References
Araya-Ajoy, Y.G., Bolstad, G.H., Brommer, J., Careau, V., Dingemanse,
N.J. & Wright, J. (2018). Demographic measures of an individual’s
“pace of life”: fecundity rate, lifespan, generation time, or a
composite variable? Behav. Ecol. Sociobiol. , 72.
Araya-Ajoy, Y.G., Ranke, P.S., Kvalnes, T., Rønning, B., Holand, H.,
Myhre, A.M., et al. (2019). Characterizing morphological
(co)variation using structural equation models: Body size, allometric
relationships and evolvability in a house sparrow metapopulation.Evolution , 73, 452–466.
Baalsrud, H.T., Sæther, B.E., Hagen, I.J., Myhre, A.M., Ringsby, T.H.,
Pärn, H., et al. (2014). Effects of population characteristics
and structure on estimates of effective population size in a house
sparrow metapopulation. Mol. Ecol. , 23, 2653–2668.
Bienvenu, F. & Legendre, S. (2015). A new approach to the generation
time in matrix population models. Am. Nat. , 185, 834–843.
Bonduriansky, R., Maklakov, A., Zajitschek, F. & Brooks, R. (2008).
Sexual selection, sexual conflict and the evolution of ageing and life
span. Funct. Ecol. , 22, 443–453.
Boyce, M.S. (1984). Restitution of r- and K-Selection as a Model of
Density-Dependent Natural Selection. Annu. Rev. Ecol. Syst. , 15,
427–447.
Brown, J.H., Gillooly, J.F., Allen, A.P., Savage, V.M. & West, G.B.
(2004). Toward a metabolic theory of ecology. Ecology , 85,
1771–1789.
Caswell, H. (2001). Matrix Population Models: Construction,
Analysis, and Interpretation . Matrix Population Models: Construction,
Analysis, and Interpretation. Sinauer Associates.
Charlesworth, B. (1994). Evolution in Age-Structured Populations .
Cambridge Studies in Mathematical Biology. 2nd edn. Cambridge University
Press.
Chevin, L.M., Lande, R. & Mace, G.M. (2010). Adaptation, plasticity,
and extinction in a changing environment: Towards a predictive theory.PLoS Biol. , 8.
Cleasby, I. & Nakagawa, S. (2012). The influence of male age on
within‐pair and extra‐pair paternity in passerines. Ibis (Lond.
1859). , 154, 318–324.
Coale, A.J. (1972). Growth and Structure of Human Populations: A
Mathematical Investigation . Princeton University Press.
Cochran, M.E. & Ellner, S. (1992). Simple Methods for Calculating
Age-Based Life History Parameters for Stage-Structured Populations.Ecol. Monogr. , 62, 345–364.
Ellner, S.P. (2018). Generation time in structured populations.Am. Nat. , 192, 105–110.
Engen, S., Lande, R. & Sæther, B.-E. (2013). A Quantitative Genetic
Model of r- and K-Selection in a Fluctuating Population. Am.
Nat. , 181, 725–736.
Engen, S. & Sæther, B.E. (2016). Optimal age of maturity in fluctuating
environments under r- and K-selection. Oikos , 125, 1577–1585.
Gaillard, J.-M., Lemaître, J.-F., Berger, V., Bonenfant, C., Devillard,
S., Douhard, M., et al. (2016). Life Histories, Axes of
Variation in . Encycl. Evol. Biol.
Gaillard, J.-M., Yoccoz, N.G., Lebreton, J.-D., Bonenfant, C.,
Devillard, S., Loison, A., et al. (2005). Generation time: a
reliable metric to measure life-history variation among mammalian
populations. Am. Nat. , 166, 119-123; discussion 124-128.
Girndt, A., Chng, C.W.T., Burke, T. & Schroeder, J. (2018). Male age is
associated with extra-pair paternity, but not with extra-pair mating
behaviour. Sci. Rep. , 8, 8378.
Hadfield, J.D. (2010). MCMC Methods for Multi-Response Generalized
Linear Mixed Models: The MCMCglmm R Package. J. Stat. Softw. , 33,
1–36.
Hansen, T.F., Pélabon, C. & Houle, D. (2011). Heritability is not
Evolvability. Evol. Biol. , 38, 258–277.
Hsu, Y.-H., Simons, M.J.P., Schroeder, J., Girndt, A., Winney, I.S.,
Burke, T., et al. (2017). Age-dependent trajectories differ
between within-pair and extra-pair paternity success. J. Evol.
Biol. , 30, 951–959.
