Rare edges and abundant cores: range-wide variation in abundance in North American birds.

Understanding how the abundance of species varies across geographical ranges is central to ecology; however, few studies test hypotheses using detailed abundance estimates across the full ranges of species on a continental scale. Here, we use unprecedented, detailed estimates of breeding abundance for North American birds (eBird) to test two hypotheses for how abundance varies across species' ranges. We find widespread support for the rare-edge hypothesis-where the abundance of species declines near the range edge-reflecting both reduced occurrence and lower local abundance near range edges. By contrast, we find mixed support for the abundant-centre hypothesis-where the abundance of species peaks in the centre of the range and declines towards the edges-with limited support in conservative tests within species, but general support in among-species tests that control for unbalanced sampling and consider a broader definition of the range centre. Overall, results are consistent with a gradual decline in suitable conditions and increase in challenge towards the range edge that eventually limit the ability of populations to persist.

Urban tolerance does not protect against population decline in North American birds.

Population declines of organisms are widespread and severe, but some species' populations have remained stable, or even increased. The reasons some species are less vulnerable to population decline than others are not well understood. Species that tolerate urban environments often have a broader environmental tolerance, which, along with their ability to tolerate one of the most human-modified habitats (i.e. cities), might allow them to persist in the face of diverse anthropogenic challenges. Here, we examined the relationship between urban tolerance and annual population trajectories for 397 North American bird species. Surprisingly, we found that urban tolerance was unrelated to species' population trajectories. The lack of a relationship between urban tolerance and population trajectories may reflect other factors driving population declines independent of urban tolerance, challenges that are amplified in cities (e.g. climate warming, disease), and other human impacts (e.g. conservation efforts, broad-scale land-use changes) that have benefitted some urban-avoidant species. Overall, our results illustrate that urban tolerance does not protect species against population decline.

Future directions in urban endocrinology - The effects of endocrine plasticity on urban tolerance.

After twenty years of studies of endocrine traits in animals living in cities, the field of urban endocrinology has built a robust literature including numerous studies looking for signatures of the effects of urban living, usually in mean circulating hormone concentrations. The findings of this past research have primarily demonstrated the absence of any generalizable endocrine responses to city life. In this opinion paper, I suggest that a strong route forward would include investigations of the role of variation in endocrine plasticity in determining the degree to which organisms tolerate urban challenges (i.e., urban tolerance). Achieving this research aim will require creative experimental and comparative studies, consideration of alternative study systems, and teasing apart of sources of variation in plastic phenotypes (plasticity, sorting, and contemporary evolution). Insight into the role of endocrine plasticity in influencing urban tolerance could help us better understand and predict impacts of expanding urbanization on biodiversity across the globe.

Among-species variation in hormone concentrations is associated with urban tolerance in birds.

As cities expand across the globe, understanding factors that underlie variation in urban tolerance is vital for predicting changes in patterns of biodiversity. Endocrine traits, like circulating hormone concentrations and regulation of endocrine responses, might contribute to variation in species' ability to cope with urban challenges. For example, variation in glucocorticoid and androgen concentrations has been linked to life-history and behavioural traits that are associated with urban tolerance. However, we lack an understanding of the degree to which evolved differences in endocrine traits predict variation in urban tolerance across species. We analysed 1391 estimates of circulating baseline corticosterone, stress-induced corticosterone, and testosterone concentrations paired with citizen-science-derived urban occurrence scores in a broad comparative analysis of endocrine phenotypes across 71 bird species that differ in their occurrence in urban habitats. Our results reveal context-dependent links between baseline corticosterone and urban tolerance, as well as testosterone and urban tolerance. Stress-induced corticosterone was not related to urban tolerance. These findings suggest that some endocrine phenotypes contribute to a species' tolerance of urban habitats, but also indicate that other aspects of the endocrine phenotype, such as the ability to appropriately attenuate responses to urban challenges, might be important for success in cities.

