Moss, C., Durant, S., Baker, L. and Sayialel, S. (2001) Matriarchs as repositories of social knowledge in African elephants. Science 292, 491-494.Google Scholar
  60. McFarlane, D.A., MacPhee, R.D.E. and Ford, D.C. (1998) Body size variability and a sangamonian extintion model for Amblyrhiza, a west indian megafaunal rodent. Quat. Res. 50, 80-89.Google Scholar
  61. Moorcroft, P.R., Albon, S.D., Pemberton, J.M., Stevenson, I.R., Mueller, L.D., Guo, P. and Ayala, F.J. (1991) Density-dependent natural selection and trade-offs in life history traits. Science 253, 433-435.Google Scholar
  62. Moss, C.J. (2001) The demography of an African elephant (Loxodonta africana) population in Amboseli, Kenya. J. Zool. Lond. 255, 145-156.Google Scholar
  63. Mueller, L.D., Guo, P. and Ayala, F.J. (1991) Density-dependent natural selection and trade-offs in life history traits. Science 253, 433-435.Google Scholar
  64. Palombo, M.R. (1996) Large Pleistocene mammals of the mediterranean islands. Vie et Milieu 46, 365-374.Google Scholar
  65. Palombo, M.R. (2001) Paedomorphic features and allometric growth in the skull of Elephas falconeri from Spinagallo (Middle Pleistocene, Sicily) Proc. 1st International Congress The World of Elephants, Rome, 16-20 October 2001.Google Scholar
  66. Pianka, E. (1970) On r and k selection. Am. Nat. 104, 592-597.Google Scholar
  67. Pianka, E. (2000) Evolutionaty Ecology. Addison Wesley Inc., San Francisco, USA, 511pp.Google Scholar
  68. Promislow, D.E.L. and Harvey, P.H. (1990) Living fast and dying young: a comparative analysis of life history variation among mammals. J. Zool. Lond. 220, 417-437.Google Scholar
  69. Purvis, A. and Harvey, P.H. (1997) The right size for a mammal. Nature 386, 332-333.Google Scholar
  70. Raubenheimer, E.J. (2000) Development of the tusk and tusklessness in African elephant (Loxodota africana). Koedoe 43(2), 57-64.Google Scholar
  71. Ro., D.A. (2000) Trade-offs between growth and reproduction: an analysis of the quantitative genetic evidence. J. Evol. Biol. 13, 434-445.Google Scholar
  72. Rollo, C.D. (2002) Growth negatively impacts the life span of mammals. Evol. Develop. 4, 55-61.Google Scholar
  73. Rosenzweig, M.L. and McCord, R.D. (1991) Incumbent replacement: evidence for long-term evolutionary progress. Paleobiology 17(3), 202-213.Google Scholar
  74. Roth, V.L. (1984) How elephants grow: heterochrony and the calibtration of developmental stages in some living and fossil species. J. Vert. Paleo. 4(1), 126-145.Google Scholar
  75. Roth, V.L. (1990) Island dwarf elephants: a case study in body mass estimation and ecological inference. In J. Damuth and J.B. MacFadden, (eds) Body size in Mammalian Paleobiology. Cambridge University Press, New York, USA, pp. 51-179.Google Scholar
  76. Roth, V.L. (1992) Inferences from allometry and fossils: dwarfing of elephants on islands. In D. Futuyma and J. Antonovics (eds) Oxford Surveys in Evolutionary Biology, Vol. 8 Oxford University Press, New York, USA.Google Scholar
  77. Roy, K., Jablonski, D. and Martien, K. (2000) Invariant size-frequency distribution along a latitudinal gradient in marine bivalves. Proc. Nat. Acad. Sci. 97, 13150-13155.Google Scholar
  78. Ruggiero, R.G. (1991) Opportunistic predation on elephant calves. Afr. J. Ecol. 29(1), 86-89.Google Scholar
  79. Seydack, A.H.V. and Bigalke, R.C. (1992) Nutritional and life history tactics in the bush pig (Potamochoerus porcus): development of an interactive model. Oecologia 90, 102-112.Google Scholar
  80. Seydack, A.H.V., Vermeulen, C. and Huisamen, J. (2000) Habitat quality and the decline of an african elephant population: implications for conservation. South. Afr. J. Wild Res. 30(1), 34-43.Google Scholar
