1. Prey color preferences of the brook stickleback Culaea inconstans Prospectus Presentation Andrew Hodgson Advisor: Dr. A. Ross Black 21 March 2012

2. Outline Sticklebacks Breeding colors Prey color preferences Sensory exploitation Methods Results and conclusions

3. Introduction Fourspine Threespine http://www.elasmodiver.com/images/Tubesnout.jpg Yellow Tubesnout Encyclopedia of Life Blackspotted Encyclopedia of life Why red ornaments? New England Aquarium Fourspine Threespine

4. Introduction Brook Sticklebacks without any red breeding signals (Reisman and Cade 1964) Ninespine http://lsfv-nds.de/cms/index.php?page=neunstachliger-stichling Fifteenspine

5. Threespine Ninespine http://lsfv-nds.de/cms/index.php?page=neunstachliger-stichling Color preferences (Smith et al. 2004) Introduction m/f

6. Introduction ninespine threespine Origin of feeding bias for red

7. ninespine threespine Origin of feeding bias for red Introduction (sensory exploitation)

9. Methods

10. Paired comparisons ~10 trials Methods

11. Measurements First bites (~30) Total bites (~130) Repeated bites (~4) Latency to bite (65%, ~6 ds) Inspections (~45, ~8.5 ds) Independent Bead color Time Methods (measures)

12. Measurements First bites (1 st color = 2 nd color) Total bites (1 st color = 2 nd color) Repeated bites (1 st color = 2 nd color) Latency to bite (1 st color = 2 nd color) Inspections (1 st color = 2 nd color) Independent Bead color Time Methods (null hypotheses)

13. Methods (video analysis) First bites Total bites Repeated attempts Latency to bite Inspections

14. Binomial tests, e.g. p(red)=0.5 Holm-Bonferroni (Holm 1979) corrected Least squares regression Specialized Wilcoxon rank sum test (Capanu et al. 2006) Methods (stats)

15. First bites 20<n<31 Results (color preferences) * * * *

16. Total bites 71<n<198 p<0.001 * * * *

17. Results (repeated bites) *

18. Results (bites vs inspections) 155<n<236 P<0.001

19. Results (latency to bite) r 2 =0.10 p<0.001 F=36 n=333

20. Is brook stickleback red/orange spine coloration a secondary sexual character? Results

21. Results (red spines) 2011 breeding season n=311 *

22. Results (red spines) *

23. Local brook stickleback prefer red and orange prey to other colors A sensory exploitation origin of red stickleback breeding signals is supported Red pelvic spines are a secondary sexual character Discussion (conclusions)

24. Conclusions Why are sticklebacks sensitive to red? Hemoglobin? Carotenoids? Immunity Cancer prevention Cell growth Liver Water balance (Smith et al. 2004) Copyright Randall & Schietzelt & Harper College https://wiki.cgb.indiana.edu/display/DGC/Projects

25. Finish paired comparisons Continue to monitor red spine color Try patterns (e.g. red and blue vs red and yellow) Try movies Saturation Brightness Movement Future plans

26. Dr. Ross Black Dr. Camille McNeely Mike Rule and TNWR Robert Holliday Nicole Wallace Joseph Peters, Anna Wu, Tomoko Bennet Acknowledgments

27. Works Cited Capanu, M., G. A. Jones, and R. H. Randles. 2006. Testing for a preference using a sum of Wilcoxon signed rank statistics. Computational Statistics and Data Analysis 51:793-796. Holm, S. 1979. A simple sequentially rejective multiple test procedure. Scandanavian Journal of Statistics 6:65-70. Proctor, H. C. 1992. Sensory exploitation and the evolution of male mating behavior: a cladistic test using water mites (Acari: Parasitengona). Animal Behaviour 44:745-752. Reisman H. M., T. J. Cade. 1967. Physiological and behavioral aspects of reproduction in the brook stickleback, Culaea inconstans. American Midland Naturalist 77:257-295. Rodd, F. H., K. A. Hughes, G. F. Grether, and C. T. Baril. 2002. A possible non-sexual origin of mate preference: are male guppies mimicking fruit? Proceedings of the Royal Society: Biological Sciences 269: 475-81. Smith C., I. Barber, R. T. Wootton, and L. Chittka. 2004. A Receiver Bias in the origin of three- spined stickleback mate choice. Proceedings: Biological Sciences 271:949-955.

28. Questions?

30. McLennan 1993

31. Mattern and McLennan 2004

32. Kawahara et al 2009

33. Results (inspections before bites)

34. Introduction (hypotheses) Hypotheses: 1) More bites at red than any other color

35. Multiple predation attempts appear to predict color preferences Brook stickleback are more likely to bite at novel prey than inspect it and leave Conclusions

36. How to deal with learning statistically Objective measurements of red coloration Controlling for color Problems to solve

37. Introduction (phylogenetics) Phylogenetic relationships of the sticklebacks (Gasterosteidae), not including Spinachia Spinachia, modified from McLennan (1996) and Smith et al (2004)

40. Three main competing models Fisher runaway Good genes Sensory Bias Introduction (sexual selection) Trait and preference components

41. Introduction (sexual selection) Fisher runaway Trait origin: response to preference Preference origin: response to trait Good genes Trait origin: indirectly indicate fitness Preference origin: indirectly increase fitness

42. Introduction (sexual selection) Sensory bias Trait origin: response to preference Preference origin: another context

43. Fisher runaway Good genes Sensory bias * ModelPrediction Preference and trait coevolve Trait evolves first Preference evolves first e.g. in predation Process Positive feedback between preference and trait Females evolve to select traits indirectly linked to fitness Trait evolves in response to a preexisting, naturally selected preference Introduction (sensory bias)

44. Introduction Ninespine http://lsfv-nds.de/cms/index.php?page=neunstachliger-stichling Preference for red prey?

45. Introduction (sensory bias) Other Examples Water mite trembling (Proctor 1992) Tungara frog mating calls (Ryan and Rand 1990) Swordtail fish tails (Basolo 1990, 1995, 1996, 1998) Costa’s hummingbird songs (Clark and Feo 2009) Cichlid egg spots (Egger et al. 2011) Bird song repertoires (Collins 1999)