Song Sparrows Research Articles

Are urbanization and brood parasitism associated with differences in telomere lengths in song sparrows?

Urbanization reflects a major form of environmental change impacting wild birds globally. Whereas urban habitats may provide increased availability of water, some food items, and reduced predation levels compared to rural, they can also present novel stressors including increased light at night, ambient noise, and reduced nutrient availability. Urbanization can also alter levels of brood parasitism, with some host species experiencing elevated levels of brood parasitism in urban areas compared to rural areas. Though the demographic and behavioral consequences of urbanization and brood parasitism have received considerable attention, their consequences for cellular‐level processes are less understood. Telomeres provide an opportunity to understand the cellular consequences of different environments as they are a well‐established metric of biological state that can be associated with residual lifespan, disease risk, and behavior, and are known to be sensitive to environmental conditions. Here we examine the relationships between urbanization, brood parasitism, and blood telomere lengths in adult and nestling song sparrows Melospiza melodia. Song sparrows are a North American songbird found in both urban and rural habitats that experience high rates of brood parasitism by brown‐headed cowbirds Molothrus ater in the urban, but not the rural, sites in our study system. Among adults and nestlings from non‐parasitized nests, we found no differences in relative telomere lengths between urban and rural habitats. However, among urban nestlings, the presence of a brood parasite in the nest was associated with significantly shorter relative telomere lengths compared to when a brood parasite was absent. Our results suggest a novel, indirect, impact of urbanization on nestling songbirds through the physiological impacts of brood parasitism.

Melospiza melodia (Song Sparrow) bill size is primarily shaped by thermoregulation on the California Channel Islands

ABSTRACT Inferring the environmental selection pressures responsible for phenotypic variation is a challenge in adaptation studies as traits often have multiple functions and are shaped by complex selection regimes. We provide indirect evidence that morphology of the multifunctional avian bill is primarily shaped by climate and thermoregulatory ability in Melospiza melodia (Song Sparrow) on the California Channel Islands. Our research builds on a study in M. melodia museum specimens that demonstrated a positive correlation between bill surface area and maximum temperature, suggesting a greater demand for dry heat dissipation in hotter, xeric environments. We sampled contemporary sparrow populations across 3 climatically distinct islands to test the hypotheses that bill morphology is influenced by habitat differences with functional consequences for foraging efficiency and is related to maximum temperature and, consequently, important for thermoregulation. Measurements of >500 live individuals indicated a significant, positive relationship between maximum temperature and bill surface area when correcting for body size. In contrast, maximum bite force, seed extraction time, and vegetation on breeding territories (a proxy for food resources) were not significantly associated with bill dimensions. While we cannot exclude the influence of foraging ability and diet on bill morphology, our results are consistent with the hypothesis that variation in M. melodia need for thermoregulatory capacity across the northern Channel Islands selects for divergence in bill surface area.

Effects of sublethal methylmercury and food stress on songbird energetic performance: metabolic rates, molt and feather quality.

Organisms regularly adjust their physiology and energy balance in response to predictable seasonal environmental changes. Stressors and contaminants have the potential to disrupt these critical seasonal transitions. No studies have investigated how simultaneous exposure to the ubiquitous toxin methylmercury (MeHg) and food stress affects birds' physiological performance across seasons. We quantified several aspects of energetic performance in song sparrows, Melospiza melodia, exposed or not to unpredictable food stress and MeHg in a 2×2 experimental design, over 3 months during the breeding season, followed by 3 months post-exposure. Birds exposed to food stress had reduced basal metabolic rate and non-significant higher factorial metabolic scope during the exposure period, and had a greater increase in lean mass throughout most of the experimental period. Birds exposed to MeHg had increased molt duration, and increased mass:length ratio of some of their primary feathers. Birds exposed to the combined food stress and MeHg treatment often had responses similar to the stress-only or MeHg-only exposure groups, suggesting these treatments affected physiological performance through different mechanisms and resulted in compensatory or independent effects. Because the MeHg and stress variables were selected in candidate models with a ΔAICc lower than 2 but the 95% confidence interval of these variables overlapped zero, we found weak support for MeHg effects on all measures except basal metabolic rate, and for food stress effects on maximum metabolic rate, factorial metabolic scope and feather mass:length ratio. This suggests that MeHg and food stress effects on these measures are statistically identified but not simple and/or were too weak to be detected via linear regression. Overall, combined exposure to ecologically relevant MeHg and unpredictable food stress during the breeding season does not appear to induce extra energetic costs for songbirds in the post-exposure period. However, MeHg effects on molt duration could carry over across multiple annual cycle stages.