  81. Shanley, D.P. and Kirchwood, T.B. (2000) Calorie restriction and aging: a life history analysis. Evolution 54(3), 740-750.Google Scholar
  82. Shoshani, J. and Einsenberg, J.F. (1982) Elephas maximus Mammal. Species 182, 1-8.Google Scholar
  83. Sinervo, B., Doughty, P., Huey, R.B. and Zamudio, K. (1992) Allometric engineering: a causal analysis of natural selection on offspring size. Science 258, 1927-1930.Google Scholar
  84. Sinervo, B., Svensson, E. and Comendant, T. (2000) Density cycles and an offspring quantity and quality game driven by natural selection. Nature 606(4069), 85-88.Google Scholar
  85. Sondaar, P.Y. (1977) Insularity and its effect on mammalian evolution. In P.C. Goody, M.K. Hecht and B.M. Hecht (eds) Major Patterns in Vertebrate Evolution. Plenum Press, New York, USA, pp. 671-707.Google Scholar
  86. Stearns, S.C. (1983) The influence of size and phylogeny on patterns of covariation among lifehistory traits in the mammals. Oikos 41, 173-187.Google Scholar
  87. Stearns, S.C., Ackermann, M., Doebeli, M. and Kaiser, M. (2000) Experimental evolution of aging, growth, and reproduction in fruitflies. Proc. Nat. Acad. Sci. 97, 3309-3313.Google Scholar
  88. Stearns, S.C. and Koella, J.C. (1986) The evolution of phenotypic plasticity in life-history traits: predictions of reaction norms for age and size at maturity. Evolution 40(5), 893-913.Google Scholar
  89. Suc, J.P., Bertini, A., Combourieau-Nebout, N., Diniz, F., Leroy, S., Russo-Ermolli, E., Zheng, Z., Bessais, E. and Ferrier, J. (1995) Structure of West Mediterranean vegetation and climate since 5.3 ma. Acta. Zool. Cracov. 38, 3-16.Google Scholar
  90. Tamarin, R.H. (1978) Dispersal, population regulation, and k-selection for field mice. Am. Nat. 112, 554-555.Google Scholar
  91. Tuljiapurkar, S. (1990) Delayed reproduction and fitness in variable environments. Proc. Nat. Acad. Sci. 87, 139-143.Google Scholar
  92. Vartanyan, S.L., Garutt, V.E. and Sher, A.V. (1993) Holocene dwarf mammoths from Wrangel islands in the Siberian artic. Nature 362, 337-340.Google Scholar
  93. Waser, P.M. (1996) Patterns and consequences of dispersal in Gregarious carnivores. In J.L. Gittleman (ed) Carnivore, Behavior, Ecology and Evolution (2). Cornell University Press, Ithaca and London, pp. 267-295.Google Scholar
  94. Western, D. (1979) Size, life history, and ecology in mammals. Afr. J. Ecol. 17, 185-204.Google Scholar
  95. Whitehouse, A.M. (2002) Tusklessness in the elephant population of the Addo Elephant National Park, South Africa. J. Zool. Lond. 257, 249-254.Google Scholar
  96. Whitehouse, A.W. and Hall-Martin, A.J. (2000) Elephants in Addo Elephant National Park, South Africa: reconstruction of the population's history. Oryx 34, 46-55.Google Scholar
  97. Wittemeyer, G. (2001) The elephant population of samburu and Buffalo springs National Reserves, Kenya. Afr. J. Ecol. 39, 357-365.Google Scholar
  98. Zar, J.J. (1984) Biostatistical Analysis. Prentice Hall, Inglewood Cliffs, New Jersey, USA, 718pp.Google Scholar
  99. Zazzo, A., Bocherenes, H., Brunet, M., Beauvilain, A., Billiou, D., Mackaye, H.T., Vignaud, P. and Mariotti, A. (2000) Herbivore paleodiet and paleoenvironmental changes in Chad during the Pliocene using stable isotope ratios of tooth enamel carbonate. Paleobiology 26(2), 294-309.Google Scholar

Copyright information

© Kluwer Academic Publishers 2003

Authors and Affiliations

  • Pasquale Raia
    • 1
    Email author
  • Carmela Barbera
    • 2
  • Maurizio Conte
    • 2
  1. 1.Dip. STATUniversità degli Studi del MoliseIsernia
  2. 2.Dip. Scienza della TerraUniversità Federico II di NapoliNapoliItaly

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