House Sparrows as Sentinels of Childhood Lead Exposure.

Our understanding of connections between human and animal health has advanced substantially since the canary was introduced as a sentinel of toxic conditions in coal mines. Nonetheless, the development of wildlife sentinels for monitoring human exposure to toxins has been limited. Here, we capitalized on a three-decade long child blood lead monitoring program to demonstrate that the globally ubiquitous and human commensal house sparrow (Passer domesticus) can be used as a sentinel of human health risks in urban environments impacted by lead mining. We showed that sparrows are a viable proxy for the measurement of blood lead levels in children at a neighborhood scale (0.28 km2). In support of the generalizability of this approach, the blood lead relationship established in our focal mining city enabled us to accurately predict elevated blood lead levels in children from another mining city using only sparrows from the second location. Using lead concentrations and lead isotopic compositions from environmental and biological matrices, we identified shared sources and pathways of lead exposure in sparrows and children, with strong links to contamination from local mining emissions. Our findings showed how human commensal species can be used to identify and predict human health risks over time and space.

Exposure to Sublethal Concentrations of Lead (Pb) Affects Ecologically Relevant Behaviors in House Sparrows (Passer domesticus).

Global contamination of environments with lead (Pb) poses threats to many ecosystems and populations. While exposure to Pb is toxic at high concentrations, recent literature has shown that lower concentrations can also cause sublethal, deleterious effects. However, there remains relatively little causal investigation of how exposure to lower concentrations of environmental Pb affects ecologically important behaviors. Behaviors often represent first-line responses of an organism and its internal physiological, molecular, and genetic responses to a changing environment. Hence, better understanding how behaviors are influenced by pollutants such as Pb generates crucial information on how species are coping with the effects of pollution more broadly. To better understand the effects of sublethal Pb on behavior, we chronically exposed adult wild-caught, captive house sparrows (Passer domesticus) to Pb-exposed drinking water and quantified a suite of behavioral outcomes: takeoff flight performance, activity in a novel environment, and in-hand struggling and breathing rate while being handled by an experimenter. Compared to controls (un-exposed drinking water), sparrows exposed to environmentally relevant concentrations of Pb exhibited decreases in takeoff flight performance and reduced movements in a novel environment following 9-10 weeks of exposure. We interpret this suite of results to be consistent with Pb influencing fundamental neuro-muscular abilities, making it more difficult for exposed birds to mount faster movements and activities. It is likely that suppression of takeoff flight and reduced movements would increase the predation risk of similar birds in the wild; hence, we also conclude that the effects we observed could influence fitness outcomes for individuals and populations altering ecological interactions within more naturalistic settings.

Mercury causes degradation of spatial cognition in a model songbird species.

Mercury is a widespread pollutant of increasing global concern that exhibits a broad range of deleterious effects on organisms, including birds. Because the developing brain is well-known to be particularly vulnerable to the neurotoxic insults of mercury, many studies have focused on developmental effects such as on the embryonic brain and resulting behavioral impairment in adults. It is not well understood how the timing of exposure, for example exclusively in ovo versus throughout life, influences the impact of mercury. Using dietary exposure to environmentally relevant methylmercury concentrations, we examined the role that timing and duration of exposure play on spatial learning and memory in a model songbird species, the domesticated zebra finch (Taeniopygia guttata castanotis). We hypothesized that developmental exposure was both necessary and sufficient to disrupt spatial memory in adult finches. We documented profound disruption of memory for locations of hidden food at two spatial scales, cage- and room-sized enclosures, but found that both developmental and ongoing adult exposure were required to exhibit this behavioral impairment. Methylmercury-exposed birds made more mistakes before mastering the spatial task, because they revisited unrewarded locations repeatedly even after discovering the rewarded location. Contrary to our prediction, hippocampal volume was not affected in birds exposed to methylmercury over their lifetimes. The disruption of spatial cognition that we detected is severe and would likely have implications for survival and reproduction in wild birds; however, it appears that individuals that disperse or migrate from a contaminated site might recover later in life if no longer exposed to the toxicant.

