Empirical studies reveal aging occurs in wild populations. Consideration of the ecological and evolutionary consequences of these findings is critical for many areas of research, including life-history evolution, sexual selection, behavior, and applied ecology. Variation in the patterns of age-dependent declines of phenotypic traits has been found both within and among individuals, and this raises future questions aimed at understanding what determines these trajectories across traits and across the tree of life. The presence of older, aging, individuals in populations can have transgenerational effects on offspring and can influence how individuals interact. In some species older individuals in populations can have positive impacts, influencing knowledge and leadership, postreproductive care, and population cycle stabilization. Aging and long life span also need to be recognized in an applied ecology context including management plans, vector-borne disease transmission, and ecotoxicology.


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Literature Cited

  1. Abrams PA. 1993. Does increased mortality favor the evolution of more rapid senescence?. Evolution 47:877–87 [Google Scholar]
  2. Aguilar A, Borrell A. 1994. Reproductive transfer and variation of body load of organochlorine pollutants with age in fin whales (Balaenoptera physalus). Arch. Environ. Contam. Toxicol. 27:546–54 [Google Scholar]
  3. Archie EA, Chiyo PI. 2012. Elephant behaviour and conservation: social relationships, the effects of poaching, and genetic tools for management. Mol. Ecol. 21:765–78 [Google Scholar]
  4. Balbontin J, De Lope F, Hermosell IG, Mousseau TA, Møller AP. 2011. Determinants of age-dependent change in a secondary sexual character. J. Evol. Biol. 24:440–48 [Google Scholar]
  5. Balbontin J, Hermosell IG, Marzal A, Reviriego M, De Lope F, Møller AP. 2007. Age-related change in breeding performance in early life is associated with an increase in competence in the migratory barn swallow Hirundo rustica. J. Anim. Ecol. 76:915–25 [Google Scholar]
  6. Balbontin J, Møller AP, Hermosell IG, Marzal A, Reviriego M, de Lope F. 2012. Geographical variation in reproductive ageing patterns and life-history strategy of a short-lived passerine bird. J. Evol. Biol. 25:2298–309 [Google Scholar]
  7. Barrett ELB, Burke TA, Hammers M, Komdeur J, Richardson DS. 2013. Telomere length and dynamics predict mortality in a wild longitudinal study. Mol. Ecol. 22:249–59 [Google Scholar]
  8. Baudisch A. 2005. Hamilton's indicators of the force of selection. Proc. Natl. Acad. Sci. USA 102:8263–68 [Google Scholar]
  9. Baudisch A, Salguero-Gómez R, Jones OR, Wrycza T, Mbeau-Ache C. et al. 2013. The pace and shape of senescence in angiosperms. J. Ecol. 101:596–606 [Google Scholar]
  10. Beamonte-Barrientos R, Velando A, Drummond H, Torres R. 2010. Senescence of maternal effects: aging influences egg quality and rearing capacities of a long-lived bird. Am. Nat. 175:469–80 [Google Scholar]
  11. Begon M. 1976. Temporal variations in the reproductive condition of Drosophila obscura Fallén and D. subobscura Collin. Oecologia 23:31–47 [Google Scholar]
  12. Berman M, Gaillard JM, Weimerskirch H. 2009. Contrasted patterns of age-specific reproduction in long-lived seabirds. Proc. R. Soc. B 276:375–82 [Google Scholar]