An Aggressive Interaction Rapidly Increases Brain Androgens in a Male Songbird during the Non-breeding Season.

Aggression is a crucial behavior that impacts access to limited resources in different environmental contexts. Androgens synthesized by the gonads promote aggression during the breeding season. However, aggression can be expressed during the non-breeding season, despite low androgen synthesis by the gonads. The brain can also synthesize steroids ("neurosteroids"), including androgens, which might promote aggression during the non-breeding season. Male song sparrows, Melospiza melodia, are territorial year-round and allow the study of seasonal changes in the steroid modulation of aggression. Here, we quantified steroids following a simulated territorial intrusion (STI) for 10 min in wild adult male song sparrows during the breeding and non-breeding seasons. Using liquid chromatography-tandem mass spectrometry (LC-MS/MS), we examined 11 steroids: pregnenolone, progesterone, corticosterone, dehydroepiandrosterone, androstenedione, testosterone, 5α-dihydrotestosterone, 17β-estradiol, 17α-estradiol, estriol, and estrone. Steroids were measured in blood and 10 microdissected brain regions that regulate social behavior. In both seasons, STI increased corticosterone in the blood and brain. In the breeding season, STI had no rapid effects on androgens or estrogens. Intriguingly, in the non-breeding season, STI increased testosterone and androstenedione in several behaviorally relevant regions, but not in the blood, where androgens remained non-detectable. Also in the non-breeding season, STI increased progesterone in the blood and specific brain regions. Overall, rapid socially modulated changes in brain steroid levels are more prominent during the non-breeding season. Brain steroid levels vary with season and social context in a region-specific manner and suggest a role for neuroandrogens in aggression during the non-breeding season.

Fear of predators reduces body and physiological condition affecting offspring survival and the ‘quality’ of the survivors

Abstract Predators affect the survival of developing young by eating them. More recently, the fear of being eaten has been uncovered as a powerful driver of offspring survival reducing recruited offspring numbers by 53%, but the mechanisms driving these effects are not well understood. We exposed song sparrows to predator or non‐predator playbacks throughout three breeding seasons. We followed offspring as eggs through development and into maturity. We quantified the repercussions of being reared in a fearful environment on body (fat, mass, size) and physiological condition using 12 different measures categorized into six systems (oxygen carrying capacity, energy reserves, immune function, antioxidant protection, oxidative stress/damage, glucocorticoids). We further tracked offspring survival in the nest to maturity and determined which measures of condition significantly predicted survival. We report that fear of predators affected offspring condition at all stages of life, with survival consequences. Predator‐playback offspring hatched from heavier eggs (8%) but quickly lost this advantage and never regained it. Nestlings reared by ‘frightened’ parents were as heavy and much leaner than non‐predator playback nestlings. This pattern extended throughout development into maturity where offspring reared by frightened parents were up to 31% leaner and 5% lighter. Predator‐playback nestlings also were evidently in poorer physiological condition, having shorter telomeres, while late dependency stage fledglings carried relatively less uric acid and more haptoglobin perhaps indicative of chronic susceptibility to infection. Only body condition (mass and fat) predicted survival during peak mortality periods (nestling and early fledging), with the leanest lightest offspring from the predator‐playback treatment being least likely to survive. Thereafter, mass predicted survival irrespective of playback treatment. Similarly, at this later stage, offspring with less telomere loss and high uric acid levels survived best regardless of playback group. Our findings reveal that fear of predators is a potent form of early life adversity which persistently alters the condition of offspring affecting early stage survival and the ‘quality’ of the survivors throughout their life. Development is fuelled by food and fear‐induced reductions in parental provisioning at the earliest stages likely instigated these effects, with consequences on the population growth rate. Read the free Plain Language Summary for this article on the Journal blog.

Multi-generation genetic contributions of immigrants reveal cryptic elevated and sex-biased effective gene flow within a natural meta-population.

