House Sparrows Passer Domesticus Research Articles

Conspecific alarm calls prevent the attenuation of neophobia behavior in wild-caught house sparrows (Passer domesticus)

Some individuals respond to new objects, foods, or environments with wariness (neophobia), whereas others are willing to approach and explore. Because novel stimuli can represent both dangers and resources, group-living species may show adaptive plasticity in neophobia in response to social cues. To better understand how conspecific calls can influence neophobia in a highly gregarious species, we exposed individual house sparrows (Passer domesticus) to either conspecific alarm calls (n = 12), conspecific contact calls (n = 12), or no playback (n = 12) and measured latency to feed in the presence of novel objects. We also measured novelty responses with no sound the week before and after the sound treatment week for all individuals. Relative to no playback and contact calls, we predicted that conspecific alarm calls would increase neophobia behavior during the acoustic trial and that these effects would persist the week after exposure. Instead, we found that individuals in the contact call and no playback groups became less neophobic as weeks progressed, while the alarm call group showed no attenuation of neophobia. There was a significant interaction between week and treatment, where neophobia responses over the three weeks were significantly different for individuals exposed to alarm calls compared to the contact and no playback groups combined. These results suggest that house sparrows learn social information about potentially threatening stimuli from conspecific alarm calls; here, that novel objects may be dangerous.

Contrasting morphometric responses to increasing urbanisation in congeneric sparrow species

Increased urbanisation influences the morphometric traits of various species, often resulting in urban individuals being smaller than their non-urban counterparts. Urbanisation can affect fundamental eco-evolutionary patterns and impact species’ ability to adapt to and occupy rapidly changing environments through morphological changes. We investigated the morphometric responses of two passerine species, the non-native house sparrow (Passer domesticus) and its native congener, the Cape sparrow (Passer melanurus), along gradients of spatial and temporal urbanisation in South Africa over a 52-year period. The house sparrow was significantly heavier, larger and in better condition with increasing urban infrastructure and lower urban vegetation cover, while the Cape sparrow showed opposing trends along these gradients. Temporally, the house sparrow’s body mass increased consistently over the 52-year study period, suggesting changes in morphology were concomitant with increasing urbanisation over time. This study demonstrates distinct differences in the morphological responses of the non-native house sparrow and the native Cape sparrow to increasing urban development. These morphological responses may also underpin community-level changes caused by urbanisation, enhancing the capabilities of non-native species to thrive over their native counterparts in these environments.

Oxidative stress across multiple tissues in house sparrows (Passer domesticus) acclimated to warm, stable cold, and unpredictable cold thermal treatments.

With climate change increasing not just mean temperatures but the frequency of cold snaps and heat waves, animals occupying thermally variable areas may be faced with thermal conditions for which they are not prepared. Studies of physiological adaptations of temperate resident birds to such thermal variability are largely lacking in the literature. To address this gap, we acclimated winter-phenotype house sparrows (Passer domesticus) to stable warm, stable cold, and fluctuating cold temperatures. We then measured several metrics of the oxidative stress (OS) system, including enzymatic and non-enzymatic antioxidants and lipid oxidative damage, in brain (post-mitotic), kidney (mitotic), liver (mitotic) and pectoralis muscle (post-mitotic). We predicted that high metabolic flexibility could be linked to increases in reactive oxygen damage. Alternatively, if variation in ROS production is not associated with metabolic flexibility, then we predict no antioxidant compensation with thermal variation. Our data suggest that ROS production is not associated with metabolic flexibility, as we found no differences across thermal treatment groups. However, we did find differences across tissues. Brain catalase activity demonstrated the lowest values compared with kidney, liver and muscle. In contrast, brain glutathione peroxidase (GPx) activities were higher than those in kidney and liver. Muscle GPx activities were intermediate to brain and kidney/liver. Lipid peroxidation damage was lowest in the kidney and highest in muscle tissue.