  13. Berube CH, Festa-Bianchet M, Jorgenson JT. 1999. Individual differences, longevity, and reproductive senescence in bighorn ewes. Ecology 80:2555–65 [Google Scholar]
  14. Bize P, Criscuolo F, Metcalfe NB, Nasir L, Monaghan P. 2009. Telomere dynamics rather than age predict life expectancy in the wild. Proc. R. Soc. B 276:1679–83 [Google Scholar]
  15. Blest AD. 1963. Longevity, palatability and natural selection in five species of New World Saturniid moth. Nature 197:1183–86 [Google Scholar]
  16. Bonduriansky R, Maklakov A, Zajitschek F, Brooks R. 2008. Sexual selection, sexual conflict and the evolution of ageing and life span. Funct. Ecol. 22:443–53 [Google Scholar]
  17. Bouwhuis S, Charmantier A, Verhulst S, Sheldon BC. 2010. Trans-generational effects on ageing in a wild bird population. J. Evol. Biol. 23:636–42 [Google Scholar]
  18. Bouwhuis S, Choquet R, Sheldon BC, Verhulst S. 2012. The forms and fitness cost of senescence: age-specific recapture, survival, reproduction, and reproductive value in a wild bird population. Am. Nat. 179:E15–27 [Google Scholar]
  19. Brockmann HJ. 1997. Cooperative breeding in wasps and vertebrates: the role of ecological constraints. Social Behavior in Insects and Arachnids JC Choe, BJ Crespi 347–71 New York: Cambridge Univ. Press [Google Scholar]
  20. Brommer JE, Rattiste K, Wilson A. 2010. The rate of ageing in a long-lived bird is not heritable. Heredity 104:363–70 [Google Scholar]
  21. Bronikowski AM, Promislow DEL. 2005. Testing evolutionary theories of aging in wild populations. Trends Ecol. Evol. 20:271–73 [Google Scholar]
  22. Carey JR. 2001. Insect biodemography. Annu. Rev. Entomol. 46:79–110 [Google Scholar]
  23. Carey JR, Gruenfelder CA. 1997. Population biology of the elderly. Biodemography of Longevity K Wachter, C Finch 127–60 Washington, DC: Natl. Acad. [Google Scholar]
  24. Carey JR, Judge DS. 2001. Life span extension in humans is self-reinforcing: a general theory of longevity. Popul. Dev. Rev. 27:411–36 [Google Scholar]
  25. Carey JR, Papadopoulos NT, Papanastasiou S, Diamanditis A, Nakas CT. 2012. Estimating changes in mean population age using the death distributions of live-captured medflies. Ecol. Entomol. 37:359–69 [Google Scholar]
  26. Carlson SM, Hilborn R, Hendry AP, Quinn TP. 2007. Predation by bears drives senescence in natural populations of salmon. PLOS ONE 2:e1286 [Google Scholar]
  27. Catry P, Phillips RA, Phalan B, Croxall JP. 2006. Senescence effects in an extremely long-lived bird: the grey-headed albatross Thalassarche chrysostoma. Proc. R. Soc. B 273:1625–30 [Google Scholar]
  28. Chu C, Adler PB. 2014. When should plant population models include age structure?. J. Ecol. 102:531–43 [Google Scholar]
  29. Clutton-Brock TH. 1991. The Evolution of Parental Care Princeton, NJ: Princeton Univ. Press352
  30. Cohen AA, McGraw KJ. 2009. No simple measures for antioxidant status in birds: complexity in inter- and intraspecific correlations among circulating antioxidant types. Funct. Ecol. 23:310–20 [Google Scholar]
  31. Congdon JD, Gibbons JW, Brooks RJ, Rollinson N, Tsaliagos RN. 2013. Indeterminate growth in long-lived freshwater turtles as a component of individual fitness. Evol. Ecol. 27:445–59 [Google Scholar]
  32. Cosantini D, Dell'Ariccia G, Lipps H. 2008. Long flights and age affect oxidative status in homing pigeons (Columba livia). J. Exp. Biol. 211:377–81 [Google Scholar]