Window films increase avoidance of collisions by birds but only when applied to external compared with internal surfaces of windows.

Window collisions are one of the largest human-caused causes of avian mortality in built environments and, therefore, cause population declines that can be a significant conservation issue. Applications of visibly noticeable films, patterns, and decals on the external surfaces of windows have been associated with reductions in both window collisions and avian mortality. It is often logistically difficult and economically prohibitive to apply these films and decals to external surfaces, especially if the windows are above the first floor of a building. Therefore, there is interest and incentive to apply the products to internal surfaces that are much easier to reach and maintain. However, there is debate as to whether application to the internal surface of windows renders any collision-reduction benefit, as the patterns on the films and decals may not be sufficiently visible to birds. To address this knowledge gap, we performed the first experimental study to compare the effectiveness of two distinct window films when applied to either the internal or external surface of double-glazed windows. Specifically, we assessed whether Haverkamp and BirdShades window film products were effective in promoting the avoidance of window collisions (and by inference, a reduction of collisions) by zebra finches through controlled aviary flight trials employing a repeated-measures design that allowed us to isolate the effect of the window treatments on avoidance flight behaviors. We chose these two products because they engage with different wavelengths of light (and by inference, colors) visible to many songbirds: the BirdShades film is visible in the ultraviolet (shorter wavelength) range, while the Haverkamp film includes signals in the orange (longer wavelength) range. We found consistent evidence that, when applied to the external surface of windows, the BirdShades product resulted in reduced likelihood of collision and there was marginal evidence of this effect with the Haverkamp film. Specifically, in our collision avoidance trials, BirdShades increased window avoidance by 47% and the Haverkamp increased avoidance by 39%. However, neither product was effective when the films were applied to the internal surface of windows. Hence, it is imperative that installers apply these products to exterior surfaces of windows to render their protective benefits and reduce the risk of daytime window collision.

Evaluating acoustic signals to reduce avian collision risk.

Collisions with human-made structures are responsible for billions of bird deaths each year, resulting in ecological damage as well as regulatory and financial burdens to many industries. Acoustic signals can alert birds to obstacles in their flight paths in order to mitigate collisions, but these signals should be tailored to the sensory ecology of birds in flight as the effectiveness of various acoustic signals potentially depends on the influence of background noise and the relative ability of various sound types to propagate within a landscape. We measured changes in flight behaviors from zebra finches released into a flight corridor containing a physical obstacle, either in no-additional-sound control conditions or when exposed to one of four acoustic signals. We selected signals to test two frequency ranges (4-6 kHz or 6-8 kHz) and two temporal modulation patterns (broadband or frequency-modulated oscillating) to determine whether any particular combination of sound attributes elicited the strongest collision avoidance behaviors. We found that, relative to control flights, all sound treatments caused birds to maintain a greater distance from hazards and to adjust their flight trajectories before coming close to obstacles. There were no statistical differences among different sound treatments, but consistent trends within the data suggest that the 4-6 kHz frequency-modulated oscillating signal elicited the strongest avoidance behaviors. We conclude that a variety of acoustic signals can be effective as avian collision deterrents, at least in the context in which we tested these birds. These results may be most directly applicable in scenarios when birds are at risk of collisions with solid structures, such as wind turbines and communication towers, as opposed to window collisions or collisions involving artificial lighting. We recommend the incorporation of acoustic signals into multimodal collision deterrents and demonstrate the value of using behavioral data to assess collision risk.

Field testing an "acoustic lighthouse": Combined acoustic and visual cues provide a multimodal solution that reduces avian collision risk with tall human-made structures.