Experimental reduction of haemosporidian infection affects maternal reproductive investment, parental behaviour and offspring condition.

When hosts have a long coevolutionary history with their parasites, fitness costs of chronic infection have often been assumed to be negligible. Yet, experimental manipulation of infections sometimes reveals effects of parasites on their hosts, particularly during reproduction. Whether these effects translate into fitness costs remains unclear. Here, we present the results of an experimental study conducted in a free-ranging population of red-winged blackbirds (Agelaius phoeniceus) naturally experiencing a high prevalence of haemosporidian infections, with more than 95% of breeding adults infected with parasites from one or more haemosporidian genus. To assess effects of infection during reproduction, we manipulated adult red-winged blackbird females' parasite burden by administering an anti-haemosporidian medication before onset of egg-laying. Experimental reduction of infection resulted in significant benefits to mothers and their offspring. Medicated females laid heavier clutches, invested more in incubation and provisioning behaviour, and produced more fledglings than control females. Nestlings of medicated females had higher haematocrit, higher blood glucose, and lower reactive oxygen metabolites than nestlings of control females. Overall, our results provide evidence that, even in a species with high prevalence of infection, parasites can lead to decreased maternal investment and offspring quality, substantially reducing fitness.

Estimates of species-level tolerance of urban habitat in North American birds.

Species vary in their responses to urban habitat; most species avoid these environments, whereas others tolerate or even thrive in them. To better characterize the extent to which species vary in their responses to urban habitat (from this point forwards "urban tolerance"), we used several methods to quantify these responses at a continental scale across all birds. Using open access community science-derived data from the eBird Status and Trends Products and two different types of high-resolution geospatial data that quantify urbanization of landscapes, we calculated urban tolerance for 432 species with breeding ranges that overlap large cities in Canada or the USA. We developed six different calculations to characterize species-level urban tolerance, allowing us to assess how each species' relative abundance across their breeding range varied with estimates of urban habitat use and intensity. We assessed correlations among these six indices, then compressed the two best-performing indices into a single principal component (multivariate urban tolerance index) that captured variation in urban tolerance among species. We assessed the accuracy of our single and multivariate urban tolerance indices using 24 test species that have been well characterized for their tolerance or avoidance of the urban habitat, as well as with previously published, independent urban tolerance estimates. Here, we provide this new dataset of species-level urban tolerance estimates that improves upon previous metrics by incorporating continental-scale, continuous estimates that better differentiate species' tolerance of urban habitat compared with existing, categorical methods. These refined metrics can be used to test hypotheses that link ecological, life history, and behavioral traits to avian urban tolerance. The dataset is licensed as CC-By Attribution 4.0 International. Users must appropriately cite the data paper and dataset if used in publications and scientific presentations.

Cytonuclear discordance in the crowned-sparrows, Zonotrichia atricapilla and Zonotrichia leucophrys.

The golden-crowned (Zonotrichia atricapilla) and white-crowned (Z. leucophrys) sparrows have been presented as a compelling case for rapid speciation. They display divergence in song and plumage with overlap in their breeding ranges implying reproductive isolation, but have almost identical mitochondrial genomes. Previous research proposed hybridization and subsequent mitochondrial introgression as an alternate explanation, but lacked robust nuclear gene trees to distinguish between introgression and incomplete lineage sorting. We test for signatures of these processes between Z. atricapilla and Z. leucophrys, and investigate the relationships among Z. leucophrys subspecies, using mitochondrial sequencing and a reduced representation nuclear genomic dataset. Contrary to the paraphyly evident in mitochondrial gene trees, we confirmed the reciprocal monophyly of Z. atricapilla and Z. leucophrys using large panels of single nucleotide polymorphisms (SNPs). The pattern of cytonuclear discordance is consistent with limited, historical hybridization and mitochondrial introgression, rather than a recent origin and incomplete lineage sorting between recent sister species. We found evidence of nuclear phylogeographic structure within Z. leucophrys with two distinct clades. Altogether, our results indicate deeper divergences between Z. atricapilla and Z. leucophrys than inferred using mitochondrial markers. Our results demonstrate the limitations of relying solely on mitochondrial DNA for taxonomy, and raise questions about the possibility of selection on the mitochondrial genome during temperature oscillations (e.g. during the Pleistocene). Historical mitochondrial introgression facilitated by past environmental changes could cause erroneous dating of lineage splitting in other taxa when based on mitochondrial DNA alone.