  33. Coulson T, Catchpole EA, Albon SD, Morgan BJ, Pemberton JM. et al. 2001. Age, sex, density, winter weather, and population crashes in Soay sheep. Science 292:1528–31 [Google Scholar]
  34. Dean R, Cornwallis CK, Lovlie H, Worley K, Richardson DS, Pizzari T. 2010. Male reproductive senescence causes potential for sexual conflict over mating. Curr. Biol. 20:1192–96 [Google Scholar]
  35. Descamps S, Boutin S, Berteaux D, Gaillard JM. 2008. Age-specific variation in survival, reproductive success and offspring quality in red squirrels: evidence of senescence. Oikos 117:1406–16 [Google Scholar]
  36. DiSilvestro RL. 1991. The hidden world of the African elephant. The African Elephant: Twilight in Eden CN Palmer 96–127 New York: Wiley [Google Scholar]
  37. Dugdale HL, Pope LC, Newman C, Macdonald DW, Burke T. 2011. Age-specific breeding success in a wild mammalian population: selection, constraint, restraint and senescence. Mol. Ecol. 20:3261–74 [Google Scholar]
  38. Ericsson G, Wallin K. 2001. Age-specific moose (Alces alces) mortality in a predator-free environment: evidence for senescence in females. Ecoscience 8:157–63 [Google Scholar]
  39. Evans HE. 1958. The evolution of social life in wasps. Proc. 10th Int. Congr. Entomol., Montreal, Aug. 17–25, 1956449–57 Ottawa: Mortimer [Google Scholar]
  40. Faulkes CG, Abbott DH. 1991. Social control of reproduction in breeding and non-breeding male naked mole-rats (Heterocephalus glaber). J. Reprod. Fert 93:427–35 [Google Scholar]
  41. Field J, Shreeves G, Sumner S, Casiraghi M. 2000. Insurance-based advantage to helpers in a tropical hover wasp. Nature 404:869–71 [Google Scholar]
  42. Field JG, Moloney CI, du Buisson L, Jarre A, Stroemme T. et al. 2008. Exploring the BOFFFF hypothesis using a model of southern African deepwater hake (Merluccius paradoxus). Proc. Fish. Glob. Welf. Environ. 5th World Fish. Congr. K Tsukamoto, T Kawamura, T Takeuchi, TD Beard Jr, MJ Kaiser 17–26 Tokyo: TERRAPUB [Google Scholar]
  43. Finch CE. 2009. Update on slow aging and negligible senescence—a mini-review. Gerontology 55:307–13 [Google Scholar]
  44. Finkelstein ME, Doak DF, Nakagawa M, Sievert PR, Klavitter J. 2010. Assessment of demographic risk factors and management priorities: Impacts on juveniles substantially affect population viability of a long-lived seabird. Anim. Conserv. 13:148–56 [Google Scholar]
  45. Foote AD. 2008. Mortality rate acceleration and post-reproductive lifespan in matrilineal whale species. Biol. Lett. 4:189–91 [Google Scholar]
  46. Foster EA, Franks DW, Mazzi S, Darden SK, Balcomb KC. et al. 2012. Adaptive prolonged postreproductive life span in killer whales. Science 337:1313 [Google Scholar]
  47. Freeman D. 1980. The living elephant. Elephants: The Vanishing Giants53–85 Hong Kong: Bison Books [Google Scholar]
  48. Fuller TK, Keith LB. 1980. Wolf population dynamics and prey relationships in Northeastern Alberta. J. Wildl. Manag. 44:583–602 [Google Scholar]
  49. Garcia MB, Dahlgren JP, Ehrlén J. 2011. No evidence of senescence in a 300-year-old mountain herb. J. Ecol. 99:1424–30 [Google Scholar]
  50. Gasparini C, Marino IAM, Boschetto C, Pilastro A. 2010. Effect of male age on sperm traits and sperm competition success in the guppy (Poecilia reticulata). J. Evol. Biol. 23:124–35 [Google Scholar]
