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.
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Abe, E. (1996) Tusklessness among the Queen Elizabeth National Park elephants, Uganda. Pachyderm 22, 46-47.
Adler, G.H. and Levins, R. (1994) The island syndrome in rodent populations. Quart. Rev. Biol. 69(4), 473-490.
Alcover, J.A. and McMinn, M. (1994) Bioscience 44, 12-19.
Ambrosetti, P. (1968) The Pleistocene dwarf elephants of Spinagallo (Siracusa, South-Eastern Sicily). Geol. Rom. 7, 277-398.
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.
Belluomini, G. and Bada, J.L. (1985) Isoleucene epimerization ages of the dwarf elephants of Sicily. Geology 13, 451-452.
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.
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.
Bonfiglio, L., Di Geronimo, S., Insacco, G. and Marra, A.C. (1996) Riv. Ital. Pal. Strat. 102(3), 375-384.
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.
Brommer, J.E. (2000) The evolution of fitness in life-history theory. Biol. Rev. 75, 377-404.
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.
Calder, W. (1996) Size, Function and Life History. Dover Publications, Olineole, New York, USA.
Calef, G.W. (1988) Maximum rate of increase in the African elephant. Afr. J. Ecol. 26, 323-327.
Caloi, L. and Palombo, M.R. (1994) Functional aspects and ecological implications in Pleistocene endemic herbivores of Mediterranean islands. Hist. Biol. 8, 151-172.
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.
Case, T.J. (1978) A general explanation for insular body size trends in terrestrial vertebrates. Ecology 59, 1-18.
Charnov, E.L. (1991) Evolution of life history variations among female mammals. Proc. Nat. Acad. Sci. USA 88, 1134-1137.
Charnov, E.L. (2001) Evolution of mammal life histories. Evol. Ecol. Res. 3, 521-535.
Chicon, M. (2001) Diversity of age-specific reproductive rates may result from ageing and optimal resource allocation. J. Evol. Biol. 14, 180-185.
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.
Damuth, J. (1982) Analysis of the preservation of community structure in assemblage of fossil mammals. Paleobiology 8, 434-446.
Damuth, J. (1993) Cope's rule, the island rule and the scaling of mammalian population density. Nature 365, 748-750.
Damuth, J.D. and MacFadden, B.J. (eds) (1990) Body Size in Mammalian Paleobiology. Cambridge University Press, 397pp.
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.
Demetrius, L. (2000) Directionality theory and the evolution of body size. Proc. Roy. Soc. Lond. B 267, 2385-2391.
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.
Dodd, J.R. and Stanton, R.J. (1990) Paleoecology, Concepts and Applications. John Wiley and Sons, New York, USA, 502pp.
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.
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.
Foster, J.B. (1964) The evolution of mammals on island. Nature 202, 234-235.
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.
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.
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.
Greenwood, P.J. (1980) Mating systems, philopatry, and dispersal in birds and mammals. Anim. Behav. 28, 1140-1162.
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.
Guthrie, R.D. (2001) Reconstruction of woolly mammoth life history. Proc. 1st International Congress The World of Elephants Rome, 16-20 October 2001.
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.
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.
Holliday, R. (1989) Food, reproduction and longevity: is the extended lifespan of calorie.restricted animal an evolutionary adaptation? Bioessays 10, 125-127.
Haynes, G. (1985) Age profiles in elephants and mammoth bone assemblage. Quat. Res. 24, 333-345.
Jachmann, H. (1980) Population dynamics of the elephants in the Kasungu National Park, Malawi. Neth. J. Zool. 30, 622-634.
Jachmann, H., Berry, P.S.M. and Imae, H. (1995) Tusklessness in African elephants: a future trend. Afr. J. Ecol. 33, 230-235.
Janis, C. (1989) A climatic explanation for patterns of diversity in ungulate mammals. Paleontology 32, 463-481.
Jarman, P. (1983) Mating system and sexual dimorphism in large, terrestrial mammalian herbivore. Biol. Rev. 58, 485-520.