Impacts of immigration on micro-evolution and population dynamics fundamentally depend on net rates and forms of resulting gene flow into recipient populations. Yet, the degrees to which observed rates and sex ratios of physical immigration translate into multi-generational genetic legacies have not been explicitly quantified in natural meta-populations, precluding inference on how movements translate into effective gene flow and eco-evolutionary outcomes. Our analyses of three decades of complete song sparrow (Melospiza melodia) pedigree data show that multi-generational genetic contributions from regular natural immigrants substantially exceeded those from contemporary natives, consistent with heterosis-enhanced introgression. However, while contributions from female immigrants exceeded those from female natives by up to three-fold, male immigrants' lineages typically went locally extinct soon after arriving. Both the overall magnitude, and the degree of female bias, of effective gene flow therefore greatly exceeded those which would be inferred from observed physical arrivals, altering multiple eco-evolutionary implications of immigration.

Effect of a Territorial Challenge on the Steroid Profile of a Juvenile Songbird

Aggression is a social behavior that is critical for survival and reproduction. In adults, circulating gonadal hormones, such as androgens, act on neural circuits to modulate aggressive interactions, especially in reproductive contexts. In many species, individuals also demonstrate aggression before reaching gonadal maturation. Adult male song sparrows, Melospiza melodia, breed seasonally but maintain territories year-round. Juvenile (hatch-year) males aggressively compete for territory ownership during their first winter when circulating testosterone is low. Here, we characterized the relationship between the steroid milieu and aggressive behavior in free-living juvenile male song sparrows in winter. We investigated the effect of a 10 min simulated territorial intrusion (STI) on behavior and steroid levels in blood, 10 microdissected brain regions, and four peripheral tissues (liver, pectoral muscle, adrenal glands, and testes). Using liquid chromatography-tandem mass spectrometry (LC-MS/MS), we quantified 12 steroids: pregnenolone, progesterone, corticosterone, 11-dehydrocorticosterone, dehydroepiandrosterone, androstenedione, testosterone, 5α-dihydrotestosterone, 17β-estradiol, 17α-estradiol, estrone, and estriol. We found that juvenile males are robustly aggressive, like adult males. An STI increases progesterone and corticosterone levels in blood and brain and increases 11-dehydrocorticosterone levels in blood only. Pregnenolone, androgens, and estrogens are generally non-detectable and are not affected by an STI. In peripheral tissues, steroid concentrations are very high in the adrenals. These data suggest that adrenal steroids, such as progesterone and corticosterone, might promote juvenile aggression and that juvenile and adult songbirds might rely on distinct neuroendocrine mechanisms to support similar aggressive behaviors.

Cultural shifts after punctuated environmental stress: a study of song distributions in Dark-eyed Junco and Song Sparrow populations

Cultural shifts after punctuated environmental stress: a study of song distributions in Dark-eyed Junco and Song Sparrow populations

Experimental manipulation of chest spotting alters territorial aggression in urban and rural song sparrows

In many taxa, melanin-based coloration is a signal of dominance or fighting ability and is associated with concentrations of hormones that may mediate aggressive behavior. Previous studies found that experimental manipulation of melanin-based signals can result in manipulated individuals receiving more social challenges in some but not all species. These differences could arise from mismatches between the signal, behavior, and hormone concentrations. In the present study, we experimentally manipulated the chest spotting of urban and rural male song sparrows (Melospiza melodia) following an assessment of their territorial aggression and initial concentrations of corticosterone and testosterone and then assessed their behavior and hormone concentrations 2 weeks later. We found that males generally displayed less territorial aggression in the second trial, consistent with our previous findings. Males in the enlarged treatment decreased aggression to a greater degree than those in the reduced treatment. The effect of the plumage manipulation was similar across the rural and urban habitats. Despite the changes in behavior we detected, we found no effects of the manipulation on concentrations of testosterone or corticosterone. Our results show that melanin-based spotting in male song sparrows is a signal of territorial aggression but the physiological mechanisms that mediate the relationships between chest spotting and behavior remain to be identified.Significance statementMany bird species use their plumage to signal their dominance status, fighting ability, or motivation during interactions with other individuals to resolve conflicts without a fight. Here, we asked whether chest spotting is a signal in territorial interactions among male song sparrows. We experimentally increased or reduced the extent of spotting in males and measured the change in their aggression. We found that reduced-spotting males showed a more moderate seasonal decrease of aggression compared to males with enlarged spotting reduced aggression, possibly because the former experienced more intrusions later on in the breeding season while the latter experienced fewer intrusions. These results are consistent with chest spotting size in song sparrows functioning as a signal of territory holding potential of the bearer.