  51. Goldwasser L. 2004. The biodemography of life span: resources, allocation and metabolism. Trends Ecol. Evol. 16:536–38 [Google Scholar]
  52. Grimaldi D, Engel MS. 2005. Evolution of the Insects Cambridge, UK: Cambridge Univ. Press
  53. Gurven M, Kaplan H. 2007. Longevity among hunter-gatherers: a cross-cultural examination. Popul. Dev. Rev. 33:321–65 [Google Scholar]
  54. Hadley GL, Rotella JJ, Garrott RA. 2007. Influence of maternal characteristics and oceanographic conditions on survival and recruitment probabilities of Weddell seals. Oikos 116:601–13 [Google Scholar]
  55. Hamilton WD. 1966. The moulding of senescence by natural selection. J. Theor. Biol. 12:12–45 [Google Scholar]
  56. Hammers M, Richardson DS, Burke T, Komdeur J. 2012. Age-dependent terminal declines in reproductive output in a wild bird. PLOS ONE 7:e40413 [Google Scholar]
  57. Harcourt AH, Stewart KJ. 2007. Gorilla society: what we know and don't know. Evol. Anthropol. 16:147–58 [Google Scholar]
  58. Hastings A. 1984. Age-dependent predation is not a simple process. II. Wolves, ungulates, and a discrete time model for predation on juveniles with a stabilizing tail. Theor. Popul. Biol. 26:271–82 [Google Scholar]
  59. Hayward AD, Wilson AJ, Pilkington JG, Clutton-Brock TH, Pemberton JM. et al. 2013. Reproductive senescence in female Soay sheep: variation across traits and contributions of individual ageing and selective disappearance. Funct. Ecol. 27:184–95 [Google Scholar]
  60. Hixon MA, Johnson DW, Sogard SM. 2013. BOFFFFs: on the importance of conserving old-growth age structure in fishery populations. J. Mar. Sci. In press. doi: 10.1093/icesjms/fst200
  61. Hrdy SB. 1981. “Nepotists” and “altruists”: the behavior of old females among macaques and langur monkeys. Other Ways of Growing Old: Anthropological Perspectives PT Amoss, S Harrell 59–76 Stanford, CA: Stanford Univ. Press [Google Scholar]
  62. Izquierdo JI. 1991. How does Drosophila melanogaster overwinter?. Entomol. Exp. Appl. 59:51–58 [Google Scholar]
  63. Johnson CM, Norris KS. 1986. Delphinid social organization and social behavior. Dolphin Cognition and Behavior: A Comparative Approach RJ Schusterman, JA Thomas, FG Wood 335–44 Hillsdale, NJ: Lawrence Erlbaum [Google Scholar]
  64. Johnson CM, Norris KS. 1994. Social behavior. The Hawaiian Spinner Dolphin KS Norris, B Wursig, RS Wells, M Wursig 243–86 Berkeley: Univ. Calif. Press [Google Scholar]
  65. Jones OR, Gaillard JM, Tuljapurkar S, Alho JS, Armitage KB. et al. 2008. Senescence rates are determined by ranking on the fast-slow life-history continuum. Ecol. Lett. 11:664–73 [Google Scholar]
  66. Jones OR, Scheuerlein A, Salguero-Gomez R, Camarda CG, Schaible R. et al. 2014. Diversity of ageing across the tree of life. Nature 505:169–73 [Google Scholar]
  67. Jorgenson JT, Festa-Bianchet M, Gaillard J-M, Wishart WD. 1997. Effects of age, sex, disease, and density on survival of bighorn sheep. Ecology 78:1019–32 [Google Scholar]
  68. Jullien M, Clobert J. 2000. The survival value of flocking in neotropical birds: Reality or fiction?. Ecology 81:3416–30 [Google Scholar]
  69. Keller L, Genoud M. 1997. Extraordinary lifespans in ants: a test of evolutionary theories of ageing. Nature 389:958–60 [Google Scholar]
  70. Kendall BE, Fox GA, Fujiwara M, Nogeire TM. 2011. Demographic heterogeneity, cohort selection, and population growth. Ecology 92:1985–93 [Google Scholar]