Keller, L. and Genoud, M. (1997) Extraordinary lifespans in ants a test of evolutionary theories of ageing. Nature 389, 958-960.
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.
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.
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.
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.
Kurten, B. (1954) Population dynamics-a new method in Paleontology. J. Paleont. 28, 286-292.
Laursen, L. and Beko., M. (1978) Loxodonta africana. Mammal. Species 92, 1-8.
Lawlor, T.E. (1982) The evolution of body size in Mammals: evidence from insular population in Mexico. Am. Nat. 119(1), 54-72.
Lister, A.M. (1989) Rapid dwarfing of red deer on Jersey in the last interglacial. Nature 342, 539-542.
Lister, A.M. (1993) Mammoths in miniature. Nature 362, 288.
Lomolino, M.V. (1985) Body size of mammals on islands: the island rule reexamined. Am. Nat. 125, 310-316.
MacArthur, R.H. and Wilson, E.O. (1967) The Theory of Island Biogeography. Princeton University Press, Princeton (NJ), USA.
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.
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.
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.
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.
Moss, C.J. (2001) The demography of an African elephant (Loxodonta africana) population in Amboseli, Kenya. J. Zool. Lond. 255, 145-156.
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.
Palombo, M.R. (1996) Large Pleistocene mammals of the mediterranean islands. Vie et Milieu 46, 365-374.
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.
Pianka, E. (1970) On r and k selection. Am. Nat. 104, 592-597.
Pianka, E. (2000) Evolutionaty Ecology. Addison Wesley Inc., San Francisco, USA, 511pp.
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.
Purvis, A. and Harvey, P.H. (1997) The right size for a mammal. Nature 386, 332-333.
Raubenheimer, E.J. (2000) Development of the tusk and tusklessness in African elephant (Loxodota africana). Koedoe 43(2), 57-64.
Ro., D.A. (2000) Trade-offs between growth and reproduction: an analysis of the quantitative genetic evidence. J. Evol. Biol. 13, 434-445.
Rollo, C.D. (2002) Growth negatively impacts the life span of mammals. Evol. Develop. 4, 55-61.
Rosenzweig, M.L. and McCord, R.D. (1991) Incumbent replacement: evidence for long-term evolutionary progress. Paleobiology 17(3), 202-213.
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.
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.
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.
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.
Ruggiero, R.G. (1991) Opportunistic predation on elephant calves. Afr. J. Ecol. 29(1), 86-89.
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.
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.
Shanley, D.P. and Kirchwood, T.B. (2000) Calorie restriction and aging: a life history analysis. Evolution 54(3), 740-750.
Shoshani, J. and Einsenberg, J.F. (1982) Elephas maximus Mammal. Species 182, 1-8.
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.
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.
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.
Stearns, S.C. (1983) The influence of size and phylogeny on patterns of covariation among lifehistory traits in the mammals. Oikos 41, 173-187.
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.
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.
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.
Tamarin, R.H. (1978) Dispersal, population regulation, and k-selection for field mice. Am. Nat. 112, 554-555.
Tuljiapurkar, S. (1990) Delayed reproduction and fitness in variable environments. Proc. Nat. Acad. Sci. 87, 139-143.
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.
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.
Western, D. (1979) Size, life history, and ecology in mammals. Afr. J. Ecol. 17, 185-204.
Whitehouse, A.M. (2002) Tusklessness in the elephant population of the Addo Elephant National Park, South Africa. J. Zool. Lond. 257, 249-254.
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.
Wittemeyer, G. (2001) The elephant population of samburu and Buffalo springs National Reserves, Kenya. Afr. J. Ecol. 39, 357-365.
Zar, J.J. (1984) Biostatistical Analysis. Prentice Hall, Inglewood Cliffs, New Jersey, USA, 718pp.
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.
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Raia, P., Barbera, C. & Conte, M. The fast life of a dwarfed giant. Evolutionary Ecology 17, 293–312 (2003). https://doi.org/10.1023/A:1025577414005
- island rule
- life history
- resource allocation