, would like to thank her dogs Flaca and Chacka for thought-provoking walks. This work was supported in part by a Breakthroughs in Gerontology grant to D.E.L.P. from the American Federation of Aging Research

V. N. Anisimov, K. G. Arbeev, I. G. Popovic, M. A. Zabezhinksi, S. V. Rosenfeld et al., Body weight is not always a good predictor of longevity in mice, Experimental Gerontology, vol.39, pp.305-319, 2004.

S. N. Austad, Animal size, metabolic rate, and survival, among and within species, The comparative biology of aging, vol.126, pp.27-41, 2005.

A. Bartke, K. Coshigano, J. Kopchick, V. Chandrashekar, J. Mattison et al., Genes that prolong life: relationships of growth hormone and growth to aging and life span, Journals of Gerontology Series A: Biological Sciences and Medical Sciences, vol.56, pp.340-349, 2001.

A. Bartke, J. C. Wright, J. A. Mattison, D. K. Ingram, R. A. Miller et al., Longevity: extending the lifespan of longlived mice, Nature, vol.414, p.412, 2001.

D. E. Berryman, J. S. Christiansen, G. Johannsson, M. O. Thorner, and J. J. Kopchick, Role of the GH/IGF-1 axis in lifespan and healthspan: lessons from animal models, Growth Hormone and IGF Research, vol.18, pp.455-471, 2008.

W. U. Blanckenhorn, The evolution of body size: what keeps organisms small?, Quarterly Review of Biology, vol.75, pp.385-407, 2000.

B. N. Bonnett, A. Egenvall, A. Hedhammar, and P. Olson, Mortality in over 350,000 insured Swedish dogs from, 1995.

I. , Breed-, gender-, age-and cause-specific rates, Acta Veterinaria Scandinavica, vol.46, pp.105-120

M. B. Bonsall, Longevity and ageing: appraising the evolutionary consequences of growing old, Philosophical Transactions of the Royal Society B: Biological Sciences, vol.361, pp.119-135, 2006.

A. R. Boyko, P. Quignon, L. Li, J. J. Schoenebeck, J. D. Degenhardt et al., A simple genetic architecture underlies morphological variation in dogs, PLoS Biology, vol.8, p.1000451, 2010.
URL : https://hal.archives-ouvertes.fr/inserm-00512737

K. P. Burnham and D. R. Anderson, Model selection and multimodel inference: a practical information-theoretic approach, 2002.

E. L. Charnov, Life history invariants, 1993.

E. L. Charnov and D. Berrigan, Dimensionless numbers and the assembly rules for life histories, Philosophical Transactions of the Royal Society B: Biological Sciences, vol.332, pp.41-48, 1991.

D. Chen, K. Z. Pan, J. E. Palter, and P. Kapahi, Longevity determined by developmental arrest genes in Caenorhabditis elegans, Aging Cell, vol.6, pp.525-533, 2007.

M. Cichon, Evolution of longevity through optimal resource allocation, Proceedings of the Royal Society B: Biological Sciences, vol.264, pp.1383-1388, 1997.

D. M. Cooley, D. L. Schlittler, L. T. Glickman, M. Hayek, and D. J. Waters, Exceptional longevity in pet dogs is accompanied by cancer resistance and delayed onset of major diseases, Journals of Gerontology Series A: Biological Sciences and Medical Sciences, vol.58, pp.1078-1084, 2003.

K. Dammrich, Relationship between nutrition and bonegrowth in large and giant dogs, Journal of Nutrition, vol.121, pp.114-121, 1991.

T. Davis and D. Kipling, Telomeres and telomerase biology in vertebrates: progress towards a non-human model for replicative senescence and ageing, Biogerontology, vol.6, pp.371-385, 2005.

J. P. De-magalhaes, J. Costa, and G. M. Church, An analysis of the relationship between metabolism, developmental schedules, and longevity using phylogenetic independent contrasts, Journals of Gerontology Series A: Biological Sciences and Medical Sciences, vol.62, pp.149-160, 2007.

M. Desai and C. N. Hales, Role of fetal and infant growth in programming metabolism in later life, Biological Reviews of the Cambridge Philosophical Society, vol.72, pp.329-348, 1997.