  71. Kim SY, Drummond H, Torres R, Velando A. 2011. Evolvability of an avian life history trait declines with father's age. J. Evol. Biol. 24:295–302 [Google Scholar]
  72. King AJ. 2010. Follow me! I'm a leader if you do; I'm a failed initiator if you don't?. Behav. Process. 84:671–74 [Google Scholar]
  73. Kirkwood TBL, Austad SN. 2000. Why do we age?. Nature 408:1006–8 [Google Scholar]
  74. Land KC, Yang Y, Yi Z. 2005. Mathematical demography. Handbook of Population D Poston, M Micklin 659–717 New York: Springer [Google Scholar]
  75. Lecomte VJ, Sorci G, Cornet S, Jaeger A, Faivre B. et al. 2010. Patterns of aging in the long-lived wandering albatross. Proc. Natl. Acad. Sci. USA 107:6370–75 [Google Scholar]
  76. Lee RD. 2003. Rethinking the evolutionary theory of aging: transfers, not births, shape senescence in social species. Proc. Nat. Acad. Sci. USA 100:9637–42 [Google Scholar]
  77. Lewontin RC. 1965. Selection for colonizing ability. The Genetics of Colonizing Species HG Baker, GL Stebbins 77–94 New York: Academic [Google Scholar]
  78. Longhurst A. 2002. Murphy's law revisited: longevity as a factor in recruitment to fish populations. Fish. Res. 56:125–31 [Google Scholar]
  79. Low M, Pärt T, Forslund P. 2007. Age-specific variation in reproduction is largely explained by the timing of territory establishment in the New Zealand stitchbird Notiomystis cincta. J. Anim. Ecol. 76:459–70 [Google Scholar]
  80. MacNulty DR, Smith DW, Vucetich JA, Mech LD, Stahler DR, Packer C. 2009. Predatory senescence in ageing wolves. Ecol. Lett. 12:1347–56 [Google Scholar]
  81. Martin JGA, Festa-Bianchet M. 2011. Age-independent and age-dependent decreases in reproduction of females. Ecol. Lett. 14:576–81 [Google Scholar]
  82. Massot M, Clobert J, Montes-Poloni L, Haussy C, Cubo J, Meylan S. 2011. An integrative study of ageing in a wild population of common lizards. Funct. Ecol. 25:848–58 [Google Scholar]
  83. McComb K, Moss C, Durant SM, Baker L, Sayialel S. 2001. Matriarchs as repositories of social knowledge in African elephants. Science 292:491–94 [Google Scholar]
  84. McNamara JM, Houston AI, Barta Z, Scheuerlein A, Fromhage L. 2009. Deterioration, death and the evolution of reproductive restraint in late life. Proc. R. Soc. B 276:4061–66 [Google Scholar]
  85. Medawar PB. 1952. An Unsolved Problem in Biology London: H.K. Lewis
  86. Monaghan P. 2010. Telomeres and life histories: the long and the short of it. Ann. N.Y. Acad. Sci. 1206:130–42 [Google Scholar]
  87. Monaghan P, Metcalfe NB, Torres R. 2009. Oxidative stress as a mediator of life history trade-offs: mechanisms, measurements and interpretation. Ecol. Lett. 12:75–92 [Google Scholar]
  88. Morales M, Onate M, Garcia MB, Munné-Bosch S. 2013. Photo-oxidative stress markers reveal absence of physiological deterioration with ageing in Borderea pyrenaica, an extraordinarily long-lived herb. J. Ecol. 101:555–65 [Google Scholar]
  89. Mota MDS, Abrahao AR, Oliveira HN. 2005. Genetic and environmental parameters for racing time at different distances in Brazilian thoroughbreds. J. Anim. Breed. Genet. 122:393–99 [Google Scholar]
  90. Munné-Bosch S, Lalueza P. 2007. Age-related changes in oxidative stress markers and abscisic acid levels in a drought-tolerant shrub, Cistus clusii grown under Mediterranean field conditions. Planta 225:1039–49 [Google Scholar]
