Renee A Duckworth

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• Duckworth, R. A., Belloni, V., & Anderson, S. R. (2015). Cycles of species replacement emerge from locally induced maternal effects on offspring behavior in a passerine bird. Science, 347, 875-877. doi:10.1126/science.1260154
• Duckworth, R. A. (2014). Evolution of genetically integrated dispersal strategies. DISPERSAL ECOLOGY AND EVOLUTION, 83-94.
• Duckworth, R. A. (2013). Epigenetic inheritance systems act as a bridge between ecological and evolutionary timescales. Behavioral Ecology, 24(2), 327-328.
• Duckworth, R. A., & Kruuk, L. E. (2013). EVOLUTION OF GENETIC INTEGRATION BETWEEN DISPERSAL AND COLONIZATION ABILITY IN A BIRD. EVOLUTION, 63(4), 968-977.
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  Discrete behavioral strategies comprise a suite of traits closely integrated in their expression with consistent natural selection for such coexpression leading to developmental and genetic integration of their components. However, behavioral traits are often also selected to respond rapidly to changing environments, which should both favor their context-dependent expression and inhibit evolution of genetic integration with other, less flexible traits. Here we use a multigeneration pedigree and long-term data on lifetime fitness to test whether behaviors comprising distinct dispersal strategies of western bluebirds-a species in which the propensity to disperse is functionally integrated with aggressive behavior-are genetically correlated. We further investigated whether selection favors flexibility in the expression of aggression in relation to current social context. We found a significant genetic correlation between aggression and dispersal that is concordant with consistent selection for coexpression of these behaviors. To a limited extent, individuals modified their aggression to match their mate; however, we found no fitness consequences on such adjustments. These results introduce a novel way of viewing behavioral strategies, where flexibility of behavior, while often aiding an organism's fit in its current environment, may be limited and thereby enable integration with less flexible traits.
• Duckworth, R. A., & Sockman, K. W. (2012). Proximate mechanisms of behavioural inflexibility: Implications for the evolution of personality traits. Functional Ecology, 26(3), 559-566.
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  Abstract: Behaviour is often assumed to be the most flexible of traits, yet recent studies show a high repeatability of behaviour within individuals even across different functional contexts. Such consistent expression of behaviour may evolve either when selection favours its integration with less flexible components of the phenotype or when pleiotropic effects produce correlations between behaviours that have different optimal timing of expression. Examining the physiological mechanisms underlying correlated expression of behaviours provides powerful insight into the evolution of personalities by establishing the extent to which pleiotropic effects might limit the independent evolution of distinct behaviours. 2.Here, we investigated proximate mechanisms behind aggressive and non-aggressive personality types in western bluebirds, Sialia mexicana, to determine whether consistency in the expression of aggression is because of shared effects of plasma-circulating androgens on aggression and mating behaviour. 3.We found that androgen concentration was unrelated to variation in both intra- and interspecific aggression even though it was closely linked to variation in male mating behaviour. These results suggest that pleiotropic effects of circulating androgens are unlikely to cause consistent differences among individuals in aggression. 4.These findings suggest that decoupling of the activational effects of hormones on behaviour is an important step in the evolution of personality traits. © 2012 The Authors. Functional Ecology © 2012 British Ecological Society.
• Duckworth, R. A., Gil, D., & Brumm, H. (2012). Human-induced changes in the dynamics of species coexistence : an example with two sister species. AVIAN URBAN ECOLOGY: BEHAVIOURAL AND PHYSIOLOGICAL ADAPTATIONS, 181-191.
• Duckworth, R. A. (2010). Aggressive behaviour affects selection on morphology by influencing settlement patterns in a passerine bird. PROCEEDINGS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES, 273(1595), 1789-1795.
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  The importance of behaviours as instigators or inhibitors of evolutionary change remains largely unresolved and this is in part because there are very few empirical examples of how behaviours affect evolutionary processes. By determining the environment of breeding, aggressive interactions over territories have the potential to strongly impact selection pressures experienced by individuals. Western bluebirds (Sialia mexicana) provide a unique opportunity to investigate the evolutionary importance of aggression, since their highly variable breeding habitat favours distinct foraging techniques and they also compete aggressively for nest boxes, a resource that is easy to manipulate. Here, I show experimentally that more aggressive males compete more effectively for territories with a high density of nest boxes and, as a consequence, aggressive and non-aggressive males are sorted into distinct breeding habitats that differ in the strength of selection on morphological traits. Specifically, males with longer tails and tarsi were favoured in open habitats where high agility is required to forage efficiently, whereas in forested habitats, where agility is less important, selection was weak. These results show that aggression can affect selection on a local scale by determining individual settlement patterns. More generally, because territorial interactions are important across a wide variety of taxa, these results suggest that aggressive behaviour has the potential to impact the evolutionary trajectory of many animal populations.
• Duckworth, R. A. (2009). Maternal effects and range expansion: A key factor in a dynamic process?. Philosophical Transactions of the Royal Society B: Biological Sciences, 364(1520), 1075-1086.
