1. Early nest environment and genes have lifetime effects on the expression of a colorful sexual signal Joey K. Hubbard, Brittany R. Jenkins, Rebecca J. Safran Joanna.Hubbard@colorado.edu Animal Behavior Society July 30, 2013

2. Phenotypic Variation

3. Genes Selective pressure Phenotypic change over generations Environment Environmental change Phenotypic change over a short time period

4. Sexually-Selected Traits Phenotypic Variation Environmental Control Genetic Control Indirect Benefits Direct Benefits

5. Colorful Plumage Traits

6. Melanin-Based Color

7. North American Barn Swallows Sexual trait Socially Monogamous Extra-pair paternity Monitoring Boulder County breeding population since 2008 Safran & McGraw 2004; Safran et al. 2005

8. Determined paternity using microsats Pedigree 530 offspring with 111 mothers and 98 fathers

10. Animal Model Univariate animal model to partition sources of variance for each color descriptor Determine key variance components Calculate variance ratios V P = V A + V T + V CE + V ME + V R

11. Variance Ratios h2h2 V A /V P : Narrow sense heritability t2t2 V T /V P : Effect of seasonal variation ce 2 V CE /V P : Effect of common environment me 2 V ME /V P : Effect of maternal environment V P = V A + V T + V CE + V ME + V R

12. Influences on Juvenile Color Variance RatiosAvg. Brightness h 2 heritability 0.10 (0.23) t 2 seasonal effect 0.05 (0.04) ce 2 common environment 0.11 (0.08) me 2 maternal environment 0.05 (0.09) Hubbard et al. in prep (Standard Error)

13. Influences on Juvenile Color Variance RatiosAvg. BrightnessHue h 2 heritability 0.10 (0.23) 1.1 -07 (7.1e -09 ) t 2 seasonal effect 0.05 (0.04)0.03 (0.04) ce 2 common environment 0.11 (0.08)0.09 (0.08) me 2 maternal environment 0.05 (0.09) 0.12 (0.07) Hubbard et al. in prep (Standard Error)

14. Influences on Juvenile Color Variance RatiosAvg. BrightnessHueChroma h 2 heritability 0.10 (0.23) 1.1 -07 (7.1e -09 ) 0.19 (0.15) t 2 seasonal effect 0.05 (0.04)0.03 (0.04)0.05 (0.04) ce 2 common environment 0.11 (0.08)0.09 (0.08)0.13 (0.08) me 2 maternal environment 0.05 (0.09) 0.12 (0.07) 4.3 -07 (7.5e -08 ) Hubbard et al. in prep (Standard Error)

15. Within-Individual Variation Hatch YearSecond Year Year 1 Year 2 54 individuals (out of ~4000 nestlings) 205 individuals

16. Within-Individual Variation Hatch YearSecond Year

17. Within-Individual Variation bR2R2 p Brightnes s 0.480.22< 0.001 Hue0.1000.49 Chroma0.870.29< 0.001 Hatch Year to Second Year

18. Within-Individual Variation bR2R2 p Brightnes s 0.480.22< 0.001 Hue0.1000.49 Chroma0.870.29< 0.001 bR2R2 p Brightnes s 0.410.21< 0.001 Hue0.210.040.003 Chroma0.530.30< 0.001 Year 1 to Year 2 Hatch Year to Second Year

19. Summary Genetic Variation Environmental Variation Maternal Environment ✔✔

20. For the Swallows Early environment has long-term effects on signal development Females gain both indirect and direct benefits

21. Heritability Estimates Ignoring other sources of non-independence results in overestimation of heritability AM with environment AM no environment P-O Regression Sibling Correlation h 2 = 0.10h 2 = 0.45h 2 = 0.36h 2 = 0.84 Average Brightness

22. Cross-fostering experiments Experimentally tease apart effects of shared genes and shared environment Explore specific environmental influences

23. Thanks! Funding: American Ornithologists Union Animal Behavior Society CU EBIO Department CU Graduate School NSF CU BURST and UROP programs Lab and Field Help: Amazing Field Crews!Site Owners Stephen AlderferMonica Brandhuber Andrew FlynnAmanda Hund Eric LordJulie Marling Liz ScordatoAudrey Tobin Ty TuffCourtney van der Linden Rachel WildrickMatt Wilkins Photo Credit: Matt Wilkins Ian Harold

24. Quantitative Genetics Variance Components Avg. BrightnessHueChroma Trait Mean 29.98% (0.015)618.94 nm (0.04)0.41 (5.04e -5 ) VAVA 5.91 (12.82) p = 0.696 5.23e -05 (3.81e -06 ) p = 0.999 1.16e -04 (9.44e -05 ) p = 0.326 VTVT 2.97 (2.55) p = 0 16.87 (20.15) p = 0.362 3.11e -05 (2.63e -05 ) p = 0.167 V CE 6.46 (4.57) p = 0.125 41.56 (38.22) p = 0.263 7.63e -05 (4.80e -05 ) p = 0.090 V ME 2.57 (5.39) p = 0.664 60.92 (35.40) p = 0.088 2.63e -10 (4.43e -11 ) p = 0.999 VRVR 38.92 (8.14)369.60 (26.93)3.86e -04 (6.50e -05 ) VPVP 56.84 (0.303)488.95 (0.088)6.09e -04 (1.06e -04 )

25. Adult Color Proxy Hubbard et al. in prep