M. D. Drapeau, E. K. Gass, M. D. Simison, L. D. Mueller, and M. R. Rose, Testing the heterogeneity theory of late-life mortality plateaus by using cohorts of Drosophila melanogaster, Experimental Gerontology, vol.35, pp.71-84, 2000.

J. E. Eigenmann, A. Amador, and D. F. Patterson, Insulin-like growth factor-1 levels in proportionate dogs, chondrodystrophic dogs and giant dogs, Acta Endocrinologica, vol.118, pp.105-108, 1988.

R. P. Favier, J. A. Mol, H. S. Kooistra, and A. Rijnberk, Large body size in the dog is associated with transient GH excess at a young age, Journal of Endocrinology, vol.170, pp.479-484, 2001.

C. E. Finch and E. M. Crimmins, Inflammatory exposure and historical changes in human life-spans, Science, vol.305, pp.1736-1739, 2004.

J. M. Fleming, K. E. Creevy, and D. E. Promislow, Mortality in North American dogs from 1984 to 2004: an investigation into age-, size-, and breed-related causes of death, Journal of Veterinary Internal Medicine, vol.25, pp.187-198, 2011.

J. Gaillard, M. Festa-bianchet, D. Delorme, and J. Jorgenson, Body mass and individual fitness in female ungulates: bigger is not always better, Proceedings of the Royal Society B: Biological Sciences, vol.267, pp.471-477, 2000.
URL : https://hal.archives-ouvertes.fr/hal-00427083

J. Gaillard, D. Pontier, D. Allainé, J. D. Lebreton, J. Trouvilliez et al., An analysis of demographic tactics in birds and mammals, Oikos, vol.56, pp.59-76, 1989.

F. Galis, I. Van-der, T. J. Sluijs, J. A. Van-dooren, M. Metz et al., Do large dogs die young, Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, vol.308, pp.119-126, 2007.

B. Gompertz, On the nature of the function expressive of the law of human mortality, and on the mode of determining the value of life contingencies, Philosophical Transactions of the Royal Society, vol.115, pp.513-585, 1825.

K. Greer, L. Hughes, and M. Masternak, Connecting serum IGF-1, body size, and age in the domestic dog, Age, vol.33, pp.475-483, 2011.

K. A. Greer, S. C. Canterberry, and K. E. Murphy, Statistical analysis regarding the effects of height and weight on life span of the domestic dog, Research in Veterinary Science, vol.82, pp.208-214, 2007.

P. H. Harvey and M. D. Pagel, The comparative method in evolutionary biology, 1991.

A. J. Hawthorne, D. Booles, P. A. Nugent, G. Gettinby, and J. Wilkinson, Body-weight changes during growth in puppies of different breeds, Journal of Nutrition, vol.134, pp.2027-2030, 2004.

O. R. Jones, J. M. Gaillard, S. Tuljapurkar, J. S. Alho, K. B. Armitage et al., Senescence rates are determined by ranking on the fast-slow life-history continuum, Ecology Letters, vol.11, pp.664-673, 2008.
URL : https://hal.archives-ouvertes.fr/hal-00318567

D. A. Juckett and B. Rosenberg, Comparison of the Gompertz and Weibull functions as descriptors for human mortality distributions and their intersections, Mechanisms of Ageing and Development, vol.69, pp.1-31, 1993.

H. M. Kang, N. A. Zaitlen, C. M. Wade, A. Kirby, D. Heckerman et al., Efficient control of population structure in model organism association mapping, Genetics, vol.178, pp.1709-1723, 2008.

H. S. Kaplan and A. J. Robson, We age because we grow, Proceedings of the Royal Society B: Biological Sciences, vol.276, pp.1837-1844, 2009.

C. Kenyon, The plasticity of aging: insights from long-lived mutants, Cell, vol.120, pp.449-460, 2005.

A. A. Khazaeli, W. J. Van-voorhies, and W. Curtsinger, The relationship between life span and adult body size is highly strainspecific in Drosophila melanogaster, Experimental Gerontology, vol.40, pp.377-385, 2005.

J. K. Kirkwood, The influence of size on the biology of the dog, Journal of Small Animal Practice, vol.26, pp.97-110, 1985.