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.

Long-distance dispersal in the short-distance dispersing house sparrow (Passer domesticus).

The house sparrow (Passer domesticus) is a small passerine known to be highly sedentary. Throughout a 30-year capture-mark-recapture study, we have obtained occasional reports of recoveries far outside our main metapopulation study system, documenting unusually long dispersal distances. Our records constitute the highest occurrence of long-distance dispersal events recorded for this species in Scandinavia. Such long-distance dispersals radically change the predicted distribution of dispersal distances and connectedness for our study metapopulation. Moreover, it reveals a much greater potential for colonization than formerly recorded for the house sparrow, which is an invasive species across four continents. These rare and occasional long-distance dispersal events are challenging to document but may have important implications for the genetic composition of small and isolated populations and for our understanding of dispersal ecology and evolution.

Light pollution affects activity differentially across breeding stages in an urban exploiter: An experiment in the house sparrow (Passer domesticus)

Artificial Light At Night (ALAN) is a major urban perturbation, which can have detrimental effects on wildlife. Recent urban planning has led to an increased use of white light emission diodes (LEDs) in cities. However, little is known about the effects of this type of ALAN on wild vertebrates, especially during reproduction. We designed an experiment to test the impact of ALAN on the activity rhythms (daily time of first activity (TFA) and time of last activity (TLA)) of captive House sparrows (Passer domesticus) during several reproductive stages (from pre-breeding to post-breeding). We also tested the impact of ALAN on reproductive performance (laying date, clutch size, hatching and fledging success). Experimental birds were active earlier in the morning (earlier TFA) relative to controls although experimental and control birds did not differ in their TLA. The effect of ALAN on TFA was apparent during specific stages only (pre-breeding and chick-rearing stages), suggesting that sparrows actively adjust their activity in response to ALAN only during specific periods. This impact of ALAN on activity did not persist through the whole breeding season, suggesting that sparrows may habituate to ALAN. Alternatively, they may not be able to sustain a long-term increased activity in response to ALAN because of sleep deprivation and related physiological costs. Finally, we did not find any impact of ALAN on the reproductive performance of captive house sparrows held under optimal conditions. This suggests that ALAN may not be dramatically detrimental to the reproduction of this urban exploiter, at least when food availability is not constraining.

Avifaunal Diversity in Telangana State: A Case Study in Campus of Professor Jayashankar Telangana State Agricultural University, Rajendranagar, Hyderabad, India

Population Studies on different agricultural and other bird species were carried out at Students farm, College farm, Agri biodiversity park, and Agricultural Research Institute farm areas of Professor Jayashankar Telangana State Agricultural University (PJTSAU) Campus in semiarid region of Telangana state covering an area of about 36 Sq.km. for a period of eleven years (2012 to 2023). Population diversity of 92 species of different birds, and their distribution within the university campus farm areas were recorded. Important among them are 9 species namely: Blue rock pigeon (Columba livia), Spotted dove (Streptopelia chinensis), Rose ringed parakeet (Psittacula krameri),Common myna (Acridotheres tristis), House crow (Corvus splendens), House sparrow (Passer domesticus), Baya weaver bird (Ploceus philippinus), White throated munia (Lonchura malabarica), and Peafowl (Pavo cristatus) which depredate experimental field crops among them only few birds such as: Rose ringed parakeets, House crows, House sparrows, Pigeons, Baya weavers, Peafowls and Munias were serious bird pests to major agricultural and horticultural crops. The University farm areas were found to be the most preferred habitat for many of these birds, it could be due to availability of food, breeding, roosting and resting sites in abundance because of more agricultural croplands and agro forestry trees.

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.

Urbanization alters the song propagation of two human-commensal songbird species.