Candidate genes under selection in song sparrows co-vary with climate and body mass in support of Bergmann’s Rule

Ecogeographic rules denote spatial patterns in phenotype and environment that may reflect local adaptation as well as a species’ capacity to adapt to change. To identify genes underlying Bergmann’s Rule, which posits that spatial correlations of body mass and temperature reflect natural selection and local adaptation in endotherms, we compare 79 genomes from nine song sparrow (Melospiza melodia) subspecies that vary ~300% in body mass (17 − 50 g). Comparing large- and smaller-bodied subspecies revealed 9 candidate genes in three genomic regions associated with body mass. Further comparisons to the five smallest subspecies endemic to California revealed eight SNPs within four of the candidate genes (GARNL3, RALGPS1, ANGPTL2, and COL15A1) associated with body mass and varying as predicted by Bergmann’s Rule. Our results support the hypothesis that co-variation in environment, body mass and genotype reflect the influence of natural selection on local adaptation and a capacity for contemporary evolution in this diverse species.

Urban and rural male song sparrows (Melospiza melodia) differ in territorial aggression and activation of vasotocin neurons in response to song challenge

When living in urban habitats, ‘urban adapter’ species often show greater aggression toward conspecifics, yet we do not understand the mechanisms underlying this behavioral shift. The neuroendocrine system regulates socio-sexual behaviors including aggression and thus could mediate behavioral responses to urbanization. Indeed, urban male song sparrows (Melospiza melodia), which are more territorially aggressive, also have greater abundance of the neuropeptide arginine vasotocin (AVT) in nodes of the brain social behavior network. Higher abundance of AVT could reflect long-term synthesis that underlies baseline territoriality or short-term changes that regulate aggression in response to social challenge. To begin to resolve the timeframe over which the AVT system contributes to habitat differences in aggression we used immediate early gene co-expression as a measure of the activation of AVT neurons. We compared Fos induction in AVT-immunoreactive neurons of the bed nucleus of the stria terminalis (BSTm) and paraventricular nucleus of the hypothalamus (PVN) between urban and rural male song sparrows in response to a short (< 5 min.) or long (> 30 min.) song playback to simulate territorial intrusion by another male. We found that urban males had a higher proportion of Fos-positive AVT neurons in both brain regions compared to rural males, regardless of the duration of song playback. Our results suggest that AVT neurons remain activated in urban males, independently of the duration of social challenge. These findings that Fos induction in AVT neurons differs between rural and urban male song sparrows further implicate this system in regulating behavioral responses to urbanization.

Indirect effects of urbanization: consequences of increased aggression in an urban male songbird for mates and offspring

Behavioral traits are often the first response to changing environmental conditions, including human induced rapid environmental change. For example, animals living in urban areas are often more aggressive than rural animals. This is especially evident in songbirds; males of several species display elevated aggression in urban habitats. Increased male aggression has been associated with reduced parental care, but the consequences of this trade-off for males, social partners, and offspring in the context of urbanization remains unclear. We explored the effects of increased urban male aggression on the life history traits, parental care, and offspring outcomes of song sparrows (Melospiza melodia). We predicted that urban males would reduce paternal investment and result in urban females providing greater nestling care or reduced fledging success in urban habitats compared to rural. Contrary to our prediction, aggressive urban males did not decrease care but visited the nest more often compared to rural males. Additionally, urban birds had higher nest and fledging success compared to rural, though this was largely due to higher nest predation in rural habitats. Our study is among the first to evaluate trade-offs associated with elevated aggression expressed by urban animals and adds to a growing body of evidence that urban habitats provide benefits to some species.

Rising to the Challenge: Mounting an Acute Phase Immune Response Has No Long-Term Negative Effects on Captive Sparrow Migratory Body Composition or Migratory Restlessness.