Jensen, H., Steinsland, I., Ringsby, T.H. & Sæther, B.E. (2008).
Evolutionary dynamics of a sexual ornament in the house sparrow (Passer
domesticus): The role of indirect selection within and between sexes.Evolution , 62, 1275–1293.
Kruuk, L.E.B. (2004). Estimating genetic parameters in natural
populations using the “animal model”. Philos. Trans. R. Soc.
Lond. B. Biol. Sci. , 359, 873–90.
Lande, R. (1982). A Quantitative Genetic Theory of Life History
Evolution. Ecology , 63, 607–615.
Lehtonen, J. & Lanfear, R. (2014). Generation time, life history and
the substitution rate of neutral mutations. Biol. Lett. , 10,
3–6.
Liker, A. & Székely, T. (2005). Mortality costs of sexual selection and
parental care in natural populations of birds. Evolution (N. Y). ,
59, 890–897.
MacArthur, R.H. & Wilson, E.O. (1967). The Theory of Island
Biogeography . Landmarks in Biology Series. Princeton University Press.
McGraw, J.B. & Caswell, H. (1996). Estimation of Individual Fitness
from Life-History Data. Am. Nat. , 147, 47–64.
Metcalfe, N.B. & Monaghan, P. (2003). Growth versus lifespan:
perspectives from evolutionary ecology. Exp. Gerontol. , 38,
935–940.
Niskanen, A.K., Billing, A.M., Holand, H., Hagen, I.J., Araya-Ajoy,
Y.G., Husby, A., et al. (2020). Consistent scaling of inbreeding
depression in space and time in house sparrows. Proc. Natl. Acad.
Sci. , in press.
van Noordwijk, A. & de Jong, G. (1986). Acquisition and allocation of
resources: their influence on variation in life history tactics.Am. Nat. , 128, 137–142.
Pärn, H., Ringsby, T.H., Jensen, H., Saether, B.-E., Parn, H., Ringsby,
T.H., et al. (2012). Spatial heterogeneity in the effects of
climate and density-dependence on dispersal in a house sparrow
metapopulation. Proc. R. Soc. B Biol. Sci. , 279, 144–152.
Pianka, E.R. (1970). On r- and K-Selection. Am. Nat. , 104,
592–597.
van de Pol, M. & Wright, J. (2009). A simple method for distinguishing
within- versus between-subject effects using mixed models. Anim.
Behav. , 77, 753–758.
R Core Team. (2015). R: A language and environment for statistical
computing.
Roff, D.A. (1993). Evolution Of Life Histories: Theory and
Analysis . The Evolution of Life Histories: Theory and Analysis.
Springer US.
Sæther, B.-E. & Engen, S. (2015). The concept of fitness in fluctuating
environments. Trends Ecol. Evol. , 30, 273–281.
Sæther, B.-E., Lande, R., Engen, S., Weimerskirch, H., Lillegard, M.,
Altwegg, R., et al. (2005). Generation time and temporal scaling
of bird population dynamics. Nature , 436, 99–102.
Stearns, S.C. (1976). Life-History Tactics: A Review of the Ideas.Q. Rev. Biol. , 51, 3–47.
Stearns, S.C. (1992). The Evolution of Life Histories . OUP
Oxford.
Steiner, U.K., Tuljapurkar, S. & Coulson, T. (2014). Generation time,
net reproductive rate, and growth in stage-age-structured populations.Am. Nat. , 183, 771–783.
Stubberud, M.W., Myhre, A.M., Holand, H., Kvalnes, T., Ringsby, T.H.,
Sæther, B.E., et al. (2017). Sensitivity analysis of effective
population size to demographic parameters in house sparrow populations.Mol. Ecol. , 26, 2449–2465.
Wright, J., Bolstad, G.H., Araya-Ajoy, Y.G. & Dingemanse, N.J. (2018).
Life-history evolution under fluctuating density-dependent selection and
the adaptive alignment of pace-of-life syndromes. Biol. Rev. , 7.
Wright, J., Bolstad, G.H., Araya-Ajoy, Y.G. & Dingemanse, N.J. (2019).
Life-history evolution under fluctuating density-dependent selection and
the adaptive alignment of pace-of-life syndromes. Biol. Rev. , 94,
230–247.
Table 1 . Univariate mixed-effect model results on the sources of
variation in life history traits measuring the timing of reproduction of
1052 individual house sparrows. Only individuals that produced at least
one recruit were included in the analyses. We present point estimates
and credible intervals in parenthesis.