Urban expansion has increased pollution, including both physical (e.g., exhaust, litter) and sensory (e.g., anthropogenic noise) components. Urban avian species tend to increase the frequency and/or amplitude of songs to reduce masking by low-frequency noise. Nevertheless, song propagation to the receiver can also be constrained by the environment. We know relatively little about how this propagation may be altered across species that (1) vary in song complexity and (2) inhabit areas along an urbanization gradient. We investigated differences in song amplitude, attenuation, and active space, or the maximum distance a receiver can detect a signal, in two human-commensal species: the house sparrow (Passer domesticus) and house finch (Haemorhous mexicanus). We described urbanization both discretely and quantitatively to investigate the habitat characteristics most responsible for propagation changes. We found mixed support for our hypothesis of urban-specific degradation of songs. Urban songs propagated with higher amplitude; however, urban song fidelity was species-specific and showed lowered active space for urban house finch songs. Taken together, our results suggest that urban environments may constrain the propagation of vocal signals in species-specific manners. Ultimately, this has implications for the ability of urban birds to communicate with potential mates or kin.

Roosting and nesting ecology of house sparrow (\(\textit{Passer domesticus}\)) in Guwahati city, Assam

Aves are unique in having a coat of feathers and in balancing on the hind limbs alone. They are known as the feathered bipeds. Roosting, nest building in birds has been considered a remarkable behaviour in aves. The nesting ecology of birds is likely the most important component of its life history and it is affected by several ecological and environmental factors. Guwahati city, a rapidly growing city in northeast India inhabits a lot of species of birds. It is located between the Brahmaputra River’s southern bank and the Shillong plateau’s foothills. Total of the 57 nests, 28% were discovered in the rolling shop shutters, followed by 27% in the tin sheet roofs of homes, followed by 19% in building crevices, 9% in building holes, 7% in advertisement hoarding and 5% in AC outdoor units and 5% in unused building pipes. It is also observed that 49% of the nests are located at a height between 2–4 metres from the ground level, 70% of the nests are located between 1–5 metres from the nearest food source, and 58% of the nests are located between 1–5 metres from the nearest water source. Out of the 57 nests observed, 50 were found to be solitary, 5 were found to be Pair, and 2 were found to be cluster type. The study found that choosing a nest is influenced directly by the location of the nearest food source and nearest water source in Guwahati city. The study presents the merit and needs for studying the nesting ecology of birds in India, also suggests best practices to conserve bird’s nest specific to the Indian context which might provide important information for the nesting ecology of House Sparrows (Passer domesticus).

Immune gene expression and epigenetic potential affect the consumption of risky food by female house sparrows

When organisms move into new areas, they are likely to encounter novel food resources. Even if they are nutritious, these foods can also be risky, as they might be contaminated by parasites. The behavioural immune system of animals could help them avoid the negative effects of contaminated resources, but our understanding of behavioural immunity is limited, particularly whether and how behavioural immunity interacts with physiological immunity. Here, we asked about the potential for interplay between these two traits, specifically how the propensity of an individual house sparrow (Passer domesticus) to take foraging risks was related to its ability to regulate a key facet of its immune response to bacterial pathogens. Previously, we found that sparrows at expanding geographic range edges were more exploratory and less risk-averse to novel foods; in those same populations, birds tended to over-express Toll-like receptor 4 (TLR4), a pattern-recognition receptor that distinguishes cell-wall components of Gram-negative bacteria, making it the major sensor of potentially lethal gut microbial infections including salmonellosis. When we investigated how birds would respond to a typical diet (i.e., mixed seeds) spiked with domesticated chicken faeces, birds that expressed more TLR4 or had higher epigenetic potential for TLR4 (more CpG dinucleotides in the putative gene promoter) ate more food, spiked or not. Females expressing abundant TLR4 were also willing to take more foraging risks and ate more spiked food. In males, TLR4 expression was not associated with risk-taking. Altogether, our results indicate that behaviour and immunity covary among individual house sparrows, particularly in females where those birds that maintain more immune surveillance also are more disposed to take foraging risks.