Migratory animals may trade-off between investing energy in immune defense versus investing in energy reserves needed for seasonal migration. However, these trade-offs are often masked by other sources of variation and may not be detected through observational field studies of free-living animals. Moreover, observational studies can rarely distinguish the costs of pathogenic infection from those of mounting an immune response. To disentangle such effects, we conducted an immune challenge experiment. We captured song sparrows (Melospiza melodia) and white-throated sparrows (Zonotrichia albicollis) in autumn migratory condition, challenged the sparrows with non-infectious antigens that induce an acute-phase immune response, then monitored body composition and migratory restlessness behavior. For both species, body mass was higher the day after exposure to keyhole limpet hemocyanin (KLH) compared to controls. White-throated sparrows, but not song sparrows, increased lean mass 1 week after exposure to lipopolysaccharide (LPS), suggesting that effects of immune upregulation on body composition may be long-lasting and specific to certain combinations of hosts and antigens. White-throated sparrows exposed to KLH increased nocturnal migratory restlessness (Zugunruhe) for the week following exposure. These findings suggest that short-term activation of the acute immune response does not constrain migratory physiology in these songbirds.

Evolutionary divergence and adaptive capacity in morphologically distinct song sparrow subspecies

Evolutionary divergence and adaptive capacity in morphologically distinct song sparrow subspecies

A perceptual field test in object experts using gaze-contingent eye tracking

A hallmark of expert object recognition is rapid and accurate subordinate-category recognition of visually homogenous objects. However, the perceptual strategies by which expert recognition is achieved is less known. The current study investigated whether visual expertise changes observers’ perceptual field (e.g., their ability to use information away from fixation for recognition) for objects in their domain of expertise, using a gaze-contingent eye-tracking paradigm. In the current study, bird experts and novices were presented with two bird images sequentially, and their task was to determine whether the two images were of the same species (e.g., two different song sparrows) or different species (e.g., song sparrow and chipping sparrow). The first study bird image was presented in full view. The second test bird image was presented fully visible (full-view), restricted to a circular window centered on gaze position (central-view), or restricted to image regions beyond a circular mask centered on gaze position (peripheral-view). While experts and novices did not differ in their eye-movement behavior, experts’ performance on the discrimination task for the fastest responses was less impaired than novices in the peripheral-view condition. Thus, the experts used peripheral information to a greater extent than novices, indicating that the experts have a wider perceptual field to support their speeded subordinate recognition.

Immune genotypes, immune responses, and survival in a wild bird population.

Individuals vary in their immune genotype, inbreeding coefficient f, immune responses, survival to adulthood, and adult longevity. However, whether immune genes predict survival or longevity, whether such relationships are mediated through immune responses, and how f affects immune genotype remain unclear. We use a wild song sparrow (Melospiza melodia) population in which survival to adulthood, adult longevity, and f were measured precisely, and in which immune responses have previously been assessed. We investigate four toll-like receptor (TLR) and the major histocompatibility complex (MHC) class IIB exon 2 genes. We test: a) whether immune genes predict fitness (survival to adulthood or adult longevity); b) whether immune genes predict immune response; c) whether immune response predicts fitness; and d) whether fitness, immune responses, or immune genotypes are correlated with f. We find that survival to adulthood is not associated with immune gene variation, but adult longevity is decreased by high MHC allele diversity (especially in birds that were relatively outbred), and by the presence of a specific MHC supertype. Immune responses were affected by specific immune genotypes. Survival to adulthood and adult longevity were not predicted by immune response, implying caution in the use of immune response as a predictor for fitness. We also found no relationship between f and immune genotype. This finding indicates that immune gene associations with longevity and immune response are not artefacts of f, and suggests that pathogen-mediated selection at functional loci can slow the loss of genetic variation arising from genetic drift and small population size.

Haemosporidian infection prevalence varies temporally and spatially and Leucocytozoon infections are male biased in Song Sparrows

Abstract Haemosporidian parasites are significant global drivers of avian disease and infections are affected by complex ecological and biological interactions. We quantified variation in haemosporidian infection prevalence of Plasmodium, Haemoproteus, and Leucocytozoon in 311 Song Sparrows (Melospiza melodia) from 2 breeding populations across years, seasons, age classes, and sexes, finding significant differences in infection prevalence across years, seasons, and sites, as well as sex differences in Leucocytozoon infection prevalence. We also found that birds were more likely to be infected with Plasmodium than Leucocytozoon and coinfections were more likely than expected. Many avian haemosporidian studies use data collected over a short period of time at one location, and few studies investigate multi-year infection dynamics across populations. However, such long-term studies are important because ecological factors can affect the distribution and prevalence of vector-borne pathogens like malaria. Because biological factors can also affect the likelihood of exposure to vector-borne pathogens, it is also important to determine how host traits influence infection dynamics. Our results demonstrate that the prevalence and composition of avian haemosporidian infections can be highly variable, making sampling regimes a key consideration when drawing conclusions from study results. To better understand the fitness consequences of haemosporidian infection in avian hosts, future work should take ecological and biological variation into account when evaluating the effects of infection on aspects of condition and quality, reproductive success, and survival.