Plasticity versus Evolutionary Divergence: What Causes Habitat Partitioning in Urban-Adapted Birds?

AbstractHabitat partitioning can facilitate the coexistence of closely related species and often results from competitive interference inducing plastic shifts of subordinate species in response to aggressive, dominant species (plasticity) or the evolution of ecological differences in subordinate species that reduce their ability to occupy habitats where the dominant species occurs (evolutionary divergence). Evidence consistent with both plasticity and evolutionary divergence exist, but the relative contributions of each to habitat partitioning have been difficult to discern. Here we use a global data set on the breeding occurrence of birds in cities to test predictions of these alternative hypotheses to explain previously described habitat partitioning associated with competitive interference. Consistent with plasticity, the presence of behaviorally dominant congeners in a city was associated with a 65% reduction in the occurrence of subordinate species, but only when the dominant was a widespread breeder in urban habitats. Consistent with evolutionary divergence, increased range-wide overlap with dominant congeners was associated with a 56% reduction in the occurrence of subordinates in cities, even when the dominant was absent from the city. Overall, our results suggest that both plasticity and evolutionary divergence play important, concurrent roles in habitat partitioning among closely related species in urban environments.

A comparison of sex, morphology, physiology and behavior of black-capped chickadees trapped using two common capture methods.

Many biological studies require the capture of individuals for sampling, for example for measurement of morphological or physiological traits, or for marking individuals for later observations. Capture methods employed often vary both within and between studies, and these differing methods could be more or less effective in capture of different individuals based on their morphology or behavior. If individuals that are prone to capture by the selected method differ with respect to traits of interest, such sampling bias could generate misleading or simply inaccurate results. The selection of capture methods could introduce two different forms of sampling bias, with the individuals that are sampled differing from the population at large or with individuals sampled via one method differing from individuals that could be sampled using a different method. We investigated this latter form of sampling bias by comparing individual birds sampled using two common capture techniques. We caught free-ranging black-capped chickadees (Poecile atricapillus) using walk-in traps baited with seed and mist nets paired with playback of an audio stimulus (conspecific mobbing calls). We measured 18 traits that we expect might vary among birds that are trappable by these differing methods-one that targets birds that are food motivated and potentially less neophobic and another that targets birds that respond readily to a perceived predation risk. We found no differences in the sex, morphology, initial and stress-induced corticosterone concentrations, behavioral response to a novel object, or behavioral response to a predator between individuals captured by these two methods. Individual variation in the behavioral response to a novel object was greater among birds caught by mist nets, suggesting this method might provide a sample that better reflects population-level individual variation. We do not know if the birds caught by these two methods provide a representative sample of the population at large, but can conclude that selection of either of these two common capture methods can similarly sample mean trait values of a population of interest. To accurately assess individual variation, particularly in behavior, mist nets might be preferable.

Population decline in tree swallows (Tachycineta bicolor) linked to climate change and inclement weather on the breeding ground.

Population decline and the threat of extinction are realities currently facing many species. Yet, in most cases, the detailed demographic data necessary to identify causes of population decline are unavailable. Using 43 years (1975-2017) of data from a box-nesting population of tree swallows (Tachycineta bicolor), we identified reduced survival of offspring as a probable demographic cause of population decline. Poor fledging success was associated with increased predation and poor weather conditions during early nestling development. Low juvenile survival and subsequent recruitment was linked to poor weather conditions during the post-fledging period and may also be linked to conditions on the wintering grounds. Regional weather conditions during critical stages of breeding (early nestling and post-fledging) have become progressively worse over the 43-year study period. None of the other factors linked to offspring survival have similarly deteriorated. Overall, our results suggest tree swallows should be added to the growing list of species challenged by climate change, and that other species of aerial insect specialists may face similar impacts of climate change.