Avian architects: Technology, domestication, and animal minds in urban America.

In the mid-nineteenth century, the house sparrow (Passer domesticus) was introduced to the United States, quickly spreading across the country. For a brief period in the late nineteenth and early twentieth centuries, the observation of sparrow behavior was something of an urban pastime. Traits such as intelligence, reason, persistence, and craftsmanship were conferred onto sparrows by American urbanites. This paper argues that sparrow intelligence was often conflated with domestication: the ability of the birds to adapt to living alongside humans. Praise for the ingenuity of sparrows generally revolved around their nest building, particularly when such structures overcame the challenges posed by urban infrastructure and technology. Sparrows were far less praiseworthy when they caused electricity outages or contaminated water supplies. The sparrow in the United States demonstrates how the relationship between these anecdotes and their implications for animal minds was mediated by the technology and infrastructure of cities. Admirers of sparrows were not measuring the birds' mental capacity, but rather their ability to adapt to human habitations. Sparrows were only granted intelligence once they had demonstrated their ability to become domesticated.

Nestling size and ornamentation interact to shape early development in house sparrow families

Abstract In many nuclear families, dependent offspring receive unequal shares of parental investment. Initial overproduction can be adaptive from the perspective of parents, but parents must be able to identify appropriate candidates for favourite status. We studied early nestling development in house sparrows (Passer domesticus), a species in which the loss of some brood members is common, testing the prediction that body mass and carotenoid-rich flange colours are important to nestling success. There was substantial variation in both traits within broods, even only 1 day after hatching. Nestlings low in the within-brood mass hierarchy gained more mass if they displayed more carotenoid-rich flanges than broodmates. Position in the colour hierarchy did not, however, predict mass gain for individuals that were heavier than their broodmates. Nestlings that were heavier or had more carotenoid-rich mouths were also less likely to be the victim of brood reduction. Our results suggest that house sparrow parents use both nestling body size and mouth colour when making allocation decisions. Understanding both how and when offspring traits and parental preferences function is key to understanding how selective pressures act on offspring–parent communication.

Water restriction increases oxidation of endogenous amino acids in house sparrows (Passer domesticus).

Animals can cope with dehydration in a myriad of ways, both behaviorally and physiologically. The oxidation of protein produces more metabolic water per kilojoule than that of fat or carbohydrate, and it is well established that birds increase protein catabolism in response to high rates of water loss. However, the fate of amino acids mobilized in response to water restriction has not been explicitly determined. While protein catabolism releases bound water, we hypothesized that water-restricted birds would also oxidize the resulting amino acids, producing additional water as a product of oxidative phosphorylation. To test this, we fed captive house sparrows (Passer domesticus) 13C-labeled leucine for 9 weeks to label endogenous proteins. We conducted weekly trials during which we measured the physiological response to water restriction as changes in lean mass, fat mass, metabolism and the enrichment of 13C in exhaled CO2 (δ13Cbreath). If water-restricted birds catabolized proteins and oxidized the resulting amino acids, we expected to simultaneously observe greater lean mass loss and elevated δ13Cbreath relative to control birds. We found that water-restricted birds catabolized more lean tissue and also had enriched δ13Cbreath in response to water restriction, supporting our hypothesis. δ13Cbreath, however, varied with metabolic rate and the length of the water restriction period, suggesting that birds may spare protein when water balance can be achieved using other physiological strategies.

Calling Structural Variants with Confidence from Short-Read Data in Wild Bird Populations.