T. B. Kirkwood, Understanding the odd science of aging, Cell, vol.120, pp.437-447, 2005.

J. P. Klein and M. L. Moeschberger, Survival analysis: techniques for censored and truncated data, 2005.

D. A. Levitis, Before senescence: the evolutionary demography of ontogenesis, Proceedings of the Royal Society B: Biological Sciences, vol.278, pp.801-809, 2011.

Y. Li, B. Deeb, W. Pendergrass, and N. Wolf, Cellular proliferative capacity and life span in small and large dogs, 1996.

, Gerontology Series A: Biological Sciences and Medical Sciences, vol.51, pp.403-408

K. Lindblad-toh, C. M. Wade, T. S. Mikkelsen, E. K. Karlsson, D. B. Jaffe et al., Genome sequence, comparative analysis and haplotype structure of the domestic dog, Nature, vol.438, pp.803-819, 2005.

N. B. Metcalfe and P. Monaghan, Growth versus lifespan: perspectives from evolutionary ecology, Experimental Gerontology, vol.38, pp.935-940, 2003.

A. R. Michell, Longevity of British breeds of dog and its relationships with sex, size, cardiovascular variables and disease, Veterinary Record, vol.145, pp.625-629, 1999.

R. A. Miller, C. Chrisp, and W. Atchley, Differential longevity in mouse stocks selected for early life growth trajectory, Journals of Gerontology Series A: Biological Sciences and Medical Sciences, vol.55, pp.455-461, 2000.

R. A. Miller, J. M. Harper, A. Galecki, and D. T. Burke, Big mice die young: early life body weight predicts longevity in genetically heterogeneous mice, Aging Cell, vol.1, pp.22-29, 2002.

P. Monaghan, A. Charmantier, D. H. Nussey, and R. E. Ricklefs, The evolutionary ecology of senescence, Functional Ecology, vol.22, pp.371-378, 2008.

J. A. Moody, L. A. Clark, and K. E. Murphy, Canine history and breed clubs. Pages 1-18 in, The dog and its genome, 2006.

J. A. Moorad, D. E. Promislow, N. Flesness, and R. A. Miller, A comparative assessment of univariate longevity measures using zoological animal records, Aging Cell, vol.11, pp.940-948, 2012.

I. J. Morgan, I. D. Mccarthy, and N. B. Metcalfe, Life-history strategies and protein metabolism in overwintering juvenile Atlantic salmon: growth is enhanced in early migrants through lower protein turnover, Journal of Fish Biology, vol.56, pp.637-647, 2000.

L. Nasir, P. Devlin, T. Mckevitt, G. Rutteman, and D. J. Argyle, Telomere lengths and telomerase activity in dog tissues: a potential model system to study human telomere and telomerase biology, Neoplasia, vol.3, pp.351-359, 2001.

J. A. Nelder and R. Mead, A simplex algorithm for function minimization, Computer Journal, vol.7, pp.308-313, 1965.

M. Olsson and R. Shine, Growth to death in lizards, Evolution, vol.56, pp.1867-1870, 2002.

H. G. Parker, L. V. Kim, N. B. Sutter, S. Carlson, T. D. Lorentzen et al., Genetic structure of the purebred domestic dog, Science, vol.304, pp.1160-1164, 2004.

L. Partridge and N. H. Barton, On measuring the rate of ageing, Proceedings of the Royal Society B: Biological Sciences, vol.263, pp.1365-1371, 1996.

G. J. Patronek, D. J. Waters, and L. T. Glickman, Comparative longevity of pet dogs and humans: implications for gerontology research, Journals of Gerontology Series A: Biological Sciences and Medical Sciences, vol.52, pp.171-178, 1997.

G. Peron, O. Gimenez, A. Charmantier, J. M. Gaillard, and P. A. Crochet, Age at the onset of senescence in birds and mammals is predicted by early-life performance, Proceedings of the Royal Society B: Biological Sciences, vol.277, pp.2849-2856, 2010.
URL : https://hal.archives-ouvertes.fr/hal-00698129

S. D. Pletcher and J. W. Curtsinger, Mortality plateaus and the evolution of senescence: why are old-age mortality rates so low, Evolution, vol.52, pp.454-464, 1998.