Do hormone manipulations reduce fitness? A meta-analytic test of the Optimal Endocrine Phenotype Hypothesis.

Endocrine traits (e.g., circulating hormone concentrations, receptor expression) can vary considerably among individuals within populations. Here, we develop two evolutionary hypotheses to explain this variation. Under the Optimal Endocrine Phenotype Hypothesis, adaptive plastic responses to environmental variation generate individual variation in endocrine traits and allow individuals to express near-optimal endocrine phenotypes. In contrast, under the Ongoing Selection Hypothesis, individual variation in endocrine traits reflects varying adaptive value, with some individuals expressing suboptimal phenotypes that are selected against. These two hypotheses generate distinct predictions for the effects of hormone manipulations on fitness. Under the Optimal Endocrine Phenotype Hypothesis, all hormone manipulations should incur fitness costs, whereas under the Ongoing Selection Hypothesis, manipulating endocrine phenotypes toward a putative optimum should increase fitness. Using a meta-analysis of findings from experimental field studies that involved manipulation of circulating glucocorticoids or androgens and measurement of fitness effects, we test and find some support for the Optimal Endocrine Phenotype Hypothesis. On average, fitness was reduced across 97 estimates of the effects of experimental hormone manipulations on fitness. However, the fitness effects of glucocorticoid manipulations varied with the sex of the individuals being studied. Fitness was more uniformly reduced by glucocorticoid manipulations in males and when both sexes were considered together. In females, effects on fitness varied from highly positive to highly negative. The effects of androgen manipulations varied across males and females, and depending upon whether fitness was estimated using measures of reproductive success or survival. Reproductive success was consistently decreased by androgen manipulation in females, but was increased almost as often as it was decreased across experiments in males. When survival was estimated as a component of fitness, it was fairly uniformly compromised by exogenous androgens in males. This variation in fitness effects of hormone manipulations across sexes and fitness metrics is consistent with the expectation that hormones differentially regulate life-history investment and that optimal endocrine phenotypes differ between males and females. Overall, our meta-analysis provides some support for the Optimal Endocrine Phenotype Hypothesis, but we await direct tests of the Ongoing Selection Hypothesis to determine the degree to which individual variation in endocrine traits continues to be shaped by natural selection.

Exogenous glucocorticoids amplify the costs of infection by reducing resistance and tolerance, but effects are mitigated by co-infection.

Individual variation in parasite defences, such as resistance and tolerance, can underlie heterogeneity in fitness and could influence disease transmission dynamics. Glucocorticoid hormone concentrations often change in response to fluctuating environmental conditions and mediate changes in immune function, resource allocation and tissue repair. Thus, changes in glucocorticoid hormone concentrations might mediate individual variation in investment in resistance versus tolerance. In this study, we experimentally increased glucocorticoid concentrations in red-winged blackbirds ( Agelaius phoeniceus) that were naturally infected with haemosporidian parasites, and assessed changes in resistance and tolerance of infection. Glucocorticoid treatment increased burdens of Plasmodium, the parasite causing avian malaria, but only in the absence of co-infection with another Haemosporidian, Haemoproteus. Thus, glucocorticoids might reduce resistance to infection, but co-infection can mitigate the negative consequences of increased hormone concentrations. Glucocorticoid treatment also decreased tolerance of infection. We found no evidence that the inflammatory immune response or rate of red blood cell production underlie the effects of glucocorticoids on resistance and tolerance. Our findings suggest that exogenous glucocorticoids can increase the costs of haemosporidian infections by both increasing parasite numbers and reducing an individual's ability to cope with infection. These effects could scale up to impact populations of both host and parasite.