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  PMID: 19324612;PMCID: PMC2666686;Abstract: Species that depend on ephemeral habitat often evolve distinct dispersal strategies in which the propensity to disperse is closely integrated with a suite of morphological, behavioural and physiological traits that influence colonizing ability. These strategies are maintained by natural selection resulting from spatial and temporal variation in resource abundance and are particularly evident during range expansion. Yet the mechanisms that maintain close alignment of such strategies with resource availability, integrate suites of dispersal traits and generate variability in dispersal propensity are rarely known. Breeding females can influence offspring phenotype in response to changes in current environmental conditions, making maternal effects uniquely suited to bridge fluctuations in resource abundance in the maternal generation and variation in offspring dispersal ability. Western bluebirds' (Sialia mexicana) dependence on nest cavities - an ephemeral resource - has led to the evolution of two distinct dispersal phenotypes: aggressive males that disperse and non-aggressive males that remain philopatric and cooperate with their relatives. Over the last 40 years, western bluebirds rapidly expanded their geographical range, providing us with an opportunity to test, in newly established populations, the importance of maternal effects for generating variability in dispersal propensity. Here, I show that, under variable resource conditions, breeding females group offspring of different competitive ability in different positions in the egg-laying order and, consequently, produce aggressive males that are more likely to disperse when resources are low and non-aggressive philopatric males when resources are abundant. I then show experimentally that the association between resource availability and sex-specific birth order is robust across populations. Thus, this maternal effect enables close tracking of resource availability and may explain how variation in dispersal is generated in newly colonized populations. More generally, these results suggest that, as a key source of variation in colonizing phenotypes, maternal effects are of crucial importance for understanding the dynamics of range expansion. © 2009 The Royal Society.
• Duckworth, R. A. (2009). The role of behavior in evolution: A search for mechanism. Evolutionary Ecology, 23(4), 513-531.
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  Abstract: Behavior has been viewed as a pacemaker of evolutionary change because changes in behavior are thought to expose organisms to novel selection pressures and result in rapid evolution of morphological, life history and physiological traits. However, the idea that behavior primarily drives evolutionary change has been challenged by an alternative view of behavior as an inhibitor of evolution. According to this view, a high level of behavioral plasticity shields organisms from strong directional selection by allowing individuals to exploit new resources or move to a less stressful environment. Here, I suggest that absence of clear mechanisms underlying these hypotheses impedes empirical evaluation of behavior's role in evolution in two ways. First, both hypotheses focus on behavioral shifts as a key step in the evolutionary process but ignore the developmental mechanisms underlying these shifts and this has fostered unwarranted assumptions about the specific types of behavioral shifts that are important for evolutionary change. Second, neither hypothesis provides a means of connecting within-individual changes in behavior to population-level processes that lead to evolutionary diversification or stasis. To resolve these issues, I incorporate developmental and evolutionary mechanisms into a conceptual framework that generates predictions about the types of behavior and types of behavioral shifts that should affect both micro and macroevolutionary processes. © Springer Science+Business Media B.V. 2008.
• Duckworth, R. A., & E., L. (2009). Evolution of genetic integration between dispersal and colonization ability in a bird. Evolution, 63(4), 968-977.
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  PMID: 19154391;Abstract: Discrete behavioral strategies comprise a suite of traits closely integrated in their expression with consistent natural selection for such coexpression leading to developmental and genetic integration of their components. However, behavioral traits are often also selected to respond rapidly to changing environments, which should both favor their context-dependent expression and inhibit evolution of genetic integration with other, less flexible traits. Here we use a multigeneration pedigree and long-term data on lifetime fitness to test whether behaviors comprising distinct dispersal strategies of western bluebirds - a species in which the propensity to disperse is functionally integrated with aggressive behavior - are genetically correlated. We further investigated whether selection favors flexibility in the expression of aggression in relation to current social context. We found a significant genetic correlation between aggression and dispersal that is concordant with consistent selection for coexpression of these behaviors. To a limited extent, individuals modified their aggression to match their mate; however, we found no fitness consequences on such adjustments. These results introduce a novel way of viewing behavioral strategies, where flexibility of behavior, while often aiding an organism's fit in its current environment, may be limited and thereby enable integration with less flexible traits. © 2009 The Author(s).
• Duckworth, R., & Duckworth, R. A. (2009). Maternal effects and range expansion: a key factor in a dynamic process?. Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 364(1520).
