Evolutionary Dynamics and Fitness in Wild Populations

Evolutionary dynamics and fitness in wild populations

Loeske Kruuk

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

“ No one supposes that all the individuals of the same species are cast in the same actual mould. These individual differences are of the highest importance for us, 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. for they are often inherited …and they thus afford materials for natural selection to act on. ”

evolutionary change: genetics and selection

Results from wild populations – genetic variance and selection – predicting evolutionary dynamics – fitness

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. “ Evolutionary biology has the most elaborate and beautiful mathematical theories in all of biology ”

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . ‘Price equation’ (G. Price, 1972) Change = response to selection + change due to other causes e.g.

‘Robertson equation’ (A. Robertson, 1966) response to selection = genetic association between trait and fitness

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited.‘Fundamental theorem of ‘Breeder’s equation’ (J. Lush, 1937) natural selection’ (R.A.Fisher, 1930) response to selection rate of adaptation = = heritability x selection additive genetic variance in fitness

+ phenotype data → genetic variance and selection

‘animal

modernfarmer.com

(Kruuk et al. 2000, 2004; Milner2018 et© II Joint al. Congress 2000; on Evolutionary R éBiologyale 2018.et All rights al. reserved 1999) - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

selection trait 2 fitness trait 1

trait 1

→ abundant genetic variance and directional selection

‘Robertson equation’ response to selection = genetic association between trait and fitness

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited.Fisher’s ‘fundamental theorem ‘Breeder’s equation’ of natural selection’ (1930) response to selection rate of adaptation = = heritability x selection genetic variance in fitness

‘Robertson equation’ response to selection = genetic association between trait and fitness

…doesn’t work in the wild:

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproductionMerilä even in part is prohibited., Sheldon & Kruuk, Genetica 2001 ‘Breeder’s equation’ response to selection = heritability x selection

‘Robertson equation’ response to selection = genetic association between trait and fitness

‘Robertson equation’ response to selection = genetic covariance between trait and fitness

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited.Why doesn’t it work? Test this! ‘Breeder’s equation’ Morrissey, Kruuk & Wilson, response to selection J. Evolutionary Biology 2010 = heritability x selection

Yes No something else

trait fitness trait fitness

Evolutionary Yes No response?

G. Dabb

Andrew Cockburn

30 years (1988 –), 9683 individuals, 2401 breeding events, 15 generations

G. Dabb

John Gould 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Average Time of year Nestling’s age mass = 7g

Number of helpers Mother’s age

Brood size heritability = 19% (13-25%, 95% CI) Weather

(Kruuk et al. Global Change Biology 2015;2018 © II Joint Hajduk Congress on Evolutionaryet al.Biology Evolution 2018. All rights reserved - 2018)Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Females Males

0,4 0,35 0,3 0,25 0,2 0,15 0,1 0,05

Early SurvivalEarly 0 [4,6] (6,6.7] (6.7,7.1] (7.1,7.6] (7.6,8.1] (8.1,10.7] Gabi Hajduk Mass (g) 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. Heavier nestlings have higher survival: CovP = 0.06 (0.04 - 0.09 95% CI)

→ genetic variance and directional selection

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . There is also genetic variance for early survival BUT no genetic covariance: -0.02 (-0.09 - 0.05, 95% CI)

Survival~size associations are driven by timing :

Gabi Hajduk

Time in season (fortnight) Time in season (fortnight)

. 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

No No trend in mean size Something else

trait fitness Mean mass (g)

heritability 23% (18 – 28%, 95% CI)

G. Dabb

The early bird gets the girls:

date of moult (log scale) 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

extra-pair within-pair

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. No genetic covariance of WPS There is genetic with plumage: covariance of EPS +0.03 (-0.03, 0.04) with plumage: +0.11 (0.05-0.17) 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

extra-pair breeding success total adult within-pair breeding breeding success success

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. Genetic covariance of longevity total breeding success and plumage: +0.01 (-0.05 - 0.06) 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

genetic variance for extra-pair success, and genetic covariance with plumage phenology

but extra-pair success isn’t everything!