Comprehensive characterization of structural variation in natural populations has only become feasible in the last decade. To investigate the population genomic nature of structural variation, reproducible and high-confidence structural variation callsets are first required. We created a population-scale reference of the genome-wide landscape of structural variation across 33 Nordic house sparrows (Passer domesticus). To produce a consensus callset across all samples using short-read data, we compare heuristic-based quality filtering and visual curation (Samplot/PlotCritic and Samplot-ML) approaches. We demonstrate that curation of structural variants is important for reducing putative false positives and that the time invested in this step outweighs the potential costs of analyzing short-read-discovered structural variation data sets that include many potential false positives. We find that even a lenient manual curation strategy (e.g. applied by a single curator) can reduce the proportion of putative false positives by up to 80%, thus enriching the proportion of high-confidence variants. Crucially, in applying a lenient manual curation strategy with a single curator, nearly all (>99%) variants rejected as putative false positives were also classified as such by a more stringent curation strategy using three additional curators. Furthermore, variants rejected by manual curation failed to reflect the expected population structure from SNPs, whereas variants passing curation did. Combining heuristic-based quality filtering with rapid manual curation of structural variants in short-read data can therefore become a time- and cost-effective first step for functional and population genomic studies requiring high-confidence structural variation callsets.

ECR Spotlight – Elizabeth Rogers

ECR Spotlight is a series of interviews with early-career authors from a selection of papers published in Journal of Experimental Biology and aims to promote not only the diversity of early-career researchers (ECRs) working in experimental biology but also the huge variety of animals and physiological systems that are essential for the ‘comparative’ approach. Elizabeth (Beth) Rogers is an author on ‘ Water restriction increases oxidation of endogenous amino acids in house sparrows (Passer domesticus)’, published in JEB. Beth is a PhD candidate in the lab of Alex Gerson at University of Massachusetts Amherst, USA, investigating the physiological and metabolic adaptations that allow animals to contend with ecological challenges such as long-distance migration, disease, environmental change and the confluence of all three.

Only rare classical MHC-I alleles are highly expressed in the European house sparrow.

The exceptional polymorphism observed within genes of the major histocompatibility complex (MHC), a core component of the vertebrate immune system, has long fascinated biologists. The highly polymorphic classical MHC class-I (MHC-I) genes are maintained by pathogen-mediated balancing selection (PMBS), as shown by many sites subject to positive selection, while the more monomorphic non-classical MHC-I genes show signatures of purifying selection. In line with PMBS, at any point in time, rare classical MHC alleles are more likely than common classical MHC alleles to confer a selective advantage in host-pathogen interactions. Combining genomic and expression data from the blood of wild house sparrows Passer domesticus, we found that only rare classical MHC-I alleles were highly expressed, while common classical MHC-I alleles were lowly expressed or not expressed. Moreover, highly expressed rare classical MHC-I alleles had more positively selected sites, indicating exposure to stronger PMBS, compared with lowly expressed classical alleles. As predicted, the level of expression was unrelated to allele frequency in the monomorphic non-classical MHC-I alleles. Going beyond previous studies, we offer a fine-scale view of selection on classical MHC-I genes in a wild population by revealing differences in the strength of PMBS according to allele frequency and expression level.

Seasonal changes in bird communities on poultry farms and house sparrow-wild bird contacts revealed by camera trapping.

Wild birds are considered reservoirs of poultry pathogens although transmission routes have not been conclusively established. Here we use camera trapping to study wild bird communities on commercial layer and red-legged partridge farms over a one-year timeframe. We also analyze direct and indirect interactions of other bird species with the house sparrow (Passer domesticus), a potential bridge host. We conducted camera trapping events between January 2018 and October 2019, in two caged layer farms, one free-range layer farm, and two red-legged partridge farms in South-Central Spain. We observed wild bird visits on all types of farms, with the significantly highest occurrence on red-legged partridge farms where food and water are more easily accessible, followed by commercial caged layer farms, and free-range chicken farms. The house sparrow (Passer domesticus) followed by spotless starlings (Sturnus unicolor) was the most encountered species on all farms, with the highest frequency in caged layer farms. On partridge farms, the house sparrow accounted for 58% of the wild bird detections, while on the free-range chicken farm, it made up 11% of the detections. Notably, the breeding season, when food and water are scarce in Mediterranean climates, saw the highest number of wild bird visits to the farms. Our findings confirm that the house sparrow, is in direct and indirect contact with layers and red-legged partridges and other wild birds independent of the type of farm. Contacts between house sparrows and other bird species were most frequent during the breeding season followed by the spring migration period. The species most frequently involved in interactions with the house sparrow belonged to the order Passeriformes. The study provides a comparative description of the composition and seasonal variations of bird communities in different types of layer/ poultry farms in Southern Spain i.e. a Mediterranean climate. It confirms the effectiveness of biosecurity measures that restrict access to feed and water. Additionally, it underscores the importance of synanthropic species, particularly the house sparrow, as potential bridge vector of avian pathogens.