Rainy springs linked to poor nestling growth in a declining avian aerial insectivore ( Tachycineta bicolor).

As species shift their ranges and phenology to cope with climate change, many are left without a ready supply of their preferred food source during critical life stages. Food shortages are often assumed to be driven by reduced total food abundance, but here we propose that climate change may cause short-term food shortages for foraging specialists without affecting overall food availability. We frame this hypothesis around the special case of birds that forage on flying insects for whom effects mediated by their shared food resource have been proposed to cause avian aerial insectivores' decline worldwide. Flying insects are inactive during cold, wet or windy conditions, effectively reducing food availability to zero even if insect abundance remains otherwise unchanged. Using long-term monitoring data from a declining population of tree swallows ( Tachycineta bicolor), we show that nestlings' body mass declined substantially from 1977 to 2017. In 2017, nestlings had lower body mass if it rained during the preceding 3 days, though females increased provisioning rates, potentially in an attempt to compensate. Adult body mass, particularly that of the males, has also declined over the long-term study. Mean rainfall during the nestling period has increased by 9.3 ± 0.3 mm decade-1, potentially explaining declining nestling body mass and population declines. Therefore, we suggest that reduced food availability, distinct from food abundance, may be an important and previously overlooked consequence of climate change, which could be affecting populations of species that specialize on foraging on flying insects.

Species interactions limit the occurrence of urban-adapted birds in cities.

Urbanization represents an extreme transformation of more natural systems. Populations of most species decline or disappear with urbanization, and yet some species persist and even thrive in cities. What determines which species persist or thrive in urban habitats? Direct competitive interactions among species can influence their distributions and resource use, particularly along gradients of environmental challenge. Given the challenges of urbanization, similar interactions may be important for determining which species persist or thrive in cities; however, their role remains poorly understood. Here, we use a global dataset to test among three alternative hypotheses for how direct competitive interactions and behavioral dominance may influence the breeding occurrence of birds in cities. We find evidence to support the competitive interference hypothesis: behaviorally dominant species were more widespread in urban habitats than closely related subordinate species, but only in taxa that thrive in urban environments (hereafter, urban adapted), and only when dominant and subordinate species overlapped their geographic ranges. This result was evident across diverse phylogenetic groups but varied significantly with a country's level of economic development. Urban-adapted, dominant species were more widespread than closely related subordinate species in cities in developed, but not developing, countries; countries in economic transition showed an intermediate pattern. Our results provide evidence that competitive interactions broadly influence species responses to urbanization, and that these interactions have asymmetric effects on subordinate species that otherwise could be widespread in urban environments. Results further suggest that economic development might accentuate the consequences of competitive interactions, thereby reducing local diversity in cities.

Should I stay, or should I go: Modeling optimal flight initiation distance in nesting birds.

Flight initiation distance (FID)-the distance at which an individual leaves in response to the approach of a perceived threat-provides a measurement of risk-taking behavior. If individuals optimize their FID, this distance should reflect the point at which the fitness resulting from leaving exceeds the fitness resulting from all other possible decisions. Previous theory of FID has often been aimed at explaining this behavior in foraging individuals. Yet flight initiation in response to approaching threats occurs in a range of contexts that might influence optimal behavior. In breeding individuals, risk-taking decisions that are made at a location of offspring care (e.g., a nest or den) can have significant effects on fitness. Here, we develop a theoretical model of distances at which a parent bird flushes from a nest in response to an approaching threat. We estimate parent fitness with regards to characteristics of the parent (reproductive values, detection distance, and cost of lost parental care cost), the nest (concealment and accessibility), and the approaching predator (detection capability and predation success), developing a dichotomous scenario between staying at the nest or leaving at varying distances. Using a generalized comparison of the benefits of leaving versus staying, we find that increasing costs of lost parental care, probability of predation of the parent due to fleeing, or current reproductive value lead to more instances of staying at the nest. In a complementary approach with specified parameters based on biologically-informed factors that likely influence a predator-prey encounter, we find that increasing the current reproductive value, concealment of the nest, or costs of lost parental care decrease optimal FID and can lead to the parent staying at the nest. Other factors, such as increasing residual reproductive value, predation success, and predator capability of detecting the nest, increase optimal FID with some instances of costs of fleeing being so great that staying becomes an optimal strategy. Our theory provides a framework to explain variation in FID among nesting species and individuals and could provide a foundation for future empirical investigations of risk-taking behavior.