  91. Novoseltsev VN, Michalski AI, Novoseltseva JA, Yashin AI, Carey JR, Ellis AM. 2012. An age-structured extension to the vectorial capacity model. PLOS ONE 7:e39479 [Google Scholar]
  92. Nussey DH, Coulson T, Delorme D, Clutton-Brock TH, Pemberton JM. et al. 2011. Patterns of body mass senescence and selective disappearance differ among three species of free-living ungulates. Ecology 92:1936–47 [Google Scholar]
  93. Nussey DH, Froy H, Lemaitre J-F, Gaillard J-M, Austad SN. 2013. Senescence in natural populations of animals: widespread evidence and its implications for bio-gerontology. Age Res. Rev. 12:214–25 [Google Scholar]
  94. Nussey DH, Pemberton JM, Pilkington JG, Blount JD. 2009. Life history correlates of oxidative damage in a free-living mammal population. Funct. Ecol. 23:809–17 [Google Scholar]
  95. Odling-Smee FJ, Laland KN, Feldman MW. 2003. Niche Construction: The Neglected Process in Evolution. Princeton, NJ: Princeton Univ. Press
  96. Olesiuk PF, Bigg MA, Ellis GM. 1990. Life history and population dynamics of resident killer whales (Orcinus orca) in the coastal waters of British Columbia and Washington State. Rep. Int. Whal. Comm. 12:Spec. Issue209–43 [Google Scholar]
  97. Packer C, Brink H, Kissui BM, Maliti H, Kushnir H, Caro T. 2011. Effects of trophy hunting on lion and leopard populations in Tanzania. Conserv. Biol. 25:142–53 [Google Scholar]
  98. Packer C, Kosmala M, Cooley HS, Brink H, Pintea L. et al. 2009. Sport hunting, predator control and conservation of large carnivores. PLOS ONE 4:e5941 [Google Scholar]
  99. Pardo D, Barbraud C, Authier M, Weimerskirch H. 2013a. Evidence for an age-dependent influence of environmental variations on a long-lived seabird's life-history traits. Ecology 94:208–20 [Google Scholar]
  100. Pardo D, Barbraud C, Weimerskirch H. 2013b. Females better face senescence in the wandering albatross. Oecologia 173:1283–94 [Google Scholar]
  101. Perez VI, Buffenstein R, Masamsetti V, Leonard S, Salmon AB. et al. 2009. Protein stability and resistance to oxidative stress are determinants of longevity in the longest-living rodent, the naked mole-rat. Proc. Natl. Acad. Sci. USA 106:3059–64 [Google Scholar]
  102. Pizzari T, Dean R, Pacey A, Moore H, Bonsall MB. 2008. The evolutionary ecology of pre- and post-meiotic sperm senescence. Trends Ecol. Evol. 23:131–40 [Google Scholar]
  103. Poole JH. 1994. Sex differences in the behavior of African elephants. The Differences Between the Sexes RV Short, E Balaban 331–46 Cambridge, UK: Cambridge Univ. Press [Google Scholar]
  104. Poole TB. 1987. Social behavior of a group of orangutans (Pongo pygmaeus) on an artificial island in Singapore Zoological Gardens. Zoo Biol. 6:315–30 [Google Scholar]
  105. Punzo F, Chavez S. 2003. Effect of aging on spatial learning and running speed in the shrew (Cryptotis parva). J. Mamm. 84:1112–20 [Google Scholar]
  106. Purves WK, Sadava D, Orians GH, Heller HC. 2004. Life: The Science of Biology New York: W.H. Freeman
  107. Radwan J. 2003. Male age, germline mutations and the benefits of polyandry. Ecol. Lett. 6:581–86 [Google Scholar]
  108. Reichert BE, Martin J, Kendall WL, Cattau CE, Kitchens WM. 2010. Interactive effects of senescence and natural disturbance on the annual survival probabilities of snail kites. Oikos 119:972–79 [Google Scholar]