Billions of birds fatally collide with human-made structures each year. These mortalities have consequences for population viability and conservation of endangered species. This source of human-wildlife conflict also places constraints on various industries. Furthermore, with continued increases in urbanization, the incidence of collisions continues to increase. Efforts to reduce collisions have largely focused on making structures more visible to birds through visual stimuli but have shown limited success. We investigated the efficacy of a multimodal combination of acoustic signals with visual cues to reduce avian collisions with tall structures in open airspace. Previous work has demonstrated that a combination of acoustic and visual cues can decrease collision risk of birds in captive flight trials. Extending to field tests, we predicted that novel acoustic signals would combine with the visual cues of tall communication towers to reduce collision risk for birds. We broadcast two audible frequency ranges (4 to 6 and 6 to 8 kHz) in front of tall communication towers at locations in the Atlantic migratory flyway of Virginia during annual migration and observed birds' flight trajectories around the towers. We recorded an overall 12-16% lower rate of general bird activity surrounding towers during sound treatment conditions, compared with control (no broadcast sound) conditions. Furthermore, in 145 tracked "at-risk" flights, birds reduced flight velocity and deflected flight trajectories to a greater extent when exposed to the acoustic stimuli near the towers. In particular, the 4 to 6 kHz stimulus produced the greater effect sizes, with birds altering flight direction earlier in their trajectories and at larger distances from the towers, perhaps indicating that frequency range is more clearly audible to flying birds. This "acoustic lighthouse" concept reduces the risk of collision for birds in the field and could be applied to reduce collision risk associated with many human-made structures, such as wind turbines and tall buildings.

Ultraviolet-reflective film applied to windows reduces the likelihood of collisions for two species of songbird.

Perhaps a billion birds die annually from colliding with residential and commercial windows. Therefore, there is a societal need to develop technologies that reduce window collisions by birds. Many current window films that are applied to the external surface of windows have human-visible patterns that are not esthetically preferable. BirdShades have developed a short wavelength (ultraviolet) reflective film that appears as a slight tint to the human eye but should be highly visible to many bird species that see in this spectral range. We performed flight tunnel tests of whether the BirdShades external window film reduced the likelihood that two species of song bird (zebra finch, Taeniopygia guttata and brown-headed cowbird, Molothrus ater) collide with windows during daylight. We paid particular attention to simulate the lighting conditions that birds will experience while flying during the day. Our results indicate a 75-90% reduction in the likelihood of collision with BirdShades-treated compared with control windows, in forced choice trials. In more ecologically relevant comparison between trials where all windows were either treated or control windows, the estimated reduction in probability of collision was 30-50%. Further, both bird species slow their flight by approximately 25% when approaching windows treated with the BirdShades film, thereby reducing the force of collisions if they were to happen. Therefore, we conclude that the BirdShades external window film will be effective in reducing the risk of and damage caused to populations and property by birds' collision with windows. As this ultraviolet-reflective film has no human-visible patterning to it, the product might be an esthetically more acceptable low cost solution to reducing bird-window collisions. Further, we call for testing of other mitigation technologies in lighting and ecological conditions that are more similar to what birds experience in real human-built environments and make suggestions for testing standards to assess collision-reducing technologies.

Sexually selected traits as bioindicators: exposure to mercury affects carotenoid-based male bill color in zebra finches.

To examine whether sexually selected traits are particularly sensitive bioindicators of environmental toxicants, we assessed the effects of exposure to environmentally relevant dietary concentrations of the pollutant methylmercury on pigment coloration in zebra finches (Taeniopygia guttata). First, we tested whether effects of methylmercury on coloration were influenced by timing of exposure. Birds were either exposed developmentally (up to 114 days after hatching), as adults (after reaching sexual maturity), or for their entire life. Bill coloration, which is a carotenoid-based, sexually selected trait, was less red in males with lifetime exposure to methylmercury, compared to controls. Neither adult, nor developmental exposure influenced bill color in adult males, with the possible exception of early exposure of nestlings. Among females, where bill color is not under strong sexual selection, neither lifetime nor adult exposure to methylmercury affected bill color. For males and females, there was no effect of either lifetime or adult methylmercury exposure on coloration of back feathers, which is a non-sexually-dimorphic, melanin-based trait that is not likely the result of sexual selection. This study is a comprehensive experimental test of the proposal that sexually selected traits may be particularly useful bioindicators of the stress imposed by environmental toxins such as methylmercury.