A New Framework for Urban Ecology: An Integration of Proximate and Ultimate Responses to Anthropogenic Change.

As urban areas continue to grow, understanding how species respond and adapt to urban habitats is becoming increasingly important. Knowledge of the mechanisms behind observed phenotypic changes of urban-dwelling animals will enable us to better evaluate the impact of urbanization on current and future generations of wildlife and predict how animals respond to novel environments. Recently, urban ecology has emerged not only as a means of understanding organismal adaptation but also as a framework for exploring mechanisms mediating evolutionary phenomena. Here, we have identified four important research topics that will advance the field of urban ecology and shed light on the proximate and ultimate causes of the phenotypic differences commonly seen among species and populations that vary in their responses to urbanization. First, we address the ecological and socio-economic factors that characterize cities, how they might interact with each other, and how they affect urban species. Second, we ask which are the proximate mechanisms underlying the emergence over time of novel traits in urban organisms, focusing on developmental effects. Third, we emphasize the importance of understanding the ultimate causations that link phenotypic shifts to function. This question highlights the need to quantify the strength and direction of selection that urban individuals are exposed to, and whether the phenotypic shifts associated with life in the city are adaptive. Lastly, we stress the need to translate how individual-level responses scale up to population dynamics. Understanding the mechanistic underpinnings of variation among populations and species in their responses to urbanization will unravel species resilience to environmental perturbation, which will facilitate predictive models for sustainability and development of green cities that maintain or even increase urban biodiversity and wildlife health and wellbeing.

Repeatability of glucocorticoid hormones in vertebrates: a meta-analysis.

We often expect that investigations of the patterns, causes, and consequences of among-individual variation in a trait of interest will reveal how selective pressures or ecological conditions influence that trait. However, many endocrine traits, such as concentrations of glucocorticoid (GC) hormones, exhibit adaptive plasticity and, therefore, do not necessarily respond to these pressures as predicted by among-individual phenotypic correlations. To improve our interpretations of among-individual variation in GC concentrations, we need more information about the repeatability of these traits within individuals. Many studies have already estimated the repeatability of baseline, stress-induced, and integrated GC measures, which provides an opportunity to use meta-analytic techniques to investigate (1) whether GC titers are generally repeatable across taxa, and (2) which biological or methodological factors may impact these estimates. From an intensive search of the literature, we collected 91 GC repeatability estimates from 47 studies. Overall, we found evidence that GC levels are repeatable, with mean repeatability estimates across studies ranging from 0.230 for baseline levels to 0.386 for stress-induced levels. We also noted several factors that predicted the magnitude of these estimates, including taxon, sampling season, and lab technique. Amphibians had significantly higher repeatability in baseline and stress-induced GCs than birds, mammals, reptiles, or bony fish. The repeatability of stress-induced GCs was higher when measured within, rather than across, life history stages. Finally, estimates of repeatability in stress-induced and integrated GC measures tended to be lower when GC concentrations were quantified using commercial kit assays rather than in-house assays. The extent to which among-individual variation in GCs may explain variation in organismal performance or fitness (and thereby inform our understanding of the ecological and evolutionary processes driving that variation) depends on whether measures of GC titers accurately reflect how individuals differ overall. Our findings suggest that while GC titers can reflect some degree of consistent differences among individuals, they frequently may not. We discuss how our findings contribute to interpretations of variation in GCs, and suggest routes for the design and analysis of future research.

Higher plasma corticosterone is associated with reduced costs of infection in red-winged blackbirds.