  109. Reid JM, Bignal EM, Bignal S, McCracken DI, Bogdanova MI, Monaghan P. 2010. Parent age, lifespan and offspring survival: structured variation in life history in a wild population. J. Anim. Ecol. 79:851–62 [Google Scholar]
  110. Reznick DN, Bryant MJ, Roff D, Ghalambor CK, Ghalambor DE. 2004. Effect of extrinsic mortality on evolution of senescence in guppies. Nature 431:1095–99 [Google Scholar]
  111. Ricklefs RE. 2010. Life-history connections to rates of aging in terrestrial vertebrates. Proc. Natl. Acad. Sci. USA 107:10314–19 [Google Scholar]
  112. Roach DA. 2012. Age, growth and size interact with stress to determine life span and mortality. Exp. Gerontol. 47:782–86 [Google Scholar]
  113. Roach DA, Ridley CE, Dudycha JL. 2009. Longitudinal analysis of Plantago: Age-by-environment interactions reveal aging. Ecology 90:1427–33 [Google Scholar]
  114. Robinson MR, Mar KU, Lummaa V. 2012. Senescence and age-specific trade-offs between reproduction and survival in female Asian elephants. Ecol. Lett. 15:260–66 [Google Scholar]
  115. Robson SKA, Vickers M, Blows MW, Crozier RH. 2006. Age determination in individual wild-caught Drosophila serrata using pteridine concentration. J. Exp. Biol. 209:3155–63 [Google Scholar]
  116. Rowe CL. 2008. “The calamity of so long life”: life histories, contaminants, and potential emerging threats to long-lived vertebrates. BioScience 58:623–31 [Google Scholar]
  117. Rowley I. 1976. Cooperative breeding in Australian birds. Proc. XVI Int. Ornithol. Congr., Canberra, Aust., Aug. 12–17, 1974657–66 Canberra: Aust. Acad. Sci. [Google Scholar]
  118. Sacher GA. 1978. Longevity and aging in vertebrate evolution. BioScience 28:497–501 [Google Scholar]
  119. Saino N, Caprioli M, Romano M, Boncoraglio G, Rubolini D. et al. 2011. Antioxidant defenses predict long-term survival in a passerine bird. PLOS ONE 6:e19593 [Google Scholar]
  120. Salomons HM, Mulder GA, van de Zande L, Haussmann MF, Linskens MH, Verhulst S. 2009. Telomere shortening and survival in free-living corvids. Proc. R. Soc. B 276:3157–65 [Google Scholar]
  121. Schumacher P, Weyeneth A, Weber DC, Dorn S. 1997. Long flights in Cydia pomonella L. (Lepidopera: Tortricidae) measured by a flight mill: influence of sex, mated status and age. Phys. Entomol. 22:149–60 [Google Scholar]
  122. Selman C, Blount JD, Nussey DH, Speakman JR. 2012. Oxidative damage, ageing, and life-history evolution: where now?. Trends Ecol. Evol. 27:570–77 [Google Scholar]
  123. Sharp SP, Clutton-Brock TH. 2010. Reproductive senescence in a cooperatively breeding mammal. J. Anim. Ecol. 79:176–83 [Google Scholar]
  124. Shefferson RP, Roach DA, Hutchings M. 2013. Longitudinal analysis in Plantago: Strength of selection and reverse age analysis reveal age-indeterminate senescence. J. Ecol. 101:577–84 [Google Scholar]
  125. Sherratt TN, Hassall C, Laird RA, Thompson DJ, Cordero-Rivera A. 2011. A comparative analysis of senescence in adult damselflies and dragonflies (Odonata). J. Evol. Biol. 24:810–22 [Google Scholar]
  126. Siegel JS. 2012. The Demography and Epidemiology of Human Health and Aging New York: Springer
  127. Slobodkin LB. 1968. How to be a predator. Am. Zool. 8:43–51 [Google Scholar]
  128. Sparkman AM, Arnold SJ, Bronikowski AM. 2007. An empirical test of evolutionary theories for reproductive senescence and reproductive effort in the garter snake Thamnophis elegans. Proc. R. Soc. B 274:943–50 [Google Scholar]