D. E. Promislow, Senescence in natural populations of mammals: a comparative study, vol.45, pp.1869-1887, 1991.

D. E. Promislow and P. H. Harvey, Living fast and dying young: a comparative analysis of life-history variation among mammals, Journal of Zoology, vol.220, pp.417-437, 1990.

D. E. Promislow, M. Tatar, S. Pletcher, and J. R. Carey, Below-threshold mortality: implications for studies in evolution, ecology and demography, Journal of Evolutionary Biology, vol.12, pp.314-328, 1999.

H. F. Proschowsky, H. Rugbjerg, and A. K. Ersboll, Mortality of purebred and mixed-breed dogs in Denmark, Preventive Veterinary Medicine, vol.58, pp.63-74, 2003.

R. Development-core and . Team, R: a language and environment for statistical computing. R Foundation for Statistical Computing, 2011.

D. Reznick, L. Nunney, and A. Tessier, Big houses, big cars, superfleas and the costs of reproduction, Trends in Ecology and Evolution, vol.15, pp.421-425, 2000.

R. E. Ricklefs, Embryo development and ageing in birds and mammals, Proceedings of the Royal Society B: Biological Sciences, vol.273, pp.2077-2082, 2006.

R. E. Ricklefs and A. Scheuerlein, Biological implications of the Weibull and Gompertz models of aging, Journals of Gerontology Series A: Biological Sciences and Medical Sciences, vol.36, pp.69-76, 2001.

C. D. Rollo, Growth negatively impacts the life span of mammals, Evolution and Development, vol.4, pp.55-61, 2002.

T. J. Samaras, Should we be concerned over increasing body height and weight?, Experimental Gerontology, vol.44, pp.83-92, 2009.

J. R. Sauer and N. A. Slade, Uinta ground squirrel demography: is body mass a better categorical variable than age, Ecology, vol.68, pp.642-650, 1987.

W. Siler, A competing-risk model for animal mortality, Ecology, vol.60, pp.750-757, 1979.

J. R. Speakman, A. Van-acker, and E. J. Harper, Age-related changes in the metabolism and body composition of three dog breeds and their relationship to life expectancy, Aging Cell, vol.2, pp.265-275, 2003.

N. Sutter, D. Mosher, M. Gray, and E. Ostrander, Morphometrics within dog breeds are highly reproducible and dispute Rensch's rule, Mammalian Genome, vol.19, pp.713-723, 2008.

N. B. Sutter, C. D. Bustamante, K. Chase, M. M. Gray, K. Y. Zhao et al., A single IGF1 allele is a major determinant of small size in dogs, Science, vol.316, pp.112-115, 2007.

N. B. Sutter and E. A. Ostrander, Dog star rising: the canine genetic system, Nature Reviews Genetics, vol.5, pp.900-910, 2004.

M. Tatar, A. Bartke, and A. Antebi, The endocrine regulation of aging by insulin-like signals, Science, vol.299, pp.1346-1351, 2003.

Y. Teuschl, C. Reim, and W. U. Blanckenhorn, Correlated response to artificial selection in growth, development, phenotypic plasticity and juvenile viability in yellow dung flies, Journal of Evolutionary Biology, vol.20, pp.87-103, 2006.

A. R. Thatcher, V. Kannisto, and J. W. Vaupel, Trade-off between growth and immune function: a metaanalysis of selection experiments, Functional Ecology, vol.25, pp.74-80, 1998.

J. W. Vaupel and A. I. Yashin, Heterogeneity ruses: some surprising effects of selection on population dynamics, American Statistician, vol.39, pp.176-185, 1985.

W. Weibull, A statistical distribution function of wide applicability, Journal of Applied Mechanics, vol.18, pp.293-297, 1951.

P. D. Williams, T. Day, Q. Fletcher, and L. Rowe, The shaping of senescence in the wild, Trends in Ecology and Evolution, vol.21, pp.458-463, 2006.

D. L. Wilson, The analysis of survival (mortality) data: fitting Gompertz, Weibull, and logistic functions, Mechanisms of Ageing and Development, vol.74, pp.15-33, 1994.