Mercury delays cerebellar development in a model songbird species, the zebra finch.

Mercury exposure can disrupt development of the cerebellum, part of the brain essential for coordination of movement through a complex environment, including flight. In precocial birds, such as fowl, the cerebellum develops embryonically, and the chick is capable of leaving the nest within hours of hatching. However, most birds, including all songbirds, are altricial, and spend weeks in the nest between hatching and fledging. The objective of this study was to describe the normal development of the cerebellum in a model altricial songbird so as to determine the effect of exposure to mercury on cerebellar maturation. Adult zebra finch (Taeniopygia guttata) pairs were fed either a control diet, or a diet augmented with one of four treatment-levels of methylmercury (0.3-2.4 µg/g wet weight), and their offspring, the subjects of this study, were fed the same diet by parents. We documented, for the first time, the schedule of cerebellar development in an altricial bird, and compared stages of development among methylmercury-exposed groups. For all treatments of methylmercury, the age of completion of cellular migration was later than for control zebra finches, indicating a delay in cerebellar maturation. Displaced (heterotopic) Purkinje neurons, a pathology typical of methylmercury exposure in developing vertebrate brains, were more numerous in methylmercury-exposed birds, and persisted at least until the age of independence. Delays in maturation of the cerebellum could delay fledging in altricial bird species, with potential serious implications for the fitness of exposed individuals, as predation rates in the nest are often very high.

Shifts in sexual dimorphism across a mass extinction in ostracods: implications for sexual selection as a factor in extinction risk.

Sexual selection often favours investment in expensive sexual traits that help individuals compete for mates. In a rapidly changing environment, however, allocation of resources to traits related to reproduction at the expense of those related to survival may elevate extinction risk. Empirical testing of this hypothesis in the fossil record, where extinction can be directly documented, is largely lacking. The rich fossil record of cytheroid ostracods offers a unique study system in this context: the male shell is systematically more elongate than that of females, and thus the sexes can be distinguished, even in fossils. Using mixture models to identify sex clusters from size and shape variables derived from the digitized valve outlines of adult ostracods, we estimated sexual dimorphism in ostracod species before and after the Cretaceous/Palaeogene mass extinction in the United States Coastal Plain. Across this boundary, we document a substantial shift in sexual dimorphism, driven largely by a pronounced decline in the taxa with dimorphism indicating both very high and very low male investment. The shift away from high male investment, which arises largely from evolutionary changes within genera that persist through the extinction, parallels extinction selectivity previously documented during the Late Cretaceous under a background extinction regime. Our results suggest that sexual selection and the allocation of resources towards survival versus reproduction may be an important factor for species extinction during both background and mass extinctions.

High male sexual investment as a driver of extinction in fossil ostracods.

Sexual selection favours traits that confer advantages in the competition for mates. In many cases, such traits are costly to produce and maintain, because the costs help to enforce the honesty of these signals and cues 1 . Some evolutionary models predict that sexual selection also produces costs at the population level, which could limit the ability of populations to adapt to changing conditions and thus increase the risk of extinction2-4. Other models, however, suggest that sexual selection should increase rates of adaptation and enhance the removal of deleterious mutations, thus protecting populations against extinction3, 5, 6. Resolving the conflict between these models is not only important for explaining the history of biodiversity, but also relevant to understanding the mechanisms of the current biodiversity crisis. Previous attempts to test the conflicting predictions produced by these models have been limited to extant species and have thus relied on indirect proxies for species extinction. Here we use the informative fossil record of cytheroid ostracods-small, bivalved crustaceans with sexually dimorphic carapaces-to test how sexual selection relates to actual species extinction. We show that species with more pronounced sexual dimorphism, indicating the highest levels of male investment in reproduction, had estimated extinction rates that were ten times higher than those of the species with the lowest investment. These results indicate that sexual selection can be a substantial risk factor for extinction.

Exposure to Dietary Methyl-Mercury Solely during Embryonic and Juvenile Development Halves Subsequent Reproductive Success in Adult Zebra Finches.

Long-term exposure to methyl-mercury has deleterious effects on avian reproduction. However, little is known about whether exposure to mercury solely during embryonic and juvenile development can have long-lasting effects on subsequent reproductive performance as adults. Birds that hatch on contaminated sites but disperse elsewhere will be exposed only during development. Hence, it is important to understand the reproductive consequences of avian exposure to methyl-mercury during early life. Accordingly, in this experiment, domesticated zebra finches ( Taeniopygia guttata) were exposed to dietary methyl-mercury (1.20 µg/g wet weight) from conception through independence (50 days post-hatching). Following maturity, developmentally exposed and control finches were paired within treatment groups and allowed to breed repeatedly for 8 months. Developmentally exposed pairs hatched 32% fewer eggs and produced 50% fewer independent juveniles despite transferring only traces of mercury into their offspring. This is the first example of mercury-related reproductive declines in birds not exposed to mercury during breeding. The magnitude of reproductive decline was similar to that of zebra finches exposed to methyl-mercury during the breeding process. Bird populations breeding in contaminated habitats may suffer from a 2-fold fitness cost of mercury exposure; adult exposure compromises parents' reproduction, while offspring exposure results in reduced reproduction in the next generation.

Experimental Infection and Clearance of Coccidian Parasites in Mercury-Exposed Zebra Finches.

Mercury is a globally distributed, persistent environmental contaminant that affects the health of many taxa. It can suppress the immune system, which often plays a role in defense against parasites. However, there have been few investigations of whether mercury affects the abilities of animals to resist parasitic infection. Here, we exposed zebra finches to a lifetime dietary exposure of methylmercury (1.2 µg/g wet weight) and experimentally infected them with coccidian parasites to examine the effect of methylmercury exposure on parasitic infection. The mercury-exposed birds did not have an altered immune response (heterophil:lymphocyte ratio) nor a reduced ability to clear the infection. However, mercury-exposed birds tended to have higher parasite loads at the time when we expected the greatest immune response (2-3 weeks post-infection). Although mercury did not greatly influence the infection-course of this parasite in captivity, responses may be more accentuated in the wild where birds face additional immune challenges.

Using a Sound Field to Reduce the Risks of Bird-Strike: An Experimental Approach.

SYNOPSIS: Each year, billions of birds collide with large human-made structures, such as building, towers, and turbines, causing substantial mortality. Such bird-strike, which is projected to increase, poses risks to populations of birds and causes significant economic costs to many industries. Mitigation technologies have been deployed in an attempt to reduce bird-strike, but have been met with limited success. One reason for bird-strike may be that birds fail to pay adequate attention to the space directly in front of them when in level, cruising flight. A warning signal projected in front of a potential strike surface might attract visual attention and reduce the risks of collision. We tested this idea in captive zebra finches (Taeniopygia guttata) that were trained to fly down a long corridor and through an open wooden frame. Once birds were trained, they each experienced three treatments at unpredictable times and in a randomized order: a loud sound field projected immediately in front of the open wooden frame; a mist net (i.e., a benign strike surface) placed inside the wooden frame; and both the loud sound and the mist net. We found that birds slowed their flight approximately 20% more when the sound field was projected in front of the mist net compared with when the mist net was presented alone. This reduction in velocity would equate to a substantial reduction in the force of any collision. In addition to slowing down, birds increased the angle of attack of their body and tail, potentially allowing for more maneuverable flight. Concomitantly, the only cases where birds avoided the mist net occurred in the sound-augmented treatment. Interestingly, the sound field by itself did not demonstrably alter flight. Although our study was conducted in a limited setting, the alterations of flight associated with our sound field has implications for reducing bird-strike in nature and we encourage researchers to test our ideas in field trials.

Correlation between investment in sexual traits and valve sexual dimorphism in Cyprideis species (Ostracoda).

Assessing the long-term macroevolutionary consequences of sexual selection has been hampered by the difficulty of studying this process in the fossil record. Cytheroid ostracodes offer an excellent system to explore sexual selection in the fossil record because their readily fossilized carapaces are sexually dimorphic. Specifically, males are relatively more elongate than females in this superfamily. This sexual shape difference is thought to arise so that males carapaces can accommodate their very large copulatory apparatus, which can account for up to one-third of body volume. Here we test this widely held explanation for sexual dimorphism in cytheroid ostracodes by correlating investment in male genitalia, a trait in which sexual selection is seen as the main evolutionary driver, with sexual dimorphism of carapace in the genus Cyprideis. We analyzed specimens collected in the field (C. salebrosa, USA; C. torosa, UK) and from collections of the National Museum of Natural History, Washington, DC (C. mexicana). We digitized valve outlines in lateral view to obtain measures of size (valve area) and shape (elongation, measured as length to height ratio), and obtained several dimensions from two components of the hemipenis: the muscular basal capsule, which functions as a sperm pump, and the section that includes the intromittent organ (terminal extension). In addition to the assessment of this primary sexual trait, we also quantified two dimensions of the male secondary sexual trait-where the transformed right walking leg functions as a clasping organ during mating. We also measured linear dimensions from four limbs as indicators of overall (soft-part) body size, and assessed allometry of the soft anatomy. We observed significant correlations in males between valve size, but not elongation, and distinct structural parts of the hemipenis, even after accounting for their shared correlation with overall body size. We also found weak but significant positive correlation between valve elongation and the degree of sexual dimorphism of the walking leg, but only in C. torosa. The correlation between the hemipenis parts, especially basal capsule size and male valve size dimorphism suggests that sexual selection on sperm size, quantity, and/or efficiency of transfer may drive sexual size dimorphism in these species, although we cannot exclude other aspects of sexual and natural selection.

Methylmercury Exposure Reduces the Auditory Brainstem Response of Zebra Finches (Taeniopygia guttata ).

Mercury contamination from mining and fossil fuel combustion causes damage to humans and animals worldwide. Mercury exposure has been implicated in mammalian hearing impairment, but its effect on avian hearing is unknown. In this study, we examined whether lifetime dietary mercury exposure affected hearing in domestic zebra finches (Taeniopygia guttata) by studying their auditory brainstem responses (ABRs). Zebra finches exposed to mercury exhibited elevated hearing thresholds, decreased amplitudes, and longer latencies in the ABR, the first evidence of mercury-induced hearing impairment in birds. Birds are a more appropriate model for the human auditory spectrum than most mammals because of similarities in frequency discrimination, vocal learning, and communication behavior. When mercury is considered in combination with other anthropogenic stressors such as noise pollution and habitat alteration, the hearing impairments we document here could substantially degrade avian auditory communication in wild birds.

Integrative behavioral ecotoxicology: bringing together fields to establish new insight to behavioral ecology, toxicology, and conservation.

The fields of behavioral ecology, conservation science, and environmental toxicology individually aim to protect and manage the conservation of wildlife in response to anthropogenic stressors, including widespread anthropogenic pollution. Although great emphasis in the field of toxicology has been placed on understanding how single pollutants affect survival, a comprehensive, interdisciplinary approach that includes behavioral ecology is essential to address how anthropogenic compounds are a risk for the survival of species and populations in an increasingly polluted world. We provide an integrative framework for behavioral ecotoxicology using Tinbergen's four postulates (causation and mechanism, development and ontogeny, function and fitness, and evolutionary history and phylogenetic patterns). The aims of this review are: 1) to promote an integrative view and re-define the field of integrative behavioral ecotoxicology; 2) to demonstrate how studying ecotoxicology can promote behavior research; and 3) to identify areas of behavioral ecotoxicology that require further attention to promote the integration and growth of the field.

Exposure to dietary mercury alters cognition and behavior of zebra finches.

Environmental stressors can negatively affect avian cognitive abilities, potentially reducing fitness, for example by altering response to predators, display to mates, or memory of locations of food. We expand on current knowledge by investigating the effects of dietary mercury, a ubiquitous environmental pollutant and known neurotoxin, on avian cognition. Zebra finches Taeniopygia guttata were dosed for their entire lives with sub-lethal levels of mercury, at the environmentally relevant dose of 1.2 parts per million. In our first study, we compared the dosed birds with controls of the same age using tests of three cognitive abilities: spatial memory, inhibitory control, and color association. In the spatial memory assay, birds were tested on their ability to learn and remember the location of hidden food in their cage. The inhibitory control assay measured their ability to ignore visible but inaccessible food in favor of a learned behavior that provided the same reward. Finally, the color association task tested each bird's ability to associate a specific color with the presence of hidden food. Dietary mercury negatively affected spatial memory ability but not inhibitory control or color association. Our second study focused on three behavioral assays not tied to a specific skill or problem-solving: activity level, neophobia, and social dominance. Zebra finches exposed to dietary mercury throughout their lives were subordinate to, and more active than, control birds. We found no evidence that mercury exposure influenced our metric of neophobia. Together, these results suggest that sub-lethal exposure to environmental mercury selectively harms neurological pathways that control different cognitive abilities, with complex effects on behavior and fitness.

Experimental exposure to urban and pink noise affects brain development and song learning in zebra finches (Taenopygia guttata).

Recently, numerous studies have observed changes in bird vocalizations-especially song-in urban habitats. These changes are often interpreted as adaptive, since they increase the active space of the signal in its environment. However, the proximate mechanisms driving cross-generational changes in song are still unknown. We performed a captive experiment to identify whether noise experienced during development affects song learning and the development of song-control brain regions. Zebra finches (Taeniopygia guttata) were bred while exposed, or not exposed, to recorded traffic urban noise (Study 1) or pink noise (Study 2). We recorded the songs of male offspring and compared these to fathers' songs. We also measured baseline corticosterone and measured the size of song-control brain regions when the males reached adulthood (Study 1 only). While male zebra finches tended to copy syllables accurately from tutors regardless of noise environment, syntax (the ordering of syllables within songs) was incorrectly copied affected by juveniles exposed to noise. Noise did not affect baseline corticosterone, but did affect the size of brain regions associated with song learning: these regions were smaller in males that had been had been exposed to recorded traffic urban noise in early development. These findings provide a possible mechanism by which noise affects behaviour, leading to potential population differences between wild animals occupying noisier urban environments compared with those in quieter habitats.

Blood Mercury Levels of Zebra Finches Are Heritable: Implications for the Evolution of Mercury Resistance.

Mercury is a ubiquitous metal contaminant that negatively impacts reproduction of wildlife and has many other sub-lethal effects. Songbirds are sensitive bioindicators of mercury toxicity and may suffer population declines as a result of mercury pollution. Current predictions of mercury accumulation and biomagnification often overlook possible genetic variation in mercury uptake and elimination within species and the potential for evolution in affected populations. We conducted a study of dietary mercury exposure in a model songbird species, maintaining a breeding population of zebra finches (Taeniopygia guttata) on standardized diets ranging from 0.0-2.4 µg/g methylmercury. We applied a quantitative genetics approach to examine patterns of variation and heritability of mercury accumulation within dietary treatments using a method of mixed effects modeling known as the 'animal model'. Significant variation in blood mercury accumulation existed within each treatment for birds exposed at the same dietary level; moreover, this variation was highly repeatable for individuals. We observed substantial genetic variation in blood mercury accumulation for birds exposed at intermediate dietary concentrations. Taken together, this is evidence that genetic variation for factors affecting blood mercury accumulation could be acted on by selection. If similar heritability for mercury accumulation exists in wild populations, selection could result in genetic differentiation for populations in contaminated locations, with possible consequences for mercury biomagnification in food webs.