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  Species that depend on ephemeral habitat often evolve distinct dispersal strategies in which the propensity to disperse is closely integrated with a suite of morphological, behavioural and physiological traits that influence colonizing ability. These strategies are maintained by natural selection resulting from spatial and temporal variation in resource abundance and are particularly evident during range expansion. Yet the mechanisms that maintain close alignment of such strategies with resource availability, integrate suites of dispersal traits and generate variability in dispersal propensity are rarely known. Breeding females can influence offspring phenotype in response to changes in current environmental conditions, making maternal effects uniquely suited to bridge fluctuations in resource abundance in the maternal generation and variation in offspring dispersal ability. Western bluebirds' (Sialia mexicana) dependence on nest cavities--an ephemeral resource--has led to the evolution of two distinct dispersal phenotypes: aggressive males that disperse and non-aggressive males that remain philopatric and cooperate with their relatives. Over the last 40 years, western bluebirds rapidly expanded their geographical range, providing us with an opportunity to test, in newly established populations, the importance of maternal effects for generating variability in dispersal propensity. Here, I show that, under variable resource conditions, breeding females group offspring of different competitive ability in different positions in the egg-laying order and, consequently, produce aggressive males that are more likely to disperse when resources are low and non-aggressive philopatric males when resources are abundant. I then show experimentally that the association between resource availability and sex-specific birth order is robust across populations. Thus, this maternal effect enables close tracking of resource availability and may explain how variation in dispersal is generated in newly colonized populations. More generally, these results suggest that, as a key source of variation in colonizing phenotypes, maternal effects are of crucial importance for understanding the dynamics of range expansion.
• Duckworth, R., Mendonca, M., & Hill, G. (2009). A condition dependent link between testosterone and disease resistance in the house finch. PROCEEDINGS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES, 268(1484), 2467-2472.
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  Testosterone has recently been proposed as a link between male quality and health and the expression of sexual traits. We investigated the relationship between testosterone and measures of the individual condition and health of males in a natural population of house finches (Carpodacus mexicanus). We also conducted a captive experiment in order to test for the effects of testosterone on resistance to coccidia, which is a common parasite of house finches. Free-living finales in better condition had higher testosterone levels and loiter corticosterone levels than free-living males iii poor condition. In our captive experiment, increased testosterone accelerated the rate of coccidial infection as compared with sham-implanted or gonadectomized males. Although the differences were not significant, free-living males infected with coccidia had louver levels of testosterone and higher levels of corticosterone than males that were not infected. Thus, experimentally elevating testosterone levels in captive males resulted in a higher percentage of infected males, while free-living males with coccidial infection had low testosterone levels. This apparent discrepancy between captive and free-living males iii the association of testosterone and disease may be explained by the condition dependence of testosterone. These results suggest that the testosterone-dependent sexual traits reliably indicate male overall condition and health and, thus, females could benefit front assessing potential mates based on these traits.
• Badyaev, A., & Duckworth, R. (2008). Context-dependent sexual advertisement: plasticity in development of sexual ornamentation throughout the lifetime of a passerine bird. JOURNAL OF EVOLUTIONARY BIOLOGY, 16(6), 1065-1076.
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  Male investment into sexual ornamentation is a reproductive decision that depends on the context of breeding and life history state. In turn, selection for state- and context-specific expression of sexual ornamentation should favour the evolution of developmental pathways that enable the flexible allocation of resources into sexual ornamentation. We studied lifelong variation in the expression and condition-dependence of a sexual ornament in relation to age and the context of breeding in male house finches (Carpodacus mexicanus) - a species that develops a new sexual ornament once a year after breeding. Throughout males' lifetime, the elaboration of ornamentation and the allocation of resources to the development of sexual ornamentation depended strongly on pairing status in the preceding breeding season - males that were single invested more resources into sexual ornamentation and changed ornamentation more than males that were paired. During the initial (post-juvenile) moult, the expression of ornamentation was closely dependent on individual condition, however the condition-dependence of ornamentation sharply decreased throughout a male's lifetime and in older males expression of sexual ornamentation was largely independent of condition during moult. Selection for early breeding favoured greater ornamentation in males that were single in the preceding seasons and the strength of this selection increased with age. On the contrary, the strength of selection on sexual ornamentation decreased with age in males that were paired in the preceding breeding season. Our results reveal strong context-dependency in investment into sexual ornamentation as well as a high flexibility in the development of sexual ornamentation throughout a male's life.
• Duckworth, R. A. (2008). Adaptive dispersal strategies and the dynamics of a range expansion. American Naturalist, 172(SUPPL. 1), S4-S17.
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  PMID: 18554143;Abstract: In species undergoing range expansion, newly established populations are often more dispersive than older populations. Because dispersal phenotypes are complex and often costly, it is unclear how highly dispersive phenotypes are maintained in a species to enable their rapid expression during periods of range expansion. Here I test the idea that metapopulation dynamics of local extinction and recolonization maintain distinct dispersal strategies outside the context of range expansion. Western bluebirds display distinct dispersal phenotypes where aggressive males are more dispersive than non-aggressive males, resulting in highly aggressive populations at the edge of their expanding range. I experimentally created new habitat interior to the range edge to show that, as on the range front, it was colonized solely by aggressive males. Moreover, fitness consequences of aggression depended on population age: aggressive males had high fitness when colonizing new populations, while nonaggressive males performed best in an older population. These results suggest that distinct dispersal strategies were maintained before range expansion as an adaptation for the continual recolonization of new habitat. These results emphasize similarities between range expansion and metapopulation dynamics and suggest that preexisting adaptive dispersal strategies may explain rapid changes in dispersal phenotypes during range expansion. © 2008 by The University of Chicago.