The impact of parental and developmental stress on DNA methylation in the avian hypothalamic-pituitary-adrenal axis.

The hypothalamic-pituitary-adrenal (HPA) axis coordinates an organism's response to environmental stress. The responsiveness and sensitivity of an offspring's stress response may be shaped not only by stressors encountered in their early post-natal environment but also by stressors in their parent's environment. Yet, few studies have considered how stressors encountered in both of these early life environments may function together to impact the developing HPA axis. Here, we manipulated stressors in the parental and post-natal environments in a population of house sparrows (Passer domesticus) to assess their impact on changes in DNA methylation (and corresponding gene expression) in a suite of genes within the HPA axis. We found that nestlings that experienced early life stress across both life-history periods had higher DNA methylation in a critical HPA axis gene, the glucocorticoid receptor (NR3C1). In addition, we found that the life-history stage when stress was encountered impacted some genes (HSD11B1, NR3C1 and NR3C2) differently. We also found evidence for the mitigation of parental stress by post-natal stress (in HSD11B1 and NR3C2). Finally, by assessing DNA methylation in both the brain and blood, we were able to evaluate cross-tissue patterns. While some differentially methylated regions were tissue-specific, we found cross-tissue changes in NR3C2 and NR3C1, suggesting that blood is a suitable tissue for assessing DNA methylation as a biomarker of early life stress. Our results provide a crucial first step in understanding the mechanisms by which early life stress in different life-history periods contributes to changes in the epigenome of the HPA axis.

Lead exposure is related to higher infection rate with the gapeworm in Norwegian house sparrows (Passer domesticus)

Anthropogenic pollution is identified as an important threat to bird and other wildlife populations. Many metals and toxic elements, along with poly- and perfluoroalkyl substances (PFASs) are known to induce immunomodulation and have previously been linked to increased pathogen prevalence and infectious disease severity. In this study, the house sparrow (Passer domesticus) was investigated at the coast of Helgeland in northern Norway. This population is commonly infected with the parasitic nematode “gapeworm” (Syngamus trachea), with a prevalence of 40–60 % during summer months. Gapeworm induces severe respiratory disease in birds and has been previously demonstrated to decrease survival and reproductive success in wild house sparrows. The aim of this study was to investigate whether a higher exposure to pollution with PFASs, metals and other elements influences gapeworm infection in wild house sparrows. We conducted PFASs and elemental analysis on whole blood from 52 house sparrows from Helgeland, including analyses of highly toxic metals such as lead (Pb), mercury (Hg) and arsenic (As). In addition, we studied gapeworm infection load by counting the parasite eggs in faeces from each individual. We also studied the expression of microRNA 155 (miR155) as a key regulator in the immune system. Elevated blood concentrations of Pb were found to be associated with an increased prevalence of gapeworm infection in the house sparrow. The expression of miR155 in the plasma of the house sparrow was only weakly associated with Pb. In contrast, we found relatively low PFASs concentrations in the house sparrow blood (∑ PFASs 0.00048–354 μg/L) and PFASs were not associated to miR155 nor infection rate. The current study highlights the potential threat posed by Pb as an immunotoxic pollutant in small songbirds.