  129. Stark JD, Banks JE. 2003. Population-level effects of pesticides and other toxicants on arthropods. Annu. Rev. Entomol. 48:505–19 [Google Scholar]
  130. Styer LM, Carey JR, Wang J-L, Scott TW. 2007. Mosquitoes do senesce: departure from the paradigm of constant mortality. Am. J. Trop. Med. Hyg. 76:111–17 [Google Scholar]
  131. Tafani M, Cohas A, Bonenfant C, Gaillard JM, Lardy S, Allaine D. 2013. Sex-specific senescence in body mass of a monogamous and monomorphic mammal: the case of Alpine marmots. Oecologia 172:427–36 [Google Scholar]
  132. Torres R, Drummond H, Velando A. 2011. Parental age and lifespan influence offspring recruitment: a long-term study in a seabird. PLOS ONE 6:e27245 [Google Scholar]
  133. Torres R, Velando A. 2007. Male reproductive senescence: the price of immune-induced oxidative damage on sexual attractiveness in the blue-footed booby. J. Anim. Ecol. 76:1161–68 [Google Scholar]
  134. Turbill C, Ruf T. 2010. Senescence is more important in the natural lives of long- than short-lived mammals. PLOS ONE 5:e12019 [Google Scholar]
  135. Tyack P. 1986. Population biology, social behavior, and communication in whales and dolphins. Trends Ecol. Evol. 1:144–50 [Google Scholar]
  136. Ungvari Z, Ridgway I, Philipp EE, Campbell CM, McQuary P. et al. 2011. Extreme longevity is associated with increased resistance to oxidative stress in Arctica islandica, the longest-living non-colonial animal. J. Gerontol. A 66:741–50 [Google Scholar]
  137. Vaupel JW, Baudisch A, Dolling M, Roach DA, Gampe J. 2004. The case for negative senescence. Theor. Popul. Biol. 65:339–51 [Google Scholar]
  138. Velando A, Noguera JC, Drummond H, Torres R. 2011. Senescent males carry premutagenic lesions in sperm. J. Evol. Biol. 24:693–97 [Google Scholar]
  139. Warner RR, Chesson PL. 1985. Coexistence mediated by recruitment fluctuations: a field guide to the storage effect. Am. Nat. 125:769–87 [Google Scholar]
  140. Wells RS. 1991. The role of long-term study in understanding the social structure of a bottlenose dolphin community. Dolphin Societies: Discoveries and Puzzles K Pryor, KS Norris 199–225 Berkeley: Univ. Calif. Press [Google Scholar]
  141. White J, Wagner RH, Helfenstein F, Hatch SA, Mulard H. et al. 2008. Multiple deleterious effects of experimentally aged sperm in a monogamous bird. Proc. Natl. Acad. Sci. USA 105:13947–52 [Google Scholar]
  142. Whitman K, Starfield AM, Quadling HS, Packer C. 2004. Sustainable trophy hunting of African lions. Nature 428:175–78 [Google Scholar]
  143. Williams GC. 1957. Pleiotropy, natural selection, and the evolution of senescence. Evolution 11:398–411 [Google Scholar]
  144. Williams PD, Day T. 2003. Antagonistic pleiotropy, mortality source interactions, and the evolutionary theory of senescence. Evolution 57:1478–88 [Google Scholar]
  145. Williams PD, Day T, Fletcher Q, Rowe L. 2006. The shaping of senescence in the wild. Trends Ecol. Evol. 21:458–63 [Google Scholar]
  146. Williams R, Lusseau D. 2006. A killer whale social network is vulnerable to targeted removals. Biol. Lett. 2:497–500 [Google Scholar]
  147. Zajitschek F, Brassil CE, Bonduriansky R, Brooks RC. 2009. Sex effects on life span and senescence in the wild when dates of birth and death are unknown. Ecology 90:1698–707 [Google Scholar]
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