• Duckworth, R. A., & Badyaev, A. V. (2008). Coupling of dispersal and aggression facilitates the rapid range expansion of a passerine bird. PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA, 104(38), 15017-15022.
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  Behaviors can facilitate colonization of a novel environment, but the mechanisms underlying this process are poorly understood. On one hand, behavioral flexibility allows for an immediate response of colonizers to novel environments, which is critical to population establishment and persistence. On the other hand, integrated sets of behaviors that display limited flexibility can enhance invasion success by coupling behaviors with dispersal strategies that are especially important during natural range expansions. Direct observations of colonization events are required to determine the mechanisms underlying changes in behavior associated with colonization, but such observations are rare. Here, we studied changes in aggression on a large temporal and spatial scale across populations of two sister taxa of bluebirds (Sialia) to show that coupling of aggression and dispersal strongly facilitated the range expansion of western bluebirds across the northwestern United States over the last 30 years. We show that biased dispersal of highly aggressive males to the invasion front allowed western bluebirds to displace less aggressive mountain bluebirds. However, once mountain bluebirds were excluded, aggression of western bluebirds decreased rapidly across consecutive generations in concordance with local selection on highly heritable aggressive behavior. Further, the observed adaptive microevolution of aggression was accelerated by the link between dispersal propensity and aggression. Importantly, our results show that behavioral changes among populations were not caused by behavioral flexibility and instead strongly implicate adaptive integration of dispersal and aggression in facilitating the ongoing and rapid reciprocal range change of these species in North America.
• Duckworth, R., Badyaev, A. V., Young, R. L., Hill, G. E., & Duckworth, R. A. (2008). Evolution of sex-biased maternal effects in birds. IV. Intra-ovarian growth dynamics can link sex determination and sex-specific acquisition of resources. Journal of evolutionary biology, 21(2).
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  The evolutionary importance of maternal effects is determined by the interplay of maternal adaptations and strategies, offspring susceptibility to these strategies, and the similarity of selection pressures between the two generations. Interaction among these components, especially in species where males and females differ in the costs and requirements of growth, limits inference about the evolution of maternal strategies from their expression in the offspring phenotype alone. As an alternative approach, we examine divergence in the proximate mechanisms underlying maternal effects across three house finch populations with contrasting patterns of sex allocation: an ancestral population that shows no sex-biased ovulation, and two recently established populations at the northern and southern boundaries of the species range that have opposite sequences of ovulation of male and female eggs. For each population, we examined how oocyte acquisition of hormones, carotenoids and vitamins was affected by oocyte growth and overlap with the same and opposite sexes. Our results suggest that sex-specific acquisition of maternal resources and sex determination of oocytes are linked in this system. We report that acquisition of testosterone by oocytes that become males was not related to growth duration, but instead covaried with temporal exposure to steroids and overlap with other male oocytes. In female oocytes, testosterone acquisition increased with the duration of growth and overlap with male oocytes, but decreased with overlap with female oocytes. By contrast, acquisition of carotenoids and vitamins was mostly determined by organism-wide partitioning among oocytes and oocyte-specific patterns of testosterone accumulation, and these effects did not differ between the sexes. These results provide important insights into three unresolved phenomena in the evolution of maternal effects - (i) the evolution of sex-specific maternal allocation in species with simultaneously developing neonates of both sexes; (ii) the link between sex determination and sex-specific acquisition of maternal products; and (iii) the evolution of context-dependent modulation of maternal effects.
• Duckworth, R., Badyaev, A., & Parlow, A. (2007). Elaborately ornamented males avoid costly parental care in the house finch (Carpodacus mexicanus): a proximate perspective. BEHAVIORAL ECOLOGY AND SOCIOBIOLOGY, 55(2), 176-183.
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  Selection should favor flexibility in reproductive tactics when the combination of sexual traits and reproductive behaviors that achieve the highest fitness differs between males within a population. Understanding the functional significance of variation in male reproductive tactics can provide insight into their evolution. Male house finches (Carpodacus mexicanus) in a Montana population display continuous variation in parental tactics: males with more elaborated (redder) plumage color provide little or no parental care compared to less elaborated (dull) males. Here, we first determined whether elevation of prolactin (a pituitary hormone) was related to variation in male parental tactics and, second, we used the relationship between prolactin levels and parental behavior to investigate why redder males avoid a high investment in parental care. We found that prolactin elevation was closely associated with paternal care. In addition, males with redder plumage color had low prolactin levels, whereas dull males, which provision twice as frequently, had high levels of prolactin. We also found that male condition was unrelated to plumage color but negatively related to prolactin levels. These results suggest that the low provisioning of redder males was not due to physiological constraints, but instead reflected a tactic to avoid the costs associated with parental care. The condition benefits accrued by redder males may explain their higher post-breeding survival compared to dull males. Moreover, dull males were previously shown to have higher pairing success than redder males, suggesting that the relationship between male plumage color and parental care may reflect individually optimized parental tactics.
• Badyaev, A., Schwabl, H., Young, R., Duckworth, R., Navara, K., & Parlow, A. (2006). Adaptive sex differences in growth of pre-ovulation oocytes in a passerine bird. PROCEEDINGS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES, 272(1577), 2165-2172.
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  Maternal modification of offspring sex in birds has strong fitness consequences, however the mechanisms by which female birds can bias sex of their progeny in close concordance with the environment of breeding are not known. In recently established populations of house finches (Carpodacus mexicanus), breeding females lay a sex-biased sequence of eggs when ambient temperature causes early onset of incubation. We studied the mechanisms behind close association of incubation and sex-determination strategies in this species and discovered that pre-ovulation oocytes that produce males and females differed strongly in the temporal patterns of proliferation and growth. In turn, sex-specific exposure of oocytes to maternal secretion of prolactin and androgens produced distinct accumulation of maternal steroids in oocyte yolks in relation to oocyte proliferation order. These findings suggest that sex difference in oocyte growth and egg-laying sequence is an adaptive outcome of hormonal constraints imposed by the overlap of early incubation and oogenesis in this population, and that the close integration of maternal incubation, oocytes' sex-determination and growth might be under control of the same hormonal mechanism. We further document that population establishment and the evolution of these maternal strategies is facilitated by their strong effects on female and offspring fitness in a recently established part of the species range.
• Duckworth, R. A. (2006). Aggressive behaviour affects selection on morphology by influencing settlement patterns in a passerine bird. Proceedings of the Royal Society B: Biological Sciences, 273(1595), 1789-1795.
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  Abstract: The importance of behaviours as instigators or inhibitors of evolutionary change remains largely unresolved and this is in part because there are very few empirical examples of how behaviours affect evolutionary processes. By determining the environment of breeding, aggressive interactions over territories have the potential to strongly impact selection pressures experienced by individuals. Western bluebirds (Sialia mexicana) provide a unique opportunity to investigate the evolutionary importance of aggression, since their highly variable breeding habitat favours distinct foraging techniques and they also compete aggressively for nest boxes, a resource that is easy to manipulate. Here, I show experimentally that more aggressive males compete more effectively for territories with a high density of nest boxes and, as a consequence, aggressive and non-aggressive males are sorted into distinct breeding habitats that differ in the strength of selection on morphological traits. Specifically, males with longer tails and tarsi were favoured in open habitats where high agility is required to forage efficiently, whereas in forested habitats, where agility is less important, selection was weak. These results show that aggression can affect selection on a local scale by determining individual settlement patterns. More generally, because territorial interactions are important across a wide variety of taxa, these results suggest that aggressive behaviour has the potential to impact the evolutionary trajectory of many animal populations. © 2006 The Royal Society.
• Duckworth, R. A. (2006). Aggressive behaviour affects selection on morphology by influencing settlement patterns in a passerine bird.. Proceedings. Biological sciences / The Royal Society, 273(1595), 1789-1795.
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  PMID: 16790412;PMCID: PMC1634784;Abstract: The importance of behaviours as instigators or inhibitors of evolutionary change remains largely unresolved and this is in part because there are very few empirical examples of how behaviours affect evolutionary processes. By determining the environment of breeding, aggressive interactions over territories have the potential to strongly impact selection pressures experienced by individuals. Western bluebirds (Sialia mexicana) provide a unique opportunity to investigate the evolutionary importance of aggression, since their highly variable breeding habitat favours distinct foraging techniques and they also compete aggressively for nest boxes, a resource that is easy to manipulate. Here, I show experimentally that more aggressive males compete more effectively for territories with a high density of nest boxes and, as a consequence, aggressive and non-aggressive males are sorted into distinct breeding habitats that differ in the strength of selection on morphological traits. Specifically, males with longer tails and tarsi were favoured in open habitats where high agility is required to forage efficiently, whereas in forested habitats, where agility is less important, selection was weak. These results show that aggression can affect selection on a local scale by determining individual settlement patterns. More generally, because territorial interactions are important across a wide variety of taxa, these results suggest that aggressive behaviour has the potential to impact the evolutionary trajectory of many animal populations.
• Duckworth, R. A. (2006). Behavioral correlations across breeding contexts provide a mechanism for a cost of aggression. Behavioral Ecology, 17(6), 1011-1019.
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  Abstract: Identifying correlations among behaviors is important for understanding how selection shapes the phenotype. Correlated behaviors can indicate constraints on the evolution of behavioral plasticity or may reflect selection for functional integration among behaviors. Obligate cavity-nesting birds provide an opportunity to examine these correlations because males must defend limited nest cavities while also competing for mating opportunities and providing parental care. Here, I investigated the role of behavioral correlations in producing a counterintuitive relationship between nest defense and reproductive success in western bluebirds (Sialia mexicana) such that males that defended their nests most intensely had the lowest reproductive success, measured as the number of within and extrapair offspring that fledged. By experimentally measuring aggression across contexts, I show that this cost of nest defense was due to the correlated expression of aggression across the contexts of nest defense and male-male competition coupled with a trade-off between male-male aggression and parental care. In particular, more aggressive males provisioned their females less during incubation and this led to disrupted incubation patterns and fewer fledged offspring. However, aggressive males did not benefit from avoiding parental investment by gaining extrapair fertilizations, and thus, it is unclear how high levels of aggression are maintained in this population despite apparent costs. These results suggest that there are constraints to the evolution of plasticity in aggression and emphasize the importance of considering the integrated behavioral phenotype to understand how variation in behavior is linked to fitness. © The Author 2006. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology. All rights reserved.
• Duckworth, R. A., & Sockman, K. W. (2006). Proximate mechanisms of behavioural inflexibility: implications for the evolution of personality traits. FUNCTIONAL ECOLOGY, 26(3), 559-566.
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  1. Behaviour is often assumed to be the most flexible of traits, yet recent studies show a high repeatability of behaviour within individuals even across different functional contexts. Such consistent expression of behaviour may evolve either when selection favours its integration with less flexible components of the phenotype or when pleiotropic effects produce correlations between behaviours that have different optimal timing of expression. Examining the physiological mechanisms underlying correlated expression of behaviours provides powerful insight into the evolution of personalities by establishing the extent to which pleiotropic effects might limit the independent evolution of distinct behaviours. 2. Here, we investigated proximate mechanisms behind aggressive and non-aggressive personality types in western bluebirds, Sialia mexicana, to determine whether consistency in the expression of aggression is because of shared effects of plasma-circulating androgens on aggression and mating behaviour. 3. We found that androgen concentration was unrelated to variation in both intra- and interspecific aggression even though it was closely linked to variation in male mating behaviour. These results suggest that pleiotropic effects of circulating androgens are unlikely to cause consistent differences among individuals in aggression. 4. These findings suggest that decoupling of the activational effects of hormones on behaviour is an important step in the evolution of personality traits.
• Duckworth, R., Badyaev, A., Farmer, K., Hill, G., & Roberts, . (2006). First case of Mycoplasma gallisepticum infection in the western range of the house finch (Carpodacus mexicanus). AUK, 120(2), 528-530.
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  We report the first case of mycoplasmosis in the western range of the House Finch (Carpodacus mexicanus). This disease originated in the eastern United States and has been previously documented only in eastern introduced House Finch populations where it reached epizootic proportions causing extensive and widespread mortality. Documentation of this disease in western Montana suggests that previously disjunct eastern and western populations of House Finches are now mixing in the northern part of their range. More importantly, as native House Finches are highly susceptible to this novel pathogen, western populations may now be at risk of high mortality, similar to that experienced by non-native eastern populations. Close monitoring of this disease in the western part of the House Finch range will provide important insight into the dynamics of the emerging disease and evolution of resistance to the pathogen.
• Duckworth, R. A. (2004). Behavioral correlations across breeding contexts provide a mechanism for a cost of aggression. BEHAVIORAL ECOLOGY, 17(6), 1011-1019.
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  Identifying correlations among behaviors is important for understanding how selection shapes the phenotype. Correlated behaviors can indicate constraints on the evolution of behavioral plasticity or may reflect selection for functional integration among behaviors. Obligate cavity-nesting birds provide an opportunity to examine these correlations because males must defend limited nest cavities while also competing for mating opportunities and providing parental care. Here, I investigated the role of behavioral correlations in producing a counterintuitive relationship between nest defense and reproductive success in western bluebirds (Sialia mexicana) such that males that defended their nests most intensely had the lowest reproductive success, measured as the number of within and extrapair offspring that fledged. By experimentally measuring aggression across contexts, I show that this cost of nest defense was due to the correlated expression of aggression across the contexts of nest defense and male-male competition coupled with a trade-off between male-male aggression and parental care. In particular, more aggressive males provisioned their females less during incubation and this led to disrupted incubation patterns and fewer fledged offspring. However, aggressive males did not benefit from avoiding parental investment by gaining extrapair fertilizations, and thus, it is unclear how high levels of aggression are maintained in this population despite apparent costs. These results suggest that there are constraints to the evolution of plasticity in aggression and emphasize the importance of considering the integrated behavioral phenotype to understand how variation in behavior is linked to fitness.
• Duckworth, R. A., Mendonça, M. T., & Hill, G. E. (2004). Condition-dependent sexual traits and social dominance in the house finch. Behavioral Ecology, 15(5), 779-784.
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  Abstract: Elaboration of costly sexual traits can reduce investment in other aspects of reproduction, such as parental care or intrasexual competition, which may lead to the evolution of alternative mating tactics. In house finches (Carpodacus mexicanus), less elaborately ornamented (dull) males tend to dominate more elaborated (redder) males, but redder males pair earlier and invest more in parental care. This suggests that males may pursue alternative parental or competitive tactics, depending on the elaboration of their sexual trait. Elevation of testosterone, a hormone that is closely associated with condition in male house finches, influences dominance and sexual behaviors but is antagonistic to parental behaviors. We tested the hypothesis that the higher dominance status of dull males reflects an alternative testosterone-dependent mating tactic. First, we experimentally manipulated the testosterone levels of captive males and measured the effect on dominance rank, and second, we measured the association of testosterone elevation and plumage hue in free-living males. We found that, as predicted, testosterone elevation increased dominance rank in captive males. However, in free-living males, testosterone levels were higher in redder males, suggesting that testosterone is dissociated from dominance status under natural circumstances. This may be because the context of social interactions and the higher motivation of dull males to access food resources have a stronger influence on the outcome of dominance interactions than does the physiological effects of testosterone elevation. In turn, the strong positive correlation between testosterone levels and plumage elaboration likely reflects the common condition dependence of these traits.
• Badyaev, A., Hill, G., Beck, M., Dervan, A., Duckworth, R., McGraw, K., Nolan, P., & Whittingham, L. (2003). Sex-biased hatching order and adaptive population divergence in a passerine bird. SCIENCE, 295(5553), 316-318.
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  Most species of birds can lay only one egg per day until a clutch is complete, and the order in which eggs are laid often has strong and sex-specific effects on offspring growth and survival. In two recently established populations of the house finch (Carpodacus mexicanus) in Montana and Alabama, breeding females simultaneously adjusted the sex and growth of offspring in relation to their position in the laying order, thereby reducing the mortality of sons and daughters by 10 to 20% in both environments. We show experimentally that the reduction in mortality is produced by persistent and sex-specific maternal effects on the growth and morphology of offspring, These strong parental effects may have facilitated the rapid adaptive divergence among populations of house finches.
• Duckworth, R. A. (2003). Adaptive dispersal strategies and the dynamics of a range expansion. AMERICAN NATURALIST, 172, S4-S17.
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  In species undergoing range expansion, newly established populations are often more dispersive than older populations. Because dispersal phenotypes are complex and often costly, it is unclear how highly dispersive phenotypes are maintained in a species to enable their rapid expression during periods of range expansion. Here I test the idea that metapopulation dynamics of local extinction and recolonization maintain distinct dispersal strategies outside the context of range expansion. Western bluebirds display distinct dispersal phenotypes where aggressive males are more dispersive than nonaggressive males, resulting in highly aggressive populations at the edge of their expanding range. I experimentally created new habitat interior to the range edge to show that, as on the range front, it was colonized solely by aggressive males. Moreover, fitness consequences of aggression depended on population age: aggressive males had high fitness when colonizing new populations, while nonaggressive males performed best in an older population. These results suggest that distinct dispersal strategies were maintained before range expansion as an adaptation for the continual recolonization of new habitat. These results emphasize similarities between range expansion and metapopulation dynamics and suggest that preexisting adaptive dispersal strategies may explain rapid changes in dispersal phenotypes during range expansion.
• Duckworth, R., Badyaev, A. V., & Duckworth, R. A. (2003). Context-dependent sexual advertisement: plasticity in development of sexual ornamentation throughout the lifetime of a passerine bird. Journal of evolutionary biology, 16(6).
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  Male investment into sexual ornamentation is a reproductive decision that depends on the context of breeding and life history state. In turn, selection for state- and context-specific expression of sexual ornamentation should favour the evolution of developmental pathways that enable the flexible allocation of resources into sexual ornamentation. We studied lifelong variation in the expression and condition-dependence of a sexual ornament in relation to age and the context of breeding in male house finches (Carpodacus mexicanus)--a species that develops a new sexual ornament once a year after breeding. Throughout males' lifetime, the elaboration of ornamentation and the allocation of resources to the development of sexual ornamentation depended strongly on pairing status in the preceding breeding season--males that were single invested more resources into sexual ornamentation and changed ornamentation more than males that were paired. During the initial (post-juvenile) moult, the expression of ornamentation was closely dependent on individual condition, however the condition-dependence of ornamentation sharply decreased throughout a male's lifetime and in older males expression of sexual ornamentation was largely independent of condition during moult. Selection for early breeding favoured greater ornamentation in males that were single in the preceding seasons and the strength of this selection increased with age. On the contrary, the strength of selection on sexual ornamentation decreased with age in males that were paired in the preceding breeding season. Our results reveal strong context-dependency in investment into sexual ornamentation as well as a high flexibility in the development of sexual ornamentation throughout a male's life.
• Duckworth, R. A., Mendonça, M., & Hill, G. E. (2001). A condition dependent link between testosterone and disease resistance in the house finch. Proceedings of the Royal Society B: Biological Sciences, 268(1484), 2467-2472.