Acute stress and restricted diet reduce bill-mediated heat dissipation in the song sparrow (Melospiza melodia): implications for optimal thermoregulation.

We used thermal imaging to show that two environmental factors - acute stress and diet - influence thermoregulatory performance of a known thermal window, the avian bill. The bill plays important roles in thermoregulation and water balance. Given that heat loss through the bill is adjustable through vasoconstriction and vasodilation, and acute stress can cause vasoconstriction in peripheral body surfaces, we hypothesized that stress may influence the bill's role as a thermal window. We further hypothesized that diet influences heat dissipation from the bill, given that body condition influences the surface temperature of another body region (the eye region). We measured the surface temperature of the bills of song sparrows (Melospiza melodia) before, during and after handling by an observer at 37°C ambient temperature. We fed five birds a restricted diet intended to maintain body mass typical of wild birds, and we fed six birds an unrestricted diet for 5months prior to experiments. Acute stress caused a decrease in the surface temperature of the bill, resulting in a 32.4% decrease in heat dissipation immediately following acute stress, before recovering over approximately 2.3 min. The initial reduction and subsequent recovery provide partial support for the hemoprotective and thermoprotective hypotheses, which predict a reduction or increase in peripheral blood flow, respectively. Birds with unrestricted diets had larger bills and dissipated more heat, indicating that diet and body condition influence bill-mediated heat dissipation and thermoregulation. These results indicate that stress-induced vascular changes and diet can influence mechanisms of heat loss and potentially inhibit optimal thermoregulation.

ECR Spotlight – Juan Zuluaga

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 during our centenary year but also the huge variety of animals and physiological systems that are essential for the ‘comparative’ approach. Juan Zuluaga is an author on ‘ Acute stress and restricted diet reduce bill-mediated heat dissipation in the song sparrow (Melospiza melodia): implications for optimal thermoregulation’, published in JEB. Juan is a PhD student in the lab of Raymond Danner at University of North Carolina Wilmington, USA, investigating the evolution of birds' adaptations to thermoregulatory challenges, and how these adaptations will fare against climate change.

Habitat associations and conservation opportunities for priority birds on small, diversified farms in the northeastern USA

Abstract Although the impacts of intensive agriculture on biodiversity and strategies for mitigating these effects have been widely described, small-scale, diversified farms and the opportunities they present for bird conservation have been less thoroughly examined. This omission is potentially significant, because this form of agriculture represents a growing sector of the industry in the populous northeastern USA, and the diverse habitats on these farms contrast with larger, structurally homogeneous intensive agriculture. To evaluate bird-habitat associations and conservation opportunities for supporting species of conservation concern on these small, diversified farms, we conducted avian point count and vegetation surveys across 23 farms in western Massachusetts during the summers of 2017 and 2018. We used Poisson-binomial mixture models and canonical correspondence analysis to assess the effects of a suite of microhabitat-, field- and landscape-scale (1 km buffer around the field) variables on the abundance of bird species. Our results confirmed that shrubland birds, a group of species of elevated conservation concern, accounted for 52% of the total observations, including song sparrow (Melospiza melodia), gray catbird (Dumatella carolinensis), common yellowthroat (Geothlypis trichas) and American goldfinch (Spinus tristis). Species–habitat relationships were diverse; however, smaller field sizes, and increased cover of tall, dense, woody or non-productive vegetation types were associated with higher abundance of shrubland species as well as lower abundance of crop pests such as European starling (Sturnus vulgaris) and house sparrow (Passer domesticus). These findings support the hypothesis that small, diversified farms are supporting birds of high conservation concern, and we provide species-specific guidelines for farmers interested in conserving birds on their land.