no genetic covariance with total breeding success

…and no change over time in plumage phenology: Mean (SE), days Mean (SE),

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproductionYear even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Results from wild populations – genetic variance and selection – predicting evolutionary dynamics – fitness

‘Robertson equation’, secondary theorem (A. Robertson, 1966) response to selection = genetic association between trait and fitness

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. ‘Fundamental theorem of ‘Breeder’s equation’ (J. Lush, 1937) natural selection’ (R.A.Fisher, 1930) response to selection rate of adaptation = = heritability x selection additive genetic variance in fitness

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. Change in mean fitness = additive genetic variance in fitness But this is very difficult to estimate ! (Hendry et al. AREES 2018 in press) 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Canada UK Netherlands Switzerland

M. Festa-Bianchet A. Ozgul J. Visser T. Bonnet France Australia

S. Morris

H. Kruuk G. Dabb Ten data-sets, ~350 years’ pedigree data S. Losdat

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. J. Visser 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . Marco Festa-Bianchet Jane Reid Dave Coltman Lukas Keller Fanie Pelletier Matthew Wolak Peter Arcese Josephine Pemberton Michael Morrissey Anne Charmantier Timothée Dan Nussey Céline Teplitsky Bonnet Chris de Franceschi Josephine Pemberton Loeske Kruuk Ben Sheldon Tim Clutton-Brock Simon Evans

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reservedErik - Any reproductionPostma even in part is prohibited. Timothée Bonnet Marcel Visser Michael Glauco Camenisch Morrissey Andrew McAdam Andrew Cockburn Stan Boutin Loeske Kruuk

How many babies did an individual have across its entire lifetime? - for every individual

Soay sheep Blue tit

Red Great tit deer (UK)

Red Superb squirrel fairy-wren

Lifetime Breeding Success Lifetime Breeding Success 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Distributions like this are challenging : combine additional 0’s with (overdispersed) Poisson process → “zero-inflated Poisson”

Model: additive genetic variance, maternal effects and cohort effects

Bighorn sheep Song sparrow Heritability Soay sheep Maternal Blue tit Cohort

Red deer Great tit (UK)

Superb Red squirrel fairy-wren

Proportion of variance (on data scale, with 95% CI) 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Heritability of fitness: 0.01 - 7.5%, median 2.5% Additive genetic variance for relative fitness: 0.004 – 0.61, median 0.056

2018 © II Joint CongressMore on Evolutionary Biology details 2018. All rights reserved in Tim- Any reproduction Bonnet’s even in part is prohibited. poster! How much are wild vertebrate populations evolving right now? Symposium 64, poster P-0260, this evening

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . Evolutionary dynamics will The theory of evolutionary dynamics be shaped by quantitative genetic parameters Results from wild populations – genetic variance and selection Heritability and selection – predicting evolutionary dynamics alone may not generate – fitness an evolutionary response

1. More taxa!

2. Wild genomics

IPCC

Andrew Cockburn Adam Hayward Ben Sheldon Arild Husby Helen Osmond Jarrod Hadfield Juha Merilä Matt Robinson Gabi Hajduk Alison Morris Anne Charmantier Jon Slate Timothée Bonnet Sean Morris Dany Garant Mick Crawley Michael Morrissey Jill Pilkington Alastair Wilson Steve Albon Josephine Pemberton Michael Jennions Dan Nussey Sue Brotherstone Tim Clutton-Brock Megan Head Tim Coulson Jeff Lane Nick Barton Regina Vega-Trejo Kathi Foerster Patrick Meir Bill Hill Janet Gardner Michelle Clements My family and Craig Walling Marco Festa-Bianchet Hannah Froy other animals Katie Stopher2018 © II Joint Congress on Evolutionary BiologyWendy 2018. All rights King reserved - Any reproduction even in part is prohibited.Ian Stevenson

© Ashley Latimer 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .