Among-population variation in telomere regulatory proteins and their potential role as hidden drivers of intraspecific variation in life history.

Biologists aim to explain patterns of growth, reproduction and ageing that characterize life histories, yet we are just beginning to understand the proximate mechanisms that generate this diversity. Existing research in this area has focused on telomeres but has generally overlooked the telomere's most direct mediator, the shelterin protein complex. Shelterin proteins physically interact with the telomere to shape its shortening and repair. They also regulate metabolism and immune function, suggesting a potential role in life history variation in the wild. However, research on shelterin proteins is uncommon outside of biomolecular work. Intraspecific analyses can play an important role in resolving these unknowns because they reveal subtle variation in life history within and among populations. Here, we assessed ecogeographic variation in shelterin protein abundance across eight populations of tree swallow (Tachycineta bicolor) with previously documented variation in environmental and life history traits. Using the blood gene expression of four shelterin proteins in 12-day-old nestlings, we tested the hypothesis that shelterin protein gene expression varies latitudinally and in relation to both telomere length and life history. Shelterin protein gene expression differed among populations and tracked non-linear variation in latitude: nestlings from mid-latitudes expressed nearly double the shelterin mRNA on average than those at more northern and southern sites. However, telomere length was not significantly related to latitude. We next assessed whether telomere length and shelterin protein gene expression correlate with 12-day-old body mass and wing length, two proxies of nestling growth linked to future fecundity and survival. We found that body mass and wing length correlated more strongly (and significantly) with shelterin protein gene expression than with telomere length. These results highlight telomere regulatory shelterin proteins as potential mediators of life history variation among populations. Together with existing research linking shelterin proteins and life history variation within populations, these ecogeographic patterns underscore the need for continued integration of ecology, evolution and telomere biology, which together will advance understanding of the drivers of life history variation in nature.

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.

The impact of mercury on North American songbirds: effects, trends, and predictive factors.

Researchers were asked to contribute new results addressing questions about the exposure and effects of mercury (Hg) in North American songbirds, a rapidly declining group of species that is the subject of enduring interest for millions of birdwatchers, the general public and conservation scientists. Important questions to be answered include: Is Hg causing or exacerbating songbird population declines? Which North American songbirds are at most risk and in which landscapes? Are there aspects of songbird natural history that pre-dispose them to risks of Hg exposure and effects, in particular, their migratory behavior? In all, 61 authors contributed 15 studies addressing aspects of these questions. Articles in this special issue address an array of topics including: (1) three studies on health effects in the laboratory using a domesticated songbird model species, the zebra finch; (2) three studies on changes in songbird exposure to Hg over time spans from less than a decade to more than a century; (3) five studies on landscape characteristics or management practices that cause the oft-noted spatial variation in Hg accumulation by resident songbirds, from the subarctic tundra to high-elevation tropical forests; (4) three papers examining the recently recognized role of migration behavior in predicting risk to songbirds from Hg; and (5) one paper on the potential pitfalls of using feather Hg concentration as a bioindicator for Hg exposure. In summary, although there are many questions still to be answered, it is clear that the effects of Hg are persistent long after exposure, Hg exposure of North American songbirds is not improving, predicting exposure requires a detailed understanding of ecosystem processes beyond simply the amount of Hg present at a site, migration behavior predisposes songbirds to risk of Hg exposure and effects, and carefully selecting appropriate bioindicator sites, species, and tissues is critical to any monitoring efforts.

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.

Synthesis of Maternal Transfer of Mercury in Birds: Implications for Altered Toxicity Risk.

Maternal transfer is a predominant route of methylmercury (MeHg) exposure to offspring. We reviewed and synthesized published and unpublished data on maternal transfer of MeHg in birds. Using paired samples of females' blood (n = 564) and their eggs (n = 1814) from 26 bird species in 6 taxonomic orders, we conducted a meta-analysis to evaluate whether maternal transfer of MeHg to eggs differed among species and caused differential toxicity risk to embryos. Total mercury (THg) concentrations in eggs increased with maternal blood THg concentrations; however, the proportion of THg transferred from females to their eggs differed among bird taxa and with maternal THg exposure. Specifically, a smaller proportion of maternal THg was transferred to eggs with increasing female THg concentrations. Additionally, the proportion of THg that was transferred to eggs at the same maternal blood THg concentration differed among taxonomic orders, with waterfowl (Anseriformes) transferring up to 382% more THg into their eggs than songbirds (Passeriformes). We provide equations to predict THg concentrations in eggs using female blood THg concentrations, and vice versa, which may help translate toxicity benchmarks across tissues and life stages. Our results indicate that toxicity risk of MeHg can vary among bird taxa due to differences in maternal transfer of MeHg to offspring.

Environmentally relevant methylmercury exposure reduces the metabolic scope of a model songbird.

For most birds, energy efficiency and conservation are paramount to balancing the competing demands of self-maintenance, reproduction, and other demanding life history stages. Yet the ability to maximize energy output for behaviors like predator escape and migration is often also critical. Environmental perturbations that affect energy metabolism may therefore have important consequences for fitness and survival. Methylmercury (MeHg) is a global pollutant that has wide-ranging impacts on physiological systems, but its effects on the metabolism of birds and other vertebrates are poorly understood. We investigated dose-dependent effects of dietary MeHg on the body composition, basal and peak metabolic rates (BMR, PMR), and respiratory quotients (RQ) of zebra finches (Taeniopygia guttata). Dietary exposure levels (0.0, 0.1, or 0.6 ppm wet weight) were intended to reflect a range of mercury concentrations found in invertebrate prey of songbirds in areas contaminated by atmospheric deposition or point-source pollution. We found adiposity increased with MeHg exposure. BMR also increased with exposure while PMR decreased, together resulting in reduced metabolic scope in both MeHg-exposed treatments. There were differences in RQ among treatments that suggested a compromised ability of exposed birds to rapidly metabolize carbohydrates during exercise in a hop-hover wheel. The elevated BMR of exposed birds may have been due to energetic costs of depurating MeHg, whereas the reduced PMR could have been due to reduced oxygen carrying capacity and/or reduced glycolytic capacity. Our results suggest that environmentally relevant mercury exposure is capable of compromising the ability of songbirds to both budget and rapidly exert energy.

Agricultural land use creates evolutionary traps for nesting turtles and is exacerbated by mercury pollution.

The rate of anthropogenic habitat conversion often exceeds the rate of natural ecological and evolutionary processes, which sometimes creates mismatches between environmental cues and adaptive behaviors. In numerous species of aquatic turtles, nest site selection is primarily based on cues related to high solar exposure, which leads females to select sites where humans have disturbed habitat. These disturbed sites are often contaminated by pollutants, such as mercury. Despite the fact that anthropogenic disturbances often co-occur, few studies have examined the interactive influence of major global changes on animal development. Using Chelydra serpentina, we investigated the individual and interactive effects of crop agriculture and mercury pollution on hatch success and offspring phenotype. We hypothesized that following nesting, rapid crop growth would shade and cool nests in agricultural fields and subsequently negatively impact embryonic development. Agricultural and control nests were similar in temperature at the time of oviposition, but temperatures diverged as crops grew: agricultural nests averaged 2.5°C cooler than control nests over the course of incubation. In laboratory and field experiments, we found that turtles incubated under agricultural thermal regimens took longer to hatch, hatched at smaller body sizes, lost more mass, and had lower posthatching growth rates. Additionally, thermal conditions associated with agricultural land use interacted with mercury contamination to decrease hatching success. To our knowledge, this is the first documentation of negative interactive effects of mercury pollution and habitat quality on early vertebrate development and highlights the importance of examining the combined influence of anthropogenic global changes on organisms.

Spatial extent of mercury contamination in birds and their prey on the floodplain of a contaminated river.

Mercury (Hg) exposure has been extensively studied in aquatic and piscivorous wildlife, but, historically, less attention has been directed towards terrestrial species. However, it has become apparent that aquatic Hg crosses ecosystem boundaries along with beneficial subsidies, thereby entering the terrestrial food chain. It is still not known how far from contaminated waterways Hg exposure remains a risk. We examined the spatial extent of exposure in terrestrial songbirds breeding in the floodplain along a 40-km stretch of Hg-contaminated river in Virginia, USA. Four songbird species were chosen that readily occupied artificial nest cavities placed at known distances from the river; Carolina chickadees (Poecile carolinensis), Carolina wrens (Thryothorus ludovicianus), eastern bluebirds (Sialia sialis), and house wrens (Troglodytes aedon). We examined Hg exposure at varying distances from the edge of the river for three endpoints: 1) adult bird blood, 2) nestling bird feather and 3) spider whole body (collected in pitfall traps and directly from the mouths of nestling birds). We used mixed effects models to understand 1) how total Hg (i.e., inorganic and organic, THg) concentrations differed between contaminated and reference sites and 2) how THg concentrations declined with distance away from the river on contaminated sites. Adult bird blood THg indicated that a species-by-distance interaction was significant, after accounting for site and year. Importantly, despite the decline with distance, we found that THg levels in some songbird species and their invertebrate prey remained elevated above reference levels for up to 400m away from the river. Our results predict a spatial extent of contamination that is an order of magnitude greater than similar studies investigating the lateral impact of other aquatically derived contaminants. To our knowledge, this study is the first to indicate that exposure to legacy aquatic Hg is possible for terrestrial wildlife across the entire floodplain, rather than being confined to narrow riparian corridors.

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.

Impacts of Sublethal Mercury Exposure on Birds: A Detailed Review.

Mercury is a ubiquitous environmental contaminant known to accumulate in, and negatively affect, fish-eating and oceanic bird species, and recently demonstrated to impact some terrestrial songbirds to a comparable extent. It can bioaccumulate to concentrations of >1 µg/g in tissues of prey organisms such as fish and insects. At high enough concentrations, exposure to mercury is lethal to birds. However, environmental exposures are usually far below the lethal concentrations established by dosing studies.The objective of this review is to better understand the effects of sublethal exposure to mercury in birds. We restricted our survey of the literature to studies with at least some exposures >5 µg/g. The majority of sublethal effects were subtle and some studies of similar endpoints reached different conclusions. Strong support exists in the literature for the conclusion that mercury exposure reduces reproductive output, compromises immune function, and causes avoidance of high-energy behaviors. For some endpoints, notably certain measures of reproductive success, endocrine and neurological function, and body condition, there is weak or contradictory evidence of adverse effects and further study is required. There was no evidence that environmentally relevant mercury exposure affects longevity, but several of the sublethal effects identified likely do result in fitness reductions that could adversely impact populations. Overall, 72% of field studies and 91% of laboratory studies found evidence of deleterious effects of mercury on some endpoint, and thus we can conclude that mercury is harmful to birds, and the many effects on reproduction indicate that bird population declines may already be resulting from environmental mercury pollution.

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.

Mercury alters initiation and construction of nests by zebra finches, but not incubation or provisioning behaviors.

Mercury is an environmental contaminant that impairs avian reproduction, but the behavioral and physiological mechanisms underlying this effect are poorly understood. The objective of this study was to determine whether lifetime dietary exposure to mercury (1.2 µg/g wet weight in food) impacted avian parental behaviors, and how this might influence reproductive success. To distinguish between the direct effects of mercury on parents and offspring, we created four treatment groups of captive-bred zebra finches (Taeniopygia guttata), with control and mercury-exposed adults raising cross-fostered control or mercury-exposed eggs (from maternal transfer). Control parents were 23% more likely to fledge young than parents exposed to mercury, regardless of egg exposure. Mercury-exposed parents were less likely to initiate nests than controls and spent less time constructing them. Nests of mercury-exposed pairs were lighter, possibly due to an impaired ability to bring nest material into the nestbox. However, nest temperature, incubation behavior, and provisioning rate did not differ between parental treatments. Unexposed control eggs tended to have shorter incubation periods and higher hatching success than mercury-exposed eggs, but there was no effect of parental exposure on these parameters. We accidentally discovered that parent finches transfer some of their body burden of mercury to nestlings during feeding through secretion in the crop. These results suggest that, in mercury-exposed songbirds, pre-laying parental behaviors, combined with direct exposure of embryos to mercury, likely contribute to reduced reproductive success and should be considered in future studies. Further research is warranted in field settings, where parents are exposed to greater environmental challenges and subtle behavioral differences might have more serious consequences than were observed in captivity.

No Effect of Lifelong Methylmercury Exposure on Oxidative Status in Zebra Finches (Taeniopygia guttata): A Demonstration of Methylmercury-Induced Selection?

Songbirds exposed to methylmercury (MeHg) often exhibit reduced reproductive success and cognitive abilities. To better understand whether oxidative stress plays a role, we dosed zebra finches (Taeniopygia guttata) with a contaminated (1.2 ppm MeHg-cysteine) or control diet for their entire lives, including during development in the egg. Levels of antioxidant enzymes [superoxide dismutase (SOD1 and SOD2)], oxidative damage (4-hydroxynonenal; 4-HNE), and antioxidant transcription factors [nuclear factor (erythroid-derived 2)-like 2; Nrf2] were measured in the liver and pectoralis muscle of adults. MeHg treatment did not affect levels of 4-HNE or liver SOD2 or Nrf2. Birds in the MeHg treatment differed significantly from controls in pectoralis SOD1 and Nrf2, and tended to differ in liver SOD1 and pectoralis SOD2; however, we detected no overall pattern of effect of MeHg on oxidative status in dosed finches. We suspect that this is a consequence of the differential survival of MeHg-tolerant birds.

Form of Dietary Methylmercury does not Affect Total Mercury Accumulation in the Tissues of Zebra Finch.

Exposure to mercury in humans, other mammals, and birds is primarily dietary, with mercury in the methylated form and bound to cysteine in the tissues of prey items. Yet dosing studies are generally carried out using methylmercury chloride. Here we tested whether the accumulation of total mercury in zebra finch blood, egg, muscle, liver, kidney or brain differed depending on whether dietary mercury was complexed with chloride or cysteine. We found no effect of form of mercury on tissue accumulation. Some previous studies have found lower accumulation of mercury in tissues of animals fed complexed mercury. Much remains to be understood about what happens to ingested mercury once it enters the intestines, but our results suggest that dietary studies using methylmercury chloride in birds will produce similar tissue accumulation levels to those using methylmercury cysteine.

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.

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.

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.

Mobilization of mercury from lean tissues during simulated migratory fasting in a model songbird.

The pollutant methylmercury accumulates within lean tissues of birds and other animals. Migrating birds catabolize substantial amounts of lean tissue during flight which may mobilize methylmercury and increase circulating levels of this neurotoxin. As a model for a migrating songbird, we fasted zebra finches (Taeniopygia guttata) that had been dosed with 0.0, 0.1, and 0.6 parts per million (ppm) dietary methylmercury and measured changes in blood total mercury concentrations (THg) in relation to reductions in lean mass. Birds lost 6-16% of their lean mass during the fast, and THg increased an average of 12% and 11% in the 0.1 and 0.6 ppm treatments, respectively. Trace amounts of THg in the 0.0 ppm control group also increased as a result of fasting, but remained extremely low. THg increased 0.4 ppm for each gram of lean mass catabolized in the higher dose birds. Our findings indicate that methylmercury is mobilized from lean tissues during protein catabolism and results in acute increases in circulating concentrations. This is a previously undocumented potential threat to wild migratory birds, which may experience greater surges in circulating methylmercury than demonstrated here as a result of their greater reductions in lean mass.

Tissue Mercury Concentrations and Survival of Tree Swallow Embryos, Nestlings and Young Adult Females on a Contaminated Site.

Tree swallows nesting on mercury-contaminated sites along the South River in Virginia, USA were monitored for reproductive success. The bodies of nestlings found deceased in their nest boxes were collected, along with blood and feather samples from the adult parents and surviving siblings. We also measured hatching and fledging success of the clutches and the annual recapture rate of adults. We found that the body feathers of deceased nestlings contained significantly higher concentrations of mercury (12.89 ± 8.42 µg/g, n = 15) than those of nestlings that survived to fledge (7.41 ± 4.79 µg/g, n = 15). However, mothers of more successful clutches (>75 % hatching) did not differ in mercury concentrations from females with less successful clutches (<50 % hatching). Additionally, adult females breeding for the first time that returned to breed the following year did not differ in blood mercury from females of the same age that bred once but never returned. Our results suggest that mercury had its greatest effect on these songbirds during the nestling stage, whereas for embryos or first-time breeding females, other factors likely played larger roles in mortality.