Glucocorticoid hormones allow individuals to rapidly adjust their physiology and behavior to meet the challenges of a variable environment. An individual's baseline concentration of glucocorticoids can reflect shifts in life history stage and resource demands while mediating a suite of physiological and behavioral changes that include immune modulation and resource allocation. Thus, glucocorticoids could facilitate a response to parasites that is optimized for an individual's specific challenges and life history stage. We investigated the relationship between endogenous circulating glucocorticoids and measures of resistance and tolerance to Haemosporidian parasites (including those that cause avian malaria) in red-winged blackbirds (Agelaius phoeniceus). We found that higher endogenous concentrations of circulating glucocorticoids were associated with reduced costs of parasite infection, which is indicative of higher tolerance, but were unrelated to parasite burden in free ranging, breeding male birds. Post-breeding, both males and females with higher glucocorticoid concentrations had higher measures of tolerance to Haemosporidian infection. Our findings suggest a potentially adaptive role for glucocorticoids in shifting the response to parasites to align with an individual's current physiological state and the challenges they face.

An experimental test of the physiological consequences of avian malaria infection.

Chronic, low-intensity parasite infections can reduce host fitness through negative impacts on reproduction and survival, even if they produce few overt symptoms. As a result, these parasites can influence the evolution of host morphology, behaviour and physiology. The physiological consequences of chronic infection can provide insight into the processes underlying parasite-driven natural selection. Here, we evaluate the physiological consequences of natural, low-intensity infection in an avian host-parasite system: adult male red-winged blackbirds (Agelaius phoeniceus) infected with haemosporidian parasites. Chronic haemosporidian infection has previously been shown to reduce both reproductive success and survival in several avian species. We used antimalarial medications to experimentally reduce haemosporidian parasitaemia (the proportion of blood cells infected with haemosporidian parasites) and measured the effect of treatment on body condition, haematology, immune function, physiological stress and oxidative state. Treatment with an antimalarial medication reduced parasitaemia for the most prevalent haemosporidian parasites from the genus Plasmodium. Treatment also increased haemoglobin and haematocrit, and decreased red blood cell production rates. We detected no effect of treatment on body condition, immune metrics, plasma corticosterone concentrations, total antioxidant capacity or reactive oxygen metabolites. Our results suggest that the damage and replacement of red blood cells during infection could be important costs of chronic haemosporidian infection. Strong links between parasitaemia and the physiological consequences of infection indicate that even for relatively low-intensity infections, measuring parasitaemia rather than only presence/absence could be important when evaluating the role of infection in influencing hosts' behaviour, physiology or fitness.

Male song sparrows have elevated testosterone in response to neighbors versus strangers.

Upon hearing a conspecific signal, animals must assess their relationship with the signaller and respond appropriately. Territorial animals usually respond more aggressively to strangers than neighbors in a phenomenon known as the "dear enemy effect". This phenomenon likely evolved because strangers represent a threat to an animal's territory tenure and parentage, whereas neighbors only represent a threat to an animal's parentage because they already possess a territory (providing territory boundaries are established and stable). Although the dear enemy effect has been widely documented using behavioral response variables, little research has been conducted on the physiological responses of animals to neighbors versus strangers. We sought to investigate whether the dear enemy effect is observed physiologically by exposing territorial male song sparrows (Melospiza melodia) to playback simulating a neighbor or a stranger, and then collecting blood samples to measure plasma testosterone levels. We predicted that song sparrows would exhibit increased testosterone levels after exposure to stranger playback compared to neighbor playback, due to the role testosterone plays in regulating aggression. Contrary to our prediction, we found that song sparrows had higher testosterone levels after exposure to neighbor playback compared to stranger playback. We discuss several explanations for our result, notably that corticosterone may regulate the dear enemy effect in male song sparrows and this may inhibit plasma testosterone. Future studies will benefit from examining corticosterone in addition to testosterone, to better understand the hormonal underpinnings of the dear enemy effect.