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  PMID: 11747565;PMCID: PMC1088901;Abstract: Testosterone has recently been proposed as a link between male quality and health and the expression of sexual traits. We investigated the relationship between testosterone and measures of the individual condition and health of males in a natural population of house finches (Carpodacus mexicanus). We also conducted a captive experiment in order to test for the effects of testosterone on resistance to coccidia, which is a common parasite of house finches. Free-living males in better condition had higher testosterone levels and lower corticosterone levels than free-living males in poor condition. In our captive experiment, increased testosterone accelerated the rate of coccidial infection as compared with sham-implanted or gonadectomized males. Although the differences were not significant, free-living males infected with coccidia had lower levels of testosterone and higher levels of corticosterone than males that were not infected. Thus, experimentally elevating testosterone levels in captive males resulted in a higher percentage of infected males, while free-living males with coccidial infection had low testosterone levels. This apparent discrepancy between captive and free-living males in the association of testosterone and disease may be explained by the condition dependence of testosterone. These results suggest that the testosterone-dependent sexual traits reliably indicate male overall condition and health and, thus, females could benefit fi-om assessing potential mates based on these traits.
• Badyaev, A. V., Young, R. L., Hill, G. E., & Duckworth, R. A. (2000). Evolution of sex-biased maternal effects in birds. IV. Intra-ovarian growth dynamics can link sex determination and sex-specific acquisition of resources. JOURNAL OF EVOLUTIONARY BIOLOGY, 21(2), 449-460.
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  The evolutionary importance of maternal effects is determined by the interplay of maternal adaptations and strategies, offspring susceptibility to these strategies, and the similarity of selection pressures between the two generations. Interaction among these components, especially in species where males and females differ in the costs and requirements of growth, limits inference about the evolution of maternal strategies from their expression in the offspring phenotype alone. As an alternative approach, we examine divergence in the proximate mechanisms underlying maternal effects across three house finch populations with contrasting patterns of sex allocation: an ancestral population that shows no sex-biased ovulation, and two recently established populations at the northern and southern boundaries of the species range that have opposite sequences of ovulation of male and female eggs. For each population, we examined how oocyte acquisition of hormones, carotenoids and vitamins was affected by oocyte growth and overlap with the same and opposite sexes. Our results suggest that sex-specific acquisition of maternal resources and sex determination of oocytes are linked in this system. We report that acquisition of testosterone by oocytes that become males was not related to growth duration, but instead covaried with temporal exposure to steroids and overlap with other male oocytes. In female oocytes, testosterone acquisition increased with the duration of growth and overlap with male oocytes, but decreased with overlap with female oocytes. By contrast, acquisition of carotenoids and vitamins was mostly determined by organism-wide partitioning among oocytes and oocyte-specific patterns of testosterone accumulation, and these effects did not differ between the sexes. These results provide important insights into three unresolved phenomena in the evolution of maternal effects - (i) the evolution of sex-specific maternal allocation in species with simultaneously developing neonates of both sexes; (ii) the link between sex determination and sex-specific acquisition of maternal products; and (iii) the evolution of context-dependent modulation of maternal effects.
• Nolan, P. M., Duckworth, R. A., Hill, G. E., & Roberts, S. R. (2000). Maintenance of a captive flock of house finches free of infection by Mycoplasma gallisepticum. Avian Diseases, 44(4), 948-952.
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  PMID: 11195652;Abstract: Since the beginning of an epidemic of conjunctivitis in wild house finches caused by Mycoplasma gallisepticum (MG), all captive colonies established by capturing free-ranging house finches from the eastern population have also either been infected at the time of capture or developed infection shortly after capture. In an attempt to avoid this infection in captive flocks being maintained for studies of the finches' behavior and ecology, we compared two different flock management strategies and were able to prevent the development of mycoplasmal conjunctivitis with one of the strategies. Single-sex flocks were built by introducing only seronegative wild-caught birds showing no clinical signs of conjunctivitis and covering their outdoor flight cages with netting to prevent interaction with other wild birds although only the female flocks were initially treated with a 6-wk course of tylosin tartrate (0.3 mg/ml). The female flocks never developed conjunctivitis although the disease did develop in the male flocks. Furthermore, serologic assessments of the healthy flock by serum plate agglutination assays for MG indicated that the females remained free of MG infection in the final 7 wk of the study, during which they were unmedicated. We conclude that any low-level MG infection not diagnosed by the initial test for seroconversion was cleared by the prolonged drug treatment.

Others

• Duckworth, R. A., & Sockman, K. W. (2015, DEC). Evolution of avian personalities: Adaptive variation or hormonal constraint?. INTEGRATIVE AND COMPARATIVE BIOLOGY.
• Duckworth, R. A. (2005, OCT). EVOLUTION OF PERSONALITY: DEVELOPMENTAL CONSTRAINTS ON BEHAVIORAL FLEXIBILITY. AUK.
• Duckworth, R., Mendonca, M., & Hill, G. (2003). The effect of testosterone manipulation on parasite resistance in the house finch.. AMERICAN ZOOLOGIST.