Date of Degree

2-2014

Document Type

Dissertation

Degree Name

Ph.D.

Program

Biology

Advisor(s)

Mark E. Hauber

Subject Categories

Behavior and Ethology | Biology

Abstract

Hosts of brood parasitic birds face fitness costs associated with rearing unrelated offspring. In response, the recognition and rejection of parasitic eggs is a common host defense. Brown-headed cowbirds (Molothrus ater) challenge coevolutionary theory, because although they exploit over 200 host species, they lay non-mimetic eggs, and most hosts do not combat cowbird parasitism with egg rejection. American robins (Turdus migratorius) are one of a handful of cowbird hosts known to recognize and remove cowbird eggs from the nest. I addressed the mechanistic and evolutionary drivers of egg rejection in this host species, by disentangling the roles of spectral tuning and visual physiology on the behavioral outcome of egg rejection, by estimating the costs of parasitism which may drive egg rejection behavior, and by addressing the reciprocal effects parasitism on host egg color variation and its role in mediating rejection decisions. I also test assumptions underlying the evolvability of host egg rejection responses in this system. In Chapter 1, I lay out an overview of brood parasitism as a reproductive strategy and brood parasite-host ecology, and highlight evolutionary mechanisms and consequences of coevolution in these systems. In Chapter 2, I test the hypothesis that foreign egg rejection is driven proximately by perceivable differences in ground color between host and parasitic eggs across the entire avian spectral sensitivity range. I show that the rejection of artificially dyed eggs is mediated by input from all four avian single-cone photoreceptors, and that more divergent model `parasitic' eggs are indeed rejected at higher rates. However, the cowbird egg does not conform to this prediction, because both model and real cowbird eggs are rejected in 100% of experimental trials despite their lower overall discriminability from robin eggs. This may indicate a cowbird-egg specific rejection response in robins. In Chapter 3, I test a critical assumption underlying the evolution of cowbird-specific egg rejection responses in robins, by assessing the hypothesis that cowbird parasitism imposes recoverable costs on robin hosts. My results indicate that cowbird chicks fare poorly when reared alongside robin chicks, but parasitism per se still reduces nesting success for robins; thus, rejection of cowbird eggs serves a function to eliminate the cost of parasitism. In Chapter 4, I examine a critical assumption underlying all of host-parasite coevolutionary theory, namely that host defenses can evolve genetically in response to parasitism. I address the hypothesis that egg rejection is repeatable in our study population, as repeatability is prerequisite to the evolution and spread of a behavioral trait, including a predictor of the trait's genetic heritability. As predicted, egg rejection behavior in American robins was found to be highly repeatable for intermediately-rejected model egg colors within the same nesting attempt, irrespective of potentially confounding ecological and temporal factors. Finally, in Chapter 5, I test predictions stemming from alternate hypotheses that egg rejection evolved in response to cowbird (non-mimetic) versus conspecific (mimetic) parasitism, by investigating the degree of color variation within robins' own clutches, and the effect of experimentally manipulating intraclutch color variation. I used both observational and experimental data, and found that egg color varies more between clutches than among egg within a single clutch, yet experimental manipulated intraclutch color variation did not affect rejection rates. These results support the scenario of historical parasitism by non-mimetic parasites. Variation among the findings of similar studies pertaining to hosts of mimetic parasites may be explained by hosts' use of different cognitive mechanisms in the decision to reject foreign eggs, However, for hosts of non-mimetic parasites, investigating egg color variation and its effect on egg rejection is not informative about different cognitive decision-making rules, as predictions under each mechanism are similar - that there will be no effect of a history of parasitism on intraclutch color variation (observational patterns) or rejection rate (experimental data). This body of research presents compelling evidence in support of egg rejection by robins as a specific response to historical cowbird parasitism, and has highlighted important components of the sensory, cognitive, functional and evolutionary processes underlying egg rejection in this paradoxical brood